Riding R., 1991. Calcareous Algae and Stromatolites

Robert Riding (Ed.
Calcareous Algae
and Stromatolites
With 16 Plates and 202 Figures
Springer-Verlag
Berlin Heidelberg New York London
Paris Tokyo Hong Kong Barcelona
Dr. ROBERf RIDING
University of Wales College of Cardiff
Cardiff CFt 3YE
United Kingdom
ISBN 978-3-642-52337-3 ISBN 978-3-642-52335-9 (eBook)

DOI 10.1007/978-3-642-52335-9
Library of Congress Cataloging in Publication Data. Calcareous algae and stromatolites / R.

Riding [editor). p. cm. Includes articles originally presented at the 4th International Symposium
on Fossil Algae, held in Cardiff, July 1987. Includes index.!. Algae, Fossil - Congresses.
2. Stromatolites - Congresses, I. Riding, Robert. II. International Symposium on Fossil Algae
(4th: 1987: Cardiff, Wales) QE955.C338 1990 561'.93 - dc20 90-10112
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Preface
"Calcareous algae and stromatolites" is shorthand for a wider array

of organisms and fabrics that also includes calcified cyanobacteria,
plus thrombolites and other microbial carbonates. Composition is the
link: these are all important components of CaC0 3 sediments, from
Archaean to present and from the ocean floor to streams and lakes.
It is hardly possible to examine limestones of any age without en-
countering them. Simultaneously they are fossils, sediments, and en-
vironmental indicators. It is the range of significance, coupled with
the breadth of their distribution in time and space, which compels
their study.
Modern calcareous marine algae mainly include reds (corallines,
squamariaceans, and the nemalialean Galaxaura) and greens (dasy-
cladaleans, udoteaceans, halimedaceans). Blue-greens, of course, are
cyanobacteria and not algae, and significantly, although they are
largely responsible for Recent tidal flat stromatolites, they are not
calcified in the same way that pre-Cenozoic marine blue-greens are.
It is in the freshwater environment of calcareous streams and lakes
that we find modern calcified cyanobacteria, and they are commonly
associated with the only major group of non-marine calcareous algae,
the charophytes.
However, in the past, and especially in the Palaeozoic and
Mesozoic, things look radically different. Mingling with the ancestors
of the modern flora are distinct, and often problematic, organisms.
Their organization could be consistent with an algal or cyanobacterial
affinity, and they have been treated as such; but sometimes they hard-
ly seem to fit conceptions founded in the Recent. This is another
world, and it informs and confuses at the same time.
Most reported calcareous "algae" in the Cambrian are actually
more likely to be cyanobacteria, and are especially important in reefs.
Never again, except briefly in the Upper Devonian, do these forms
assume such importance in marine environments. Dasycladaleans
and halimedacean-like greens, strikingly similar to Recent forms, ap-
pear in the Ordovician. But any semblance of modernity is destroyed
by other newcomers which raise possibilities and problems for algal
evolution which are still unresolved. For the greens there are Ordovi-
cian receptaculitids and cyclocrinitids to be considered, together with
Late Palaeozoic "green" phylloids, and also "erect udoteaceans",
VI Preface
which so often resemble cyanobacteria. For the reds there are Soleno-
poraceae and Moniliporellaceae to be taken into account in the Mid-
dle Palaeozoic, and these are augmented by "ancestral coralline" and
squamariacean-like phylloid algae, as well as by Gymnocodiaceae, in
the Late Palaeozoic. Along with these a host of more minor proble-
matica, which for better or worse have been, entrusted to palaeolago-
logists for study, create a systematists jungle in the Carboniferous and
Permian.
End-Palaeozoic extinctions failed to eliminate this disorder. Dur-
ing the Mesozoic, despite dasycladalean diversification and the ap-
pearance of coralline red algae, the undercurrent of problematic
forms is still evident, with bizarre newcomers mingling with survivors
from the Late Palaeozoic.
No less do stromatolites present problems and show dramatic
changes with time. The striking differences between Recent forms and
those of the Precambrian are often glossed over. More than most
other fossils, stromatolites exist at two levels in the minds of those
who study them. On the one hand, there are the tangible deposits:
laminated layers, domes, and columns which are fairly easy to discern
in isolation but whose clotted, spongy, sometimes filamentous,
micro fabrics also often, since the Early Cambrian, lurk obscurely in
skeletal reefs. On the other hand, there are the shifting dogmas of ex-
planation: environment of origin (freshwater, intertidal, subtidal); mi-
crobial community (cyanobacterial, bacterial, algal); process of for-
mation (trapping and binding, calcification); reason for decline (ox-
ygen levels, invertebrate grazing, coarse bioclastic sediment); reason
for survival (saline refuges and desiccated tidal flats). Descriptive em-
phases have also varied: geometric or Linnean classifications, macro-
structures or micro fabrics.
The problem with stromatolites is not their complexity, but their
simplicity. They offer so little to grasp. But, like Plautus's Pseudolus,
we so often seem to have overlooked the certainties while seeking
uncertainties. Stromatolites are just the tip of a microbial carbonate
iceberg: lamination varies and may disappear; there are gradations
between clotted and peloidal spongiostrome fabrics, porostromate
fabrics, and macroscopically clotted thrombolite fabrics; calcification
or cement encrustation of the microbes may radically alter the ap-
pearance of the deposit. The clues have always been there in the car-
bonate fabrics. The challenge is not only to correctly interpret them
but also to objectively and accurately describe them.
The incentive for this book came about because of the 4th Interna-
tional Symposium on Fossil Algae, held at Cardiff in July 1987. But
I decided early on that there was much more need for an overview of
the subject as a whole, than for a compilation of conference papers.
Only just over half of the articles here were originally presented as
talks at the meeting, so this is not strictly a symposium volume.
Preface VII
Nevertheless, were it not for the stimulus of the Symposium this book
would not have appeared.
The title of the Fossil Algae Symposium is an overstatement. The
four meetings so far (Erlangen 1975, Paris 1979, Denver 1983, Cardiff
1987) have mainly attracted researchers interested in benthic calcified
algae, cyanobacteria, stromatolites, and other microbial carbonates.
But even within this much more limited area there is an enormous
variety of information and it is very scattered. Calcareous algae and
prokaryotes have yet to benefit from the kind of general synthetic
treatment applied to invertebrate fossils, on the one hand, and to vas-
cular plant fossils, on the other.
Precisely because of this, it is worth trying to draw together a
review volume. I cannot claim to have achieved anything close to an
even, comprehensive coverage. What has emerged, however, does
demonstrate the breadth of the subject, the variety of approach, and
the state of the science at present. Despite omissions due to con-
straints on length or non-availability of authors, this volume does in-
clude discussions of major groups of algae and cyanobacteria,
stromatolites, and other microbial carbonates, of all ages, both extant
and extinct, marine and non-marine and, where appropriate, benthic
and planktonic.
I am very grateful indeed to the following colleagues who gener-
ously helped by giving their time and expertise to referee manuscripts:
Stanley M. Awramik, Jack A. Babcock, Michael A. Borowitzka, Can-
dace M. Brooke, Yvonne M. Chamberlain, Barrie Dale, Graham F.
Elliott, Monique Feist, Erik Flugel, Richard A. Fortey, Patrick Genot,
Nicole Grambast-Fessard, John R. Groves, Llewellya Hillis-Colin-
vaux, Hans 1. Hofmann, David M. John, Andrew R. Leitch, Bernard
L. Mamet, Xi-nan Mu, Ernst Ott, Allan Pentecost, Katharina Perch-
Nielsen, Brian R. Pratt, Alain Roux, Daniel Vachard, Malcolm R.
Walter, V. Paul Wright.
It is a pleasure to thank the authors for their willingness to con-
tribute to this volume, and for their patience with my suggestions.
The skill and care of the staff of the Springer Geosciences editorial
office in Heidelberg brought the whole to fruition.
Cardiff, August 1990 ROBERI' RIDING

Contents
I Introduction...................................... 1
1 Calcification Processes in Algae and Cyanobacteria
A. Pentecost (with 4 Figures) ....................... 3
2 Classification of Microbial Carbonates
R. Riding (with 11 Figures) ......................... 21
II Major Groups .................................... 53

3 Calcified Cyanobacteria
R. Riding (with 15 Figures) ......................... 55
4 The Solenoporaceae: A General Point of View
A.-F. Poignant .................................... 88
5 Coralline Algae: Mineralization, 'Thxonomy, and
Palaeoecology
D. W. J. Bosence (with 3 Figures) .................... 98
6 Cyclocrinitids
S. C. Beadle (with 4 Figures) ........................ 114
7 Dasycladalean Algae of the Palaeozoic and Mesozoic
G. F. Elliott ....................................... 125
8 Cenozoic and Recent Dasycladales
P. Genot (with 43 Figures) .. . . ... . . . .. .. .... . . . . . . .. 131
9 Fossil Udoteaceae and Gymnocodiaceae
x. Mu (with I Plate and 3 Figures) .................. 146
10 Recent Calcified Halimedaceae
L. Hillis (with 12 Figures) .......................... 167
11 The Genus Concept in Charophyta: Evidence from
Palaeozoic to Recent
M. Feist and N. Grambast-Fessard (with 6 Figures) 189
12 Calcification of the Charophyte Oosporangium
A. R. Leitch (with I Plate and 2 Figures) ............. 204
13 Calcareous Nannofossils
S.D. Houghton (with 19 Figures) .................... 217
14 Fossil Calcareous Dinoflagellate Cysts
H. Keupp (with 17 Figures) ......................... 267
x Contents
III Algae and StromatoUtes Through Time .............. 287
15 Archaean and Proterozoic Stromatolites

S. M. Awramik (with 7 Figures) ..................... 289
16 Cambrian Calcareous Cyanobacteria and Algae
R. Riding (with 6 Figures) .......................... 305
17 Ordovician Algae and Global Thctonics
A. Roux (with 5 Figures) ........................... 335
18 Ordovician to Devonian Marine Calcareous Algae
A. Roux (with 6 Figures) ........................... 349
19 Carboniferous Calcareous Algae
B. Mamet (with III Plates and 3 Figures) ............. 370
20 Permian Marine Calcareous Algae
R. Riding and L. Guo (with 22 Figures) ............. , 452
21 Triassic and Jurassic Marine Calcareous Algae:
A Critical Review
E. Flugel (with 8 Figures) .......................... 481
22 Mesozoic and Cenozoic Marine Benthic Calcareous
Algae with Particular Regard to Mesozoic
Dasycladaleans
F. Barattolo (with VII Plates and 6 Figures) .......... 504
23 Modern Stromatolites: A Review
S. Golubic (with II Plates) .......................... 541
Index ................................................ 563

List of Contributors
Awramik, S. M. 289 Hillis, L. 167

Barattolo, F. 504 Houghton, S. D. 217
Beadle, S. C. 114 Keupp, H. 267
Bosence, D. W. J. 98 Leitch, A. R. 204
Elliott, G. F. 125 Mamet, B. 370
Feist, M. 189 Mu X. 146
Flugel, E. 481 Pentecost, A. 3
Genot, P. 131 Poignant, A.-F. 88
Golubic, S. 541 Riding, R. 21, 55, 305, 452
Grambast-Fessard, N. 189 Roux, A. 335, 349
Guo L. 452
1 Introduction
Chapter 1
Calcification Processes in Algae and Cyanobacteria
A. PENTECOST!
Abstract
In algae and cyanobacteria, calcification is always associated with biosynthetic products and, where
analyses have been made, these products have been found to consist predominantly of polysac-
charide, although no specific polymer seems to be associated with calcification.
Two aspects of the mineralization merit special consideration: photosynthesis has long been
considered the main driving force causing a depletion in aqueous carbon dioxide and an increase
in C~-. Photosynthesis will always favour calcification and the magnitude of the effect will depend
upon photosynthetic rates and the lengths of diffusion paths for the ions and molecules involved in
calcification. Second, at the ocean surface the precipitation of calcium carbonate is thermodynami-
cally favoured even in the absence of photosynthesis, which may indicate either a widespread
evolution of calcification inhibitors or absence of suit able catalysts. Today, the majority of aquatic
plants remain uncalcified.
1 Introduction
During the past 15 years the topic of algal calcification has been reviewed many
times. A detailed overview ofthe subject is not the purpose of this essay. Instead,
a brief description of calCification processes in the algal phyla will be presented
with emphasis on recent progress in the field. The earlier work has been
discussed in detail in a number of papers, a brief outline of which is given below.
Reviews of algal calcification with an emphasis on mechanisms have been
presented by Borowitzka (1977, 1982a), Krumbein (1979), Pentecost (1980),
Okazaki and Furuya (1985) and Watabe (1989). Information on carbon sources
and anatomy can be obtained from Borowitzka (1982b) and Pentecost (1985).
A few earlier reviews, notably Lewin (1962) and Arnott and Pautard (1970) are
also useful sources of information. In addition there are several specialized
reviews dealing with individual phyla or classes plus a large body of information
on calcification processes in animals. Some important principles have now been
established by combining theory with some experimentation. Our knowledge is
sufficient to state that the evolution of calcification within the algae is poly-
phyletic and even within genera the process can be remarkably diverse.
Much has been written about the degree of control exercised by organisms
over calcification. Lowenstam (1981, 1986) distinguished 'organic matrix-
mediated' biomineralization, which is biologically controlled by means of an
Division of Biosphere Sciences, King's College London, Campden Hill Road, London W8 7 AH,
I
UK
4 A. Pentecost
organically constructed template or framework, from 'biologically induced'

mineralization, where there is apparently no biological control over the process.
This allows mineralizing organisms to be classified according to the degree of
control they exercise over the process. The importance of organic frameworks
for calcification has been stressed by several researchers, stimulated in part by
the close association between mineral and organic phases in bone formation and
many types of invertebrate calcification (Lowenstam and Wiener 1983; Simkiss
1986). Research into algal calcification proceeds at a slow rate and we owe much
to research in the medical field despite the hiatus resulting from photosynthesis
which separates algal calcification from any other mineralization process. In
some ways, photosynthesis is a bete noire, as it adds the complicating factor of
a diurnal, light-dependent sink for carbon dioxide. Nevertheless, as the fol-
lowing discussion will show, photosynthesis plays some part, large or small, in
all of the calcification processes so far investigated in the cyanobacteria and
algae.
Figure 1 illustrates the approximate relative abundance of modern calcified
genera within the algal phyla divided between freshwater and the sea. Most
phyla are seen to possess only small numbers of genera with calcified species.
Considerably more cyanobacteria have been observed calcified in freshwater
compared with the sea, but in the Chlorophyta the situation is reversed. Some
species of all six charophyte genera calcify, although the degree of miner-
alization is often slight and sometimes confined to the reproductive organs.
This phylum is barely represented in salt water but is probably the major
contributor towards modern freshwater algal carbonate.
One phylum, the Prymnesiophyta, holds a pre-eminent position, with more
than 50% of the genera having calcified species (the coccolithophorids). These
minute algae are major primary producers in the oceans and must form more
biogenic carbonate than any other group of plants. A small but significant
contribution is made by another planktonic group, the Dinophyta, though
unfortunately nothing is known of the calcification processes in this great
phylum. The two remaining groups are benthic, mainly neritic algae. In the
Phaeophyta there is one genus, Padina, whose calcified species are often
abundant in warmer waters. The Rhodophyta is a large phylum containing
algae of great diversity with over 500 genera currently recognised. The com-
paratively small number of calcified genera probably gives a false impression of
the actual quantity of calcium carbonate produced, as several genera are found
in all of the world's oceans.
2 Calcification Processes in Cyanobacteria
In common with the eukaryotic algae. the cyanobacteria have left an extensive
fossil record primarily as a result of their intimate association with calcium
carbonate. Unlike many algae, these microbes have a simple organization and
lack the complex anatomy which facilitates calcification in some chlorophycean
genera such as Halimeda. Pentecost and Riding (1986) defined calcification in
Calcification Processes in Algae and Cyanobacteria 5
Key for lettering

Freshwater Marine
Freshwater
Cyanobacteria
Chlorophyta
•
Charophyta
Prymnesiophyta
C) Chrysophyta
0 Dinophyta
Fig. I. Pie charts illustrating the approx- Phaeophyta

imate relative proportion of calcified gen-
era to uncalcified genera in the major al-
o
gal ph yla. Circle size is approximately
proportional to the known abundance of Rhodophyta
the groups in freshwater and the oceans
respectively. Minor groups are omitted.
Calcified genera black. uncalcified while
cyanobacteria as "the nucleation of calcium carbonate upon or within the

mucilaginous sheath". It is necessary to make this distinction so that trapping
and binding of carbonate can be excluded as a calcification process. The
problem arises because the sheaths surrounding the cells of most cyanobacteria
promote the adhesion of particles suspended in the surrounding water. irre-
spective of their origin. Larger particles can also become trapped between the
open framework produced by filamentous genera. This is assisted by the motility
of the filaments which secrete a sticky mucilage whilst gliding phototactically to
the mat surface. Calcification is unlikely to occur with the complete exclusion of
trapping and binding. The problem is illustrated in Fig. 2 where three
calcification processes are shown in addition to binding. It is assumed, in this
model, that the surrounding water is supersaturated with respect to a CaCO:!
mineral and that nucleation begins both upon and within the sheath. In Fig. 2a,
6 A. Pentecost
s[~
W__~~~~~~~~
Fig. 2a-d. Calcification and carbonate-binding by the cyanobacterium sheath. a Calcification

resulting from biogenic nucleation within and upon the sheath. b Calcification at the sheath surface
resulting from the adhesion of seed crystals or heteronuclei originally present in the surrounding
water. Note secondary nucleation at right caused by the fragmentation of larger crystals. c
Calcification associated with bacteria upon and within the sheath. d Carbonate binding at the sheath
surface. This process is non-selective and includes non-carbonate grains. Time increases from left
to right along the abscissa. S sheath. W cell wall
heterogeneous nucleation occurs at favourable sites in a sheath composed of

diverse polysaccharides and some protein (Schrader et al. 1982). Sheath
chemistry is known to be complex, but no detailed studies have so far been made
on calcified species.]n Fig. 2b, abiogenic nucleation results from the attachment
of crystal seeds originating from the surrounding milieu. Calcification would
begin superficially but could proceed into the sheath by crystal growth or
secondary nucleation. Finally, precipitation could be initiated by bacteria which
are invariably present in natural mats. The sheath either provides a substrate or
a suitable attachment point for epiphytic bacteria. The association between
certain bacteria and cyanobacteria is well known (Paerl 1982). Calcification
could be induced by the presence of a low-energy bacterium surface reducing
the energy barrier to nucleation, or result from metabolic activities, e.g. base
formation by ammonification or nitrate-reduction (Krumbein (979). The

binding of particles (Fig. 2d) is by contrast, non-selective but difficult to
distinguish from 2b. In fact, there is no certain way of distinguishing the
processes if binding is accompanied by the growth of mineral grains in a
supersaturated solution.
Recent studies of the freshwater cyanobacterium Rivularia haematites
(D.C.) Ag. have demonstrated a pronounced seasonal pattern of growth and
calcification (Pentecost (987). Calcification results from two separate processes.
During winter, when growth was less than 2 JLm day-I, a surface precipitate of
calcite formed on the hemispherical colonies with crystallization proceeding
inwards. The process involved could be any of those described a bove. During
spring, growth was more rapid, reaching a maximum in midsummer when the
trichomes overgrew the winter layer but were initially less heavily calcified (Fig.
3). As growth continued, narrow calcified layers, termed 'summer bands'
appeared (Figs. 3, 4a). These bands could result from a pH rise induced by
photosynthesis in the region of active growth , 1-2 mm below the colony surface.
Since the entire colony is invested in a polysaccharide gel, the flow of ions is
diffusion-controlled. Counter-diffusion ofCa 2 + and CO~- ions is possible and
periodic precipitation (Keller and Rubinow (981) might occur. In any case,
calcification during this period would seem to be induced by photosynthesis.
The water at these sites was supersaturated with respect to calcite. During
......:..... - ..... .. .. ... _.
s·:·· ···_.... ·..;· .... ·
a b c d
Fig. 3a-«l. Diagram illustrating the progress of growth and calcification in Rivularia colonies.
Calcification shown by slipplingconsistsofa thick 'winter' band and a numberofnarrower'summer'
bands. Inselshows a section through a Rivularia colony. Wwinter bands; Ssurface of mucilaginous
sheath. a Spring; b early summer; c mid-summer; d winter
8 A. Pentecost
Fig. 4. a Section through calcified Rivularia colony showing a series of calcified 'summer' bands.
Crossed nicols. x250. Mastiles Stream. N. Yorks. b Calcified Scytonema (cyanobacterium) trichome
from a subaerial site. Note the dendritic crystals radiating out from the sheath surface. SEM
photograph. x 1250. Cowside Beck. N. Yorks. c Part of the articulated thallus of Cymopolia barbata
(L.) Harv. (Chlorophyta. Dasycladales) showing intense ICS calcification adjacent to the outer
cortex. Tufts of conglutinated uncalcified hairs appear at the branch apices. x5 , Jamaica. d Section
through the infolded margin of Padina japonica showing crystals of aragonite perpendicular to the
winter, the calcification is abiogenic, whilst in summer, it results from a com-

bination of abiogenic and biogenic (photosynthetic) processes. In contrast,
regular banding is not apparent in marine species of Rivularia which are
occasionally found calcified with aragonite and amorphous carbonate (Golubic
and Campbell 1981).
A second colonial, calcified cyanobacterium, Homoeothrix crustacea Wo-
ronichin, is often associated with Rivularia. Despite the close association, the
progress of calcification is quite different in this species. During winter a soft
spongy deposit is the norm, whilst in summer the deposit is more dense, and
forms 100 /-tm below a surface layer of radiating trichomes (Pentecost 1988a).
Summer bands also occur, and probably relate to periods of high photosynthetic
activity, as in Rivularia.
In Calothrix, a closely related genus, calcification follows neither of these
patterns. Growth rates of Calothrix oncoids have been found to be an order of
magnitude lower than Rivularia and Homoeothrix even where the genera
coexist (Pentecost 1989). Calothrix does not form mucilaginous colonies and
calcification is confined to the sheath which adheres closely to the trichomes.
Irregular carbonate grains permea te the shea th and extend little, if at all, beyond
the sheath boundary. There are no narrow 'summer' bands and inorganic
nucleation appears to predominate throughout the year. Photosynthesis must
playa subordinate role, considering the low rates of summer growth, amounting
to just I /-tm day-'.
Other calcified cyanobacteria include Geitleria, Lyngbya, Phormidium,
Plectonema, Schizothrix and Scytonema (Golubic 1973; Pentecost and Riding
1986). In Geitleria the discovery of regular arrays of calcite crystals within and
external to the sheath has led to the idea that the sheath may regulate this orderly
form of mineralization (Lowenstam 1986), although fine structure studies in
other genera have not revealed pronounced periodicity in the sheath. The
conditions under which such calcification occurs are uncertain. Similar ex-
tracellular calcification is found in Scytonema in cave entrances (Fig. 4b) where
condensation water may be the only significant vehicle for the reacting ions.
Dendritic calcite associated with algal (probably cyanobacterial) sheaths has
been described by Jones and Kahle (1986).
The problem of species-specificity of calcification remains, but some in-
teresting patterns are emerging. For example, clear differences in the
calcification of Calothrix, Homoeothrix and Rivularia are apparent. By placing
cyanobacteria into a gel containing counterdiffusing calcium and carbonate
ions, Pentecost and Bauld (1988) demonstrated sheath nucleation of calcite in
cell wall surface. Note the association between the fibrous wall and the crystals. Electron micrograph
Pb stained. x22000 (Okazaki et al. 1986). e Calcified colonies of Chrysonebula holmesii Lund.
Prismatic calcite crystals, growing along their c-axes develop in diffluent mucilage surrounding the
cells. Note penetration twins (?on 0112). x140. Waterfall Beck. N. Yorks. fCalcification associated
with freshwater pennate diatoms attached to a moss leaf. The carbonate crystals (identified with
EPMA) are 2-3/lm in diameter. SEM photograph x1150. Matlock. Derbyshire
10 A. Pentecost
Lyngbya and Scytonema. Under the same conditions, Entophysalis and

Pseudanabaena failed to nucleate the mineral, providing further evidence for
'species' specificity.
3 Calcification Processes in Algae
3.l Chlorophyta
Most calcified genera are marine macrophytes belonging to the Dasycladales

and Caulerpales. Whilst the former have an extensive fossil record, it is only in
the latter that there has been much experimental work (Borowitzka 1986). The
process of calcification is well understood in Halimeda and is dependent upon
thallus anatomy. The outermost cortex is composed of apressed clavate cells
enclosing a series of aqueous spaces adjacent to the photosynthetic cortex. These
intercellular spaces (ICS) are believed to contain seawater which exchanges
carbon dioxide rapidly with the adjacent cells as a result of photosynthesis and
respiration. The resulting alkalinization during CO 2 removal favours the
precipitation of aragonite. Although no direct measurements have been made
on these spaces the model is supported by several lines of evidence using
radioactive tracers and metabolic inhibitors (Borowitzka and Larkum 1976a,b;
Borowitzka 1976, 1986). This model can be readily extended to green algae
possessing a similar anatomy, e.g. Cymopolia (Fig. 4c), Dasyc!adus, Neomeris
and some species of Udotea (Borowitzka 1982b; Pentecost 1985) but the
mechanism cannot explain calcification in Acetabularia, Rhipocephalus and
Penicillus. Here, calcification is generally superficial but associated with a
'sheath-like' layer of the cell wall. In Acetabularia aragonite precipitation is
initiated within the wall and is possibly augmented by the secretion of Ca-rich
material from the cytosol (Okazaki and Furuya 1985). Wall material is also
associated with the initial calcification sites of Halimeda but the composition of
the material has yet to be determined in any of the calcified genera. Calcium-
binding polysaccharides have been reported from Halimeda (Bahm and Goreau
1973; Misonou et al. 1980a,b) but polysaccharides with similar properties are
also known in non-calcifying genera. Further studies are therefore needed in
some of these genera. particularly A cetabularia, Penicillus and Udotea.
A small number of calcified freshwater species occasionally produce
significant amounts of calcareous sediment (Muller and Oti 1981). The most
interesting of these are the planktonic species Phacotus and Coccomonas often
seen in hard-water lakes. In these unicellular forms anatomy plays no part in
calcification and it is doubtful whether these organisms alone could always
remove sufficient CO 2 from the water to cause precipitation. Phacotus pos-
sesses a thickened, calcified mucilage envelope. The breadth of the envelope
undoubtedly increases the diffusion path of dissolved gases entering the cell but
there are similar uncalcified genera with thickened envelopes e.g. Sphaerel-
lopsis. In common with some other algae. calcification of Phacollis is not always
observed in culture but a non-motile colonial phytoplankter, GloeOlaenilll11.
Calcification Processes in Algae and Cyanobacteria II
calcifies in vitro at high pH, possibly stimulated by photosynthesis (Devi-Prasad

and Chowdary 1979). Calcification also occurs in three epilithic freshwater
Chlorophytes, Cladophora (Wood 1975), Gongrosira (Golubic and Fischer
1975; Pentecost 1988b) and Oocardium (Golubic and Marcenko 1965). These
algae are often associated with calcified cyanobacteria in highly calcareous
streams, and like them, the precipitation is thought to be partly the result of
photosynthesis and partly by CO 2 evasion to the atmosphere.
3.2 Charophyta
Extracellular deposits of calcium carbonate occur on the internodal and some

reproductive cells of charophytes. Calcification has been reported in all genera
but is most apparent in Chara. Most of our knowledge of calcification comes
from studies ofthe internodal cells which have long been favourite experimental
objects for plant physiologists. Regular bands of calcite are often a conspicuous
feature of certain ecorticate species and these are known to be associated with
alkaline regions of the cell wall where the pH attains values of9 or more. This
high pH is thought to result primarily from a passive influx of H+ through the
plasmalemma (Raven et al. 1986). An active efflux of H+ occurs in adjacent
uncalcified acid bands resulting in the circulation of electric current. This efflux
is powered by ATP, the ultimate energy source being provided by photo-
synthetic reactions. This circulatory behaviour is unusual and has not been
detected in any other algae and is probably linked to photosynthetic inorganic
carbon uptake.
In the acid bands, the extra H + may combine with bicarbonate, which is the
predominant anion in most fresh waters, to form unionized carbon dioxide. The
plasma membrane is known to be permeable to CO 2 , allowing the gas to diffuse
into the cells, ultimately forming photosynthetic products within the chloroplast
stroma. This hypothetical mechanism is called 'external conversion' (Walker
1983; Price et al. 1985). An alternative hypothesis involves the combined active
uptake ofH+ and HCO; via an electrochemical pump followed by intracellular
combination perhaps catalyzed by carbonic anhydrase. Whichever hypothesis
is correct, quantitatively the effect is the same and the precipitation ofCaC0 3 in
the alkaline bands is seen to be indirectly the result of photosynthetically driven
processes. Whether such a mechanism operates in the oogonia of charophytes
(Daily 1975) remains to be seen, but this appears unlikely, and calcification may
instead result from a favourable anatomy.
3.3 Prymnesiophyta
The physiology and fine structure of coccolith formation has been studied in
depth in three coccolithophorid species; Emiliania huxleyi (Lohmann)
Kamptner, Pleurochrysis carterae Braarud and Fagerl. and Coccolith us
(Crystallolithus) pelagicus (Wallich) Schiller. The calcification process differs in
12 A. Pentecost
each case but there are similarities between Emiliania and Pleurochrysis. E.
huxleyi is the best known species. Although a common phytoplankter in the
oceans, the organic scale upon which all coccoliths form is not as apparent in this
species as it is in many others. There has been littlework to date on scale analysis
(KIa veness and Paasche 1979) and many coccolithophorids possess uncalcified
scales in addition to coccoliths. Sonication of the coccolith-bearing scales of
Emiliania (the 'c' cells, see Westbroek et al. 1986) followed by decalcification has
yielded a water-soluble polysaccharide intimately associated with the carbonate
(de Vrind de Jong et al. 1986). The polysaccharide is complex, with a molecular
weight ca. 60 000, and contains carboxyl and ester-sulphate groups.
In Emiliania c-cells, calcification is intracellular and begins in small
membrane-bound vesicles. These bodies appear to develop into more obvious
'coccolith' vesicles within which an organic scale is formed (Klaveness 1976).
Calcite is observed forming along the rim of the scale until the fully developed
coccolith, consisting of a mosaic of calcite crystals (Parker et al. 1983), is released
onto the cell surface. Staining has shown that polysaccharide is in close contact
with the vesicle membrane and the mineral surface and appears to penetrate the
crystalline areas (van der Wal et al. 1983a; Westbroek et al. 1986). Intracellular
calcification resembling that found in Emiliania has also been reported in the
genus Cruciplacolithus(FresneI1986), and is probably widespread in the group.
In Pleurochrysis carterae the vesicles are associated with the Golgi appa-
ratus and contain coccolithosomes (granular particles about 25 nm in diameter)
which, by vesicle fusion with scale-containing cisternae, appear to initiate
crystallization. Two polysaccharides have been isolated from coccolith-rich
fractions of Pleurochrysis and they show some similarity to the polysaccharide
of Emiliania (van der Wal et al. 1983b; Westbroek et al. 1986).
It has been known for some time, either through direct observation (Manton
and Leedale 1963; Green 1986; Manton 1986) or by inference (Pentecost 1980),
that scales of certain species can undergo extracellular calcification. In a study
of the 'Crystallolithus' stage of Coccolith us hyalinus Gaarder and Markali,
recalcification of decalcified scales has been observed in vitro (Green 1986;
Rowson et al. 1986). Organic matter is again closely associated with the calcite
rhombs and the entire assemblage is covered with an organic 'skin' marking the
periplast surface. An important observation here is the recognition of two
apparently disparate mechanisms operating in the same genotype, namely
intracellular calcification in the Coccolith us stage and extracellular calcification
in the Crystallolithus stage (Green 1986).
Elaborate intracellular construction seems to be required for heterococ-
coliths, whilst the simpler holococcoliths seem able to calcify at the cell surface.
Other forms of calcification are known in the Prymnesiophyta. These include
the 'vacuolar clusters' found in Pleurochrysis (Westbroek et al. 1986) and the
sheath calcification of Chrysotila (Green and Course 1983; Green 1986). In the
latter, calcification is initiated within mucilage surrounding the cells in vitro.
Minor differences in deposition have been observed between strains and may be
attributable to different polysaccharide compositions, though none has yet been
analyzed. Within the Prymnesiophyta as a whole, the link between specific

polysaccharides and calcification is strong, but mechanisms supported by a rigid
theoretical background are still lacking.
3.4 Phaeophyta
Calcification has only been reported in one genus, Padina (Dictyotales). Padina
species are widely distributed in warmer waters but only about half of them
calcify. The calcification is superficial and consists of band-like deposits of
aragonite on the thallus surface. In some species, calcification occurs on both
dorsal (outward-facing) and ventral surfaces (P. japonica Yamada), whilst in
others it occurs only on the ventral surface (P. pavonica Gaillon). In both species,
calcification has been shown to be initiated in the enrolled thallus margin, which
in section, resembles a fern crozier and contains meristematic cells (Okazaki et
al. 1986). This circinate margin behaves as a semi-enclosed space, containing a
film of sea water a few micrometres in thickness. The surrounding cells contain
chloroplasts, and calcification is probably initiated within this space as a result
of photosynthesis. Initially, crystals are found within an outer fibrous wall layer
(Fig. 4d) and remain associated with the surface after thallus expansion. Further
calcifica tion occurs on the expanded thallus, perha ps seeded by crystals already
present on the surface. Most species of Padina calcify only on their ventral
surface suggesting that the surfaces differ in composition. Of greater interest,
however, is the observation that a non-calcifying species, P. arborescens Holmes,
lacks the fibrous wall layer previously referred to, but possesses a smooth
'cuticular' layer instead. This suggests that the wall plays an important part in the
calcification either by inhibiting or directly promoting deposition. The origins
of the concentric calcified bands are still uncertain but they are sometimes
associated with zones of developing tetrasporangia showing enhanced photo-
synthetic activity.
3.5 Rhodophyta
Most intensively studied are the coralline algae, the most widely distributed
calcified benthic algae in the seas today. Most vegetative cells ofthe crustose and
non-articulated branched species are calcified to some extent. Calcification is
observed to begin immediately after segmentation of the tetraspores and is
entirely extracellular (Vesk and Borowitzka 1984). Two phases of miner-
alization are apparent (Cabioch and Giraud 1986). First, acicular crystals,
probably of calcite, form in the 'middle lamella' and outer cell walls and in some
cases these grow into tabular crystals tangential to the wall surface. More intense
calcification follows within the inner cell walls and perpendicular to the wall
surface. The growth of crystals follows the direction of polysaccharide fibrils in
the outer wall and middle lamella, but this is less apparent in the inner wall.
14 A. Pentecost
Nevertheless there does seem to be a close association between the wall material
and deposition. The coralline algae have a complex organization and compact
structure but there are no obvious differences between the cellular organization
of the corraline algae and many other Florideophyceae which might account for
the mineralization. A calcification mechanism has been proposed by Digby
(1977) which involves the photosynthetic oxidation of water followed by export
of the hydrogen ions to sea water. Intracellular hydroxyl ions react with
bicarbonate and the resulting carbonate ion is exported to the cell wall where it
reacts with calcium to form calcite. An alternative model involving localized
export of H + and OH- has also been proposed (Pentecost 1985). There is some
evidence for H+ efflux to seawater (Digby 1977) and, if this is a local effect,
parallels can be seen with the Charophyta.
Rates of calcification and photosynthesis are significantly correlated in
Corallina (Pentecost 1978; Borowitzka 1981) but correlation is not evidence for
direct participation. Lavelle (1979) has demonstrated the transport of calcium
to the calcifying tips of Calliarthron which considerably complicates the picture.
Intercalary growth occurs in the Corallinaceae but it is not known whether the
meristematic cells are always photosynthetically active. The link between
calcification and photosynthesis might be further explored through a careful
examination of these cells.
Calcification also occurs in a small number of genera belonging to other
groups. For example, in Galaxaura [astigiata Decaisne, ICS calcification occurs
in the open spaces of the young cortex (Okazaki et al. 1982) and the same
mechanism might account for calcification in a related genus, Liagora (Okazaki
and Furuya 1985).
3.6 Dinophyta, Chrysophyta, and Bacillariophyta
Calcification is rarely reported in these groups and deposition is varied, unusual,

and in the Dinophyta, may be on a sufficient scale to produce substantial
amounts of sediment. The best documented case of dinophyte calcification is
afforded by Thoracosphaera, which until recently was classified as a coc-
colithophorid (Tangen et al. 1982). The oceanic species T. heimii Kamptner has
a calcified cell wall in the vegetative coccoid stage. Calcite deposition begins, in
vitro, at numerous crystallization centres on the wall surface and eventually
covers the entire cell. The mineralization is extracellular and no evidence of
intracellular calcification has been obtained. Cyst calcification also occurs
(Keupp, this Vol.).
In the other phyla, calcification has been observed only in freshwater and
is usually associated with mucilage coatings. Calcite deposition within a
diffluent gel surrounding unicells has been observed in Tefrasporopsis
(Tschermak-Woess 1980) and Chrysonebula (Fig. 4e). In Chrysonebula nu-
cleation is confined to the gel rather than the cell walls and deposition only
occurs in waters supersaturated with respect to calcium carbonate. An unusual
case of calcification has been reported in the lorica of some Pseudokephyrion
species (Tappan 1980), a genus normally characteristic of aggressive, oligo-

trophic waters.
Diatom calcification is rarely observed but a close association between
carbonate deposition and diatom mucilage is sometimes found (Winsborough
and Golubic 1987), again, in calcite-supersaturated water (Fig. 4f). However, in
both phyla, bacteria can be observed in the mucilage and may provide nu-
cleation centres, although Green (1986) found no evidence for bacterial nu-
cleation in Chrysotila (Prymnesiophyta) which also produces copious mucilage.
4 Discussion and Conclusions
Most calcifying algal regimes are characterized by a water pH of7.8-9.5. Logic

dictates that photosynthesising cells will utilize HCO; rather than CO 2 under
these conditions because the concentration of CO 2 will be low, typically 5-10
p.M. Uptake ofHCO; may be either active or passive, depending upon the sign
of the free energy change resulting from the transport of the ion across the
diffusion gradient of an electrically charged membrane. Calculations made by
Walker (1983) and others employing estimated cytosol pH, membrane poten-
tial, and the Km of ribulose biphosphate carboxylase-oxidase indicate that
HCO; transport is almost certainly an active process. To maintain electro-
neutrality in the external medium, H+ must also be removed resulting in an
effective net loss ofH 2 C03 • In an unbounded medium this will be replaced by
CO 2 arriving through laminar and turbulent diffusion, with the length of the
diffusion path, depending upon relative water movement. Within the diffusion
zone, depletion of H+ and HCO; results in a pH rise and CO~- formation,
assuming instantaneous equilibration. Such conditions favour CaC03
precipitation and increasing the photosynthesis rate will therefore favour
calcification.
As the volume of the medium decreases relative to that of the cell, the CO 2
exchange will be more rapid, resulting in a greater pH rise. In a semi-enclosed
space, high pH may be maintained between two diffusion gradients, namely that
across the cell wall plus plasma membrane and that between the intercellular
space and the external medium, separated by a layer of wall material
(Borowitzka 1982a; 1986).
The medium surrounding calcifying algae is normally supersaturated with
respect to mineral phases of calcium carbonate. Although the degree of
supersaturation at the ocean surface is not as great as originally supposed
(Cooke and Kepkay 1984) the formation of (magnesian) calcite is still accom-
panied by a favourable free energy change of about -1 Kcal Mole-!. The
supersaturation comes from five sources whose relative significance depends
upon the local environment. These are photosynthesis, CO 2 evasion to the
atmosphere, evaporation and changes in water temperature and pressure. In
freshwater the same factors operate, apart from pressure, and high supersatu-
rations are sometimes achieved. Calcified algae have not been found in waters
undersaturated with CaC03 • This is probably due to the lack of suitable
16 A. Pentecost
protective coatings preventing dissolution of the mineral phase, which is often

extracellular, and perhaps the extra energy required for mineralization results
in no benefits for the species involved.
Calcification should occur spontaneously at the ocean surface and in many
calcium-rich freshwaters. This is sometimes observed but it is by no means
universal and the majority of algae are uncalcified (Fig. I), even where there is
additional CO 2 depletion caused by photosynthesis. Two principles are at work
here. First, the kinetics of calcite nucleation are slow, especially in seawater
(Pytkowicz 1965). Furthermore, nucleation and subsequent crystal growth
can be inhibited by a variety of substances. Inhibitors of carbonate precipi-
tation include a wide range of compounds. Particularly effective are poly
(meta)phosphates, phosphonates (Gill and Varsanik 1986), phosphate esters
and orthophosphate (House and Donaldson 1986). Other substances include
polyphenols and a range of polysaccharides. Of particular interest are polymers
isolated from calcified species. A polysaccharide associated with Emiliania was
found to be an effective inhibitor of calcite nucleation and uronic acid groups are
thought to be involved (Borman et al. 1982). The polysaccharide binds several
ions including calcium and the degree of inhibition varies with the ion bound.
Conformational changes seem able to alter the inhibitory effect which suggests
that such substances might control mineralization. A polysaccharide purified
from Rivularia also completely inhibited crystal growth at25 ppm concentration
(unpublished observations) and the sheaths of some cyanobacteria have Ca-
binding properties (Somers and Brown 1978). Further evidence for the role of
inhibitors has been obtained from fine structure studies. In A cetabularia,
calcification is confined to the inner regions of the cell wall (Okazaki and Furuya
1985) but in the coralline algae, mineralization is initially only present in the
outer wall. Deposition would normally be expected close to the cell surface
where large concentration gradients would occur through photosynthetic ac-
tivity or ion pumping.
In the absence of inhibitors, calcification may proceed, although rates could
be low. No catalysts are known to facilitate the direct combination ofCa 2 + and
CO~- IHC0:i but they may exist. Such activity might occur in the extracellular
calcification ofthe crystalloliths of Coccolith us pelagicus. Here there is a paradox
because in one stage of the life cycle, calcification is intracellular and must
involve the transport of ions across a minimum of two membranes with the
expenditure of metabolic energy, whilst in the other, mineralization seems to
proceed in the complete absence of membrane transport. Rowson et al. (1986)
noted calcification in the dark, in which case a local photosynthetic depletion of
CO 2 cannot be involved.
In all of the algae studied so far, calcification occurs associated with organic
matter, primarily polysaccharide. In some groups, particularly the Corallin-
aceae and the coccolithophorids, the association is intimate and mechanisms
involving epitaxial growth are often invoked. It is possible for an organic matrix
to stimulate nucleation and control the subsequent growth of crystals but the
mechanisms are far from clear. It is too early to state how widespread matrix
control is in the algae, and a continuum of organization levels seems to exist from
the synthesis of intracellular coccoliths down to the random deposition of calcite

in a Chrysotila colony. If organic matrices do playa major role, then we would
not expect to see them at these lower levels. Calcite is often grown in artificial gels
and this mineral appears to be unusual in maintaining growth whilst incor-
porating the gel matrix (Henisch 1970), so this intimate association in nature
may be to some extent accidental. If calcification occurred in an initially
undersaturated medium the problem would be more satisfactorily defined and
ironically, easier to tackle.
Acknowledgements. Thanks are extended to Jim Price, John C. Green, and John Dodge for useful
discussion and suggestions.
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de Vrind de Jong EW, Borman AH, Thierry R, Westbroek P, Gruter M, Kamerling JP (1986)
Calcification in the coccolithophorids Emiliania huxleyi and Pleurochrysis carterae. II. Bio-
chemical aspects. In: Leadbeater BSC, Riding R (eds) Biomineralization in lower plants and
van der Wal P, de Jong L, Westbroek P, de Bruijn WC (l983a) Ultrastructural polysaccharide
20 A. Pentecost: Calcification Processes in Algae and Cyanobacteria
localization in calcifying and naked cells of the coccolithophorid Emiliania huxleyi. Protoplasma
118:157-168
van der Wal P, de Jong L, Westbroek P, de Bruijn WC, Mulder-Stapel AA (l983b) Polysaccharide
localization, coccolith formation and Golgi dynamics in the coccolithophorid Hymenomonas
carterae. J Ultrastruct Res 85:1139-1158
Walker NA (1983) The uptake of inorganic carbon by freshwater plants. Plant Cell Environ
6:323-328
Watabe N (1989) Intracellular mineralization of invertebrates and algae. In: Crick RE (ed) 5th Int
ConfBiomineralization: evolution of ocean chemistry and its significance to biomineralization.
(in press)
Westbroek P, van der Wal P, van Emburg PR, de Vrind de Jong EW, de Bruijn WC (1986)
Calcification in the coccolithophorids Emiliania huxleyi and Pleurochrysis carterae. I. Ultra-
structural aspects. In: Leadbeater BSC, Riding R (eds) Biomineralization in lower plants and
Wins borough BM, Golubic S (1987) The role of diatoms in stromatolite growth: two examples from
modern freshwater settings. J Phycol23: 195-20 I
Wood KG (1975) Photosynthesis of Cladophora in relation to light, and CO, limitation: CaCO,
precipitation. Ecology 56:479-484
Chapter 2
Classification of Microbial Carbonates
R. RIDING!
Abstract
The main groups of organisms participating in the formation of benthic microbial carbonates are
bacteria, cyanobacteria and algae. Three main processes are involved in the formation of microbial
carbonates: (I) trapping of sedimentary particles; (2) biomineralization of organic tissue; (3)
surficial precipitation of minerals on organisms and sediments. Of these, biomineralization pre-
serves the most information concerning the organisms involved, and trapping preserves the least.
Surface mineralization is intermediate, in that it preserves information concerning the exterior size
and shape of the organisms.
The principal categories of benthic microbial carbonate deposits are:

I. stromatolite, laminated microbial deposit:
a) agglutinated stromatolite
(i) fine-grained, well-laminated
(ii) coarse-grained, crudely laminated
b) tufa stromatolite
c) skeletal stromatolite
d) subaerial stromatolite
2. dendrolite, dendritic microbial deposit
3. thrombolite, clotted microbial deposit
4. travertine, layered microbial deposit with a dendritic macrofabric
5. cryptic microbial carbonates: microbial deposits with micritic, clotted, peloidal or sparitic
micro fabrics, but lacking distinctive macrofabrics.
Only agglutinated stromatolites are dominated by trapping. Skeletal stromatolites, dendrolites, and
thrombolites are dominated by biomineralization. Tufa stromatolites, subaerial stromatolites and
travertine are dominated by surficial mineralization.
Tufa stromatolites, subaerial stromatolites, and travertine are essentially non-marine, whereas
skeletal stromatolites, dendrolites, and thrombolites are essentially marine and restricted to the
Palaeozoic and Mesozoic. Agglutinated stromatolites occur in both marine and non-marine
environments. Travertine, and especially tidal flat and subaerial stromatolites, are strongly
influenced by desiccation.
1 Introduction
The history of the development of ideas concerning stroma tolites has often been
reviewed (Maslov 1960; Hoffman 1973; Hofmann 1973; Monty 1977), and is
instructive in understanding these and other benthic microbial deposits. But it
is also true that the intricacies of the uncertainties and confusions along this path
readily obscure the results of this research during the past century. The fact is
that we can now recognize a variety of benthic calcareous deposits formed
1 Department of Geology, University of Wales College of Cardiff, Cardiff CF I 3YE, UK

22 R.Riding
primarily by microbial organisms such as bacteria, cyanobacteria, and algae,

and I propose to concentrate here on describing in a straightforward way what
I consider to be the main processes, components, and diagnostic features of the
principal types of benthic calcareous microbial deposits, together with an out-
line of the organisms most likely to be involved in their formation.
The term microbolite (adapted from microbialite of Burne and Moore
1987) is useful because it encourages us to view benthic microbial carbonate
deposits as a whole, rather than concentrating on one type, such as stromatolites,
alone. However, this broad view requires both comparative terminology and a
system of classification which can accurately, but without unnecessary com-
plication, provide basic information linking the organisms, processes and
products involved. In particular, terms are required for non-calcified, non-
mineralized "classic" stromatolites formed by trapping of particulate sediment
which I here propose to call agglutinated stromatolites; and a term is required
for calcified but non-laminated microbial deposits with a dendritic fabric which
I propose to call dendrolites (Riding 1988). Travertine and some types of tufa
need to be included in microbial carbonates, although much still remains to be
learned about these non-marine deposits.
Microbial carbonates are defined here as the carbonate deposits produced
or localized by benthic (to exclude coccolith and foraminiferal oozes) microbial
communities. The term microbe is used here for any microscopic organism. The
principal benthic microbes involved in these deposits are bacteria, cyanobac-
teria (both of which are prokaryotic) and algae such as diatoms and various
chlorophytes and rhodophytes. The principal invertebrate microbes locally
involved in construction of benthic microbial carbonates are foraminifers.
Cyanobacteria (cyanophytes, blue-greens) are especially important, and
reviews of the groups prominent in carbonate deposition have been made by
Golubic (1973) and Pentecost and Riding (1986).
The communities of organisms creating these deposits are often termed
"microbial mats" in the current literature (Gerdes and Krumbein 1987, p. 5),
and "algal mats" in earlier literature, a reference to the densely interlayered,
intertwined and variously prostrate or erect orientations of the coccoid and
filamentous cells involved. The composition of these communities will have
changed through time (Fig. I).
2 Processes
The main processes of microbial carbonate formation are the same as for
stromatolite formation: (1) trapping (agglutination) of sedimentary particles;
(2) biomineralization of organic tissues; (3) mineralization, i.e. surficial
precipitation of minerals on organisms and/or sediment (Fig. 2). The most
important distinction to be made here is between biomineralization (2 above)
and mineralization (3 above). Biomineralization here signifies precipitation
within organisms or organic tissue during life and implies a significant degree of
biological control over precipitation. Mineralization, in contrast, is here
regarded as precipitation outside organisms.
Classification of Microbia l Carbonates 23
modern hot spring

stromatolites, travertine,
salt stromatolites &
? Archaean stromatolites
? Proterozoic
stromatolites
Palaeozoic / Mesozoic
dendrolltes, thrombolltes
& skeletal stromatolites
coarse modern
stromatolites
grained tidal flat
stromatolites stromatolites
Fig.1. ABC (algae, bacteria, cyanobacteria) diagram showing possible variations in time and space
of the principal constructional organisms in benthic microbial carbonate deposits. A rrow indicates
likely time trend for marine forms . Subaerial stromatolites (not shown) contain fungi and lichens in
addition to prokaryotes and algae
S T S
a b c
Fig. 2a-c. Processes of benthic microbial carbonate deposition exemplified by a filamentous
cyanobacterium; T trichome ; S sheath . aTrapping of sediment particles on the sheath exterior; b
calcification (biomineralization) during life of the sheath interior; c precipitation ofa surficial crust
ofCaCO, upon the sheath exterior (mineralization). None of these processes is mutually exclusive,
although trapp ing will be by baffling rather than by adhesion ifeithercalcification or mineralization
of the sheath has occurred
24 R. Riding
2.1 Trapping
Stickiness and relief (even if the relief is only microscopic) on a surface can both
result in retention of sedimentary particles. These two sub-processes of trapping
are theoretically separate: adhesion due to the sticky or soft nature of mucilage
produced by microbes, and baffling due to organisms impeding movement of
particles by physical obstruction. In practice, adhesion and baffling may not be
readily distinguishable. For example, erect cyanobacterial filaments will act as
baffles and may also be encased in a sticky mucilaginous sheath. On a larger
scale, pustular colonies of cyanobacteria and/ or diatoms can create microrelief
and may also be sufficiently soft for particles to become imbedded within the
sticky gel which they create.
Sedimentary particles will be transported by currents onto the microbolite
surface. They may then be removed again by currents which are sufficiently
strong, or they may be trapped. The longer the grain remains on the microbolite
surface, the greater is the chance that it will be incorporated into the deposit. This
is due to the vertical growth of the organisms, a process generally referred to as
binding. "Trapping and binding" is a phrase widely used in the stromatolite
literature to indicate the initial retention of particles on a microbial substrate
and their subsequent incorporation.
The sizes of particles trapped will depend mainly on: (1) size and abun-
dance of particles introduced by currents; (2) strength and frequency of cur-
rents; (3) stickiness of the surface; and (4) relief on the surface. Smaller, micrite-
grade, particles will be preferentially tra pped. The stickiness and relief of most
microbial surfaces are generally insufficient to trap sand-size particles (Fig. 3).
An exception to this is the more complex, more sticky, and more irregular
surfaces of algal-cyanobacterial communities (Awramik and Riding 1988).
This relationship between particle size and trapping potential of the
microbolite community is probably important in permitting long-term accre-
tion. Iflarge particles were deposited relatively rapidly on a surface occupied by
small, relatively slow-growing organisms, then the survival of the community
would be threatened. Pratt (1982) is probably correct to suggest that the
widespread production of coarse bioclastic sediment due to metazoan
biomineralization has been a factor of equal significance to the grazing effects
of herbivorous animals in the Phanerozoic decline of stromatolites formed by
tra pping of sediment.
Trapping appears to operate in all types of microbial carbonate deposition,
but it is the pre-eminent process only where calcification, whether by biomin-
eralization or by inorganic mineralization, is relatively unimportant. It is
therefore most conspicuous in environments, or at times, where calcification is
reduced. In non-marine environments this is more likely to be in lakes than in
rivers. In shallow marine environments it has mainly been during the
Precambrian and Cenozoic (see Distribution, below).
Classification of Microbial Carbonates 25
microbial mat
a
o
c:;:> 0 "' O~ ' ~ ... .. .
~ ~ 'o
0 '.: : ••
---
• ...
~
---
o 0 .
~~J<l;~~~~
0 . '.
preferential trapping
C of smaller particles
Fig.3a-c. Selective tra pping of smaller particles by a benthic microbial mat community. Larger
particles are not easily retained by small organisms, whereas small particles are
2.2 Biomineralization
The degree of control over mineralogy and crystal growth in biomineralization

varies with the organism. Attempts to clarify these processes, and the termi-
nology used to describe them, are still being debated. Lowenstam (1981) and
Lowenstam and Weiner (1983) recognized biologically-induced mineralization
(resulting in random crystal aggregates) and organic matrix-mediated miner-
alization (where minerals adopt unique crystal habits with well-defined
orientation) as end-members of an intergrading spectrum . Simkiss (1986)
emphasized four factors controlling biomineralization: I) modification of
ion-activity, 2) presence of a diffusion limited site, 3) presence of a nucleating
surface, 4) lattice modifiers and crystal inhibitors.
In general, microbial calcification appears to be induced rather than
mediated, although details of the operation of Simkiss' factors still need to be
elucidated in most of the groups involved. Bacterial calcification (Krumbein
1986) is less well understood than cyanobacterial calcification but principal
products are spherulites (Danielli and Edington 1983) and bushy growths
(Chafetz and Folk 1984). Cyanobacterial calcification (Pentecost and Riding
26 R. Riding
1986) produces a wide range of bushy, dendritic and tubular fabrics. Macro-
scale deposits of these , such as skeletal stromatolites, dendrolites, etc. are
discussed here, but discrete microfossils, such as Girvanella, Cayeuxia, and so
on, are not included (for details see Riding, calcified cyanobacteria, this
Vol.). Calcification of eukaryotic algae is less important in microbial carbonate
formation , not because in place deposits produced by benthic algae are un-
common, but because the large size of the algae generally involved precludes
them from being regarded as microbes.
Calcification during life of the organisms enhances the accretion rate ofthe
microbial deposit and, by preserving organisms ingrowth position, it commonly
increases micro-relief of the surface and thereby enhances trapping of particles.
At the same time, it may disrupt lamination ifthe calcified organisms have erect
orientation. Consequently, calcification often results in a coarse, unlaminated or
at best crudely laminated fabric in which the original organisms are preserved
to some degree; usually their size distribution and general morphology are
retained . Closely spaced erect calcified tufts and bushes may create intervening
shelter cavities. Where these remain unfilled by particulate sediment, coarse
fenestrae then develop which differ in size and orientation from those associated
with desiccation in, for example, tidal fiat stromatolites. Another product of
calcification is post-mortem nucleation of cements upon the microbes. These are
often conspicuous, for example, in Early Palaeozoic dendrolites formed by
A ngulocellularia Vologdin , Renalcis Vologdin and Epiphyton Bornemann (Fig.
4; see Dendrolites below). Thus, microbial carbonates formed by biominer-
alization basically consist of:
2. post-mortem
1. biominerallsed
early cement fringe
organisms
3.shelter cavity &

intervening micrite matrix
Fig. 4. Composition of calcified microbial deposits exemplified by dendrolite, consisting of 1

biomineralized organism ; 2cement crusts or void fills which , in this example, often predate or are
contemporaneous with the trapping of sediment; 3 trapped particulate sediment. The erect,
branched or bushy fossils in Lower Palaeozoic dendrolites typically belong to the Epiphyton ,
Renalcis . and/or A ngulocelllliaria groups
1. Calcified organisms;
2. Trapped particulate sediment;
3. Cement crusts or void fills.
In terms of the fabrics and degree of lamination they possess, these microbial
carbonates may be subdivided into skeletal stromatolites (crudely laminated,
with prostrate or erect calcified organisms), dendrolites (unlaminated, with
erect, or pendent dendritic calcified organisms), and thrombolites
(unlaminated, with a macroscopic clotted fabric which may have a variety of
origins, including the presence of rounded, or irregular, calcified organisms).
Composition and fabrics of microbial carbonates are discussed further below
(see Composition and fabrics).
2.3 External Surficial Precipitation of Minerals
This process, here termed mineralization, is, in effect, early cementation. In

microbial carbonates it is important in the formation of subaerial and tufa
stromatolites and travertine. It results in crystalline veneers on the surfaces of
organisms, irrespective of whether or not they are biomineralized, and on
associated particulate sediment. Mineralization in marine microbial environ-
ments is most conspicuous in reefal and intertidal situations. In non-marine
environments it is most prominent in areas of rapid CO 2 evasion, such as springs
and waterfalls, or high evaporation.
3 Types of Benthic Microbial Carbonates
3.l Microbolites
The term microbia lite was proposed by Burne and Moore (1987), and defined
as "organosedimentary deposits that have accreted as a result of a benthic
microbial community trapping and binding detrital sediment and/or forming
the locus of mineral precipitation". Here I replace the adjectival prefix
"microbial" by the noun-based prefix "microbo-" to conform with similar
established usage, as in stromatolite and thrombolite. Burne and Moore's (1987)
definition is, in fact, very similar to Awramik and Margulis' definition of
stromatolite (see Stromatolites, below).
The processes of microbial carbonate formation outlined above can be
related directly to the principal calcareous microbolite types: stromatolites,
dendrolites, thrombolites, and travertine (Table I). Sediment trapping results in
agglutinated stromatolites, characterized by a particulate microstructure which
preserves few, if any, details of original size, shape or orientation of the
organisms responsible for localizing deposition. Biomineralization, in contrast,
retains relatively considerable detail of the organisms, which are preserved as
skeletal microfossils. These can often be recognized as cyanobacteria or,
28 R. Riding
'Illble 1. Possible main relationships between the processes and products of calcareous microbial
deposits"
Process Product
Subaerial stromatolite
Mineralization ---------------------------------------
Tufa
stromatolite Travertine
-------------------------- ---------------
Biomineralization Skeletal Dendrolite Thrombolite
stromatolite
- - - - - - - -------------------------- - - - - - - - - - - - - - - -
Trapping Agglutinated stromatolite
Microbial growth Prostrate Erect Coccoid

Microbial mat
"Note that (I) all thrombolites are not necessarily produced by biomineralization of coccoid
microorganisms; (2) biomineralization influences subaerial stromatolites, but they are dominated
by mineralization processes; (3) prostrate, erect and coccoid growth styles strongly influence the
fabrics ofbiomineralized microbial deposits (skeletal stromatolites, dendrolites, thrombolites), but
are less well reflected in agglutinated and subaerial stromatolites; (4) lamination (the characteristic
feature uniting stromatolites) is due to episodic sedimentation, episodic growth, or prostrate habit.
or a combination of these.
especially in the Early Palaeozoic, as probable cyanobacteria. Consequently,

skeletal stromatolites, dendrolites and thrombolites generally reveal much more
information concerning the microbes responsible for their formation than do
agglutinated stromatolites (non-skeletal stromatolites of Riding 1977).
Mineralization is especially important in non-marine microbolites. Tufa
stromatolites form in freshwater environments where inorganic CaC0 3
precipitation is enhanced by CO 2 evasion. This results in surficial veneering of
microbes, mainly cyanobacteria, and sediment by CaC03 which would be
termed cement if it were interstitial. There is no general term for this surface
crystalline veneer; I term it here "surface cement". Processes of travertine
formation are as yet poorly understood, but it is likely that mineralization is also
a major factor for these non-marine, warm spring, deposits and for subaerial
stromatolites. Exposure results in desiccation, which promotes mineralization
and early lithification. There is thus an important relationship between the
environment and the processes of formation of microbial carbonates. Den-
drolites, skeletal stromatolites, subtidal agglutinated stromatolites and,
probably, thrombolites are subaqueous in origin. Tufa stromatolites are es-
sentially subaqueous, but may be occasionally exposed by falls in lake or river
levels. On the other hand, tidal flat agglutinated stromatolites, travertine and
subaerial stromatolites represent a trend towards increasing exposure which will
be reflected in their processes of formation and the fabrics they contain.
The important distinction between stromatolites on the one hand and

dendrolites and thrombolites on the other is based upon the presence or absence
of lamination. Any of the three principal processes (trapping, biomineraliza-
tion, mineralization) may produce stromatolites, provided that lamination (due
to episodic sedimentation and / or growth, or to prostrate habit ofthe organisms)
is also developed, even though the quality of the lamination will vary.
The principal categories of benthic microbial carbonate deposits recog-
nized here are as follows:
I. Stromatolite: macroscopically laminated benthic microbial deposit.

a) Agglutinated stromatolite, produced by trapping/binding of par-
ticulate sediment:
(i) fine-grained, well-laminated;
(ii) coarse-grained, crudely laminated.
b) Tufa stromatolite, produced by precipitation of minerals on (rather than
within) organic tissue.
c) Skeletal stromatolite, produced by calcified in place organisms.
d) Subaerial stromatolite, dominated by mineralization in a desiccating
environment.
2. Dendrolite: macroscopically dendritic benthic microbial deposit.
3. Thrombolite: macroscopically clotted benthic microbial deposit.
4. Travertine: layered benthic microbial deposit with a dendritic macrofabric.
5. Cryptic microbial carbonate: benthic microbial carbonates which have
micritic, clotted, peloidal or sparitic microfabrics, but which lack distinctive
macrofabrics.
3.2 Stromatolites
Kalkowsky (1908) introduced the term stromatolith (for details see Monty
1977), but research on stromatolites preceded his paper by at least twenty-five
years. Hall's (1883) description of Cryptozoon from the Cambrian of New York
marks the beginning of close attention to stromatolites. Definition of the term
has oscillated between genetic and descriptive approaches. Genetically,
stromatolites may be defined as: "organosedimentary structures produced by
sediment trapping, binding and/or precipitation as a result of the growth and
meta bolic activi ty of microorganisms, princi pally cyanoph ytes". This defini tion
was presented by S.M. Awramik and L. Margulis in the Stromatolite Newsletter
in 1974 (see Walter 1976a, p. I). It is noteworthy that it makes no reference to
lamination and is thus applicable to benthic microbial deposits generally. In
fact, Burne and Moore (1987) use virtually the same definition for microbolites.
Descriptively, stromatolites may be defined as: "attached, laminated, lithified,
sedimentary growth structures, accretionary away from a point or limited
surface of initiation" (Semikhatov et al. 1979, p. 993), an all-embracing view
which would readily include many ooids and other coated grains, as well as
30 R.Riding
sinter and some calcrete fabrics. A more simple, but combined, definition
proposed here is "stromatolites are laminated benthic microbial deposits".
In Recent marine stromatolites trapping of sediment is by far the most
important process involved. In non-marine stromatolites at the present day, in
contrast, biomineralization and mineralization are usually dominant. Walcott
(1914) compared Proterozoic stromatolites with modern tufa, and this fresh-
water analogue was popular until Black's (1933) recognition of sediment
trapping in intertidal and supratidal stromatolites led to the widely held
"trapping and binding" view of stromatolite which persists to the present. It is
not difficult, in hindsight, to see that conflicting views of stromatolite formation
are largely due to failure to discriminate between space and time variations in
stromatolite development. As has already been stated, stromatolites may be
formed by one of several processes: trapping, biomineralization, or miner-
alization. Furthermore, these processes have in the past operated in environ-
ments in which they are now rare. For example, cyanobacterial biominer-
alization has clearly taken place in marine environments in the Palaeozoic and
Mesozoic, but has been rare during the Cenozoic (Riding 1982).
3.2.1 Agglutinated Stromatolites
This term is introduced here for stromatolites produced by trapping of par-

ticulate sediment. Two types can be distinguished: (1) fine-grained, well-
laminated and (2) coarse-grained, crudely laminated (Awramik and Riding
1988).
Fine-Grained, Well-Laminated. This is the predominant Precambrian stroma-

tolite (Fig. 5), but it persists in marine and non-marine environments until
the present-day. Fine grain size is often due to a combination of supply and
selection: the microbes involved (probably bacteria and/or cyanobacteria) will
have difficulty in coping with coarse sediment and fine-grained stromatolites
can be observed to develop even in environments where coarse sediment is
available. Coarse, sand-sized material will not be retained by these small
organisms, just as a football is unlikely to adhere to glue-covered strands if they
are only a few millimetres in diameter. And if a cyanobacterial layer should be
covered by coarse sediment, it is unlikely to successfully colonize such large
grains, whereas micritic particles may be readily trapped and bound as the
organic filaments move up and over them.
Episodic sedimentation, linked to periodic microbial growth, will result in
good lamination, and this is a characteristic feature of many fine-grained
stromatolites. Lamination, even when well-developed, is however not neces-
sarily smooth, and we can distinguish:
1. Smooth lamination, of subaqueous origin:

2. Crinkly lamination, with fenestrae, due to exposure and dessication on tidal
flats or lake margins.
Fig. 5. Preca mbrian columnar stromatolites, ca . 1100 million years; Tieling Formation, near
Tielingzi, Jixian County, Tianjian Province, northern China. Note the good lamination. Scale bar
is 5 cm long
Coarse-Grained, Crudely Laminated. Awramik and Riding (1988) recognized a

probable link between an algal (as opposed to only cyanobacterial) floral
component and coarse grain size in Shark Bay subtidal columnar stromatolites.
They contrasted coarse-grained, crudely laminated, algal-cyanobacterial
stromatolites with fine , well-laminated cyanobacterial stromatolites and cast
doubt on the previously widely established view that Shark Bay columnar
stromatolites are analogues for most Precambrian stromatolites. Subsequently,
this link between coarse grain size and the presence of eukaryotic algae has been
supported by study of "giant" Bahamian stromatolites at Lee Stocking Island,
which have sand-size grains and a flora showing an approximately equal mix of
algae and cyanobacteria (Riding et aI., in preparation).
No ancient examples of similar coarse-grained stromatolites have yet been
reported, and it remains to be seen whether these may date from the evolution
of algae in general (probably 1000 Ma) or from the appearance of diatoms
during the Cretaceous.
32 R. Riding
3.2.2 Tufa Stromatolites
This term is introduced here for stromatolites dominated by precipitation of

minerals on (as opposed to within) organic substrates. These deposits are
characterized by crystalline precipitated veneers of calcium carbonate upon the
external surfaces of the organisms. These crusts interfere with organic growth
and result in rapid sedimentary accretion. Cyanobacteria are the principal
organisms involved in tufa stromatolite formation. Other types of tufa, not
considered here because they are not laminated, include algal and moss tufa.
Tufa stromatolites are not known from marine environments. They are
common in modern calcareous freshwa ter lakes and streams (e.g. Eggleston and
Dean 1976; Winsborough 1987; Wins borough et aI., in prep.). In particular they
are prominent here as oncoids (e.g. Clarke 1900; Roddy 1915) which in
freshwater may combine features of tufa stromatolites and skeletal stroma-
tolites. These are produced by both biomineralization and inorganic miner-
alization. The former process is likely to be dominant in lakes, the latter in
streams. These modern freshwater tufa oncoids have been compared with
ancient marine oncoids, but there are often pronounced differences: ancient
marine oncoids are usually either agglutinated oncoids or skeletal oncoids, and
the latter normally lack the component of surficial inorganic mineralization
which is characteristic of tufa stromatolites. Nevertheless, there is intergrada-
tion between tufa stromatolites and skeletal stromatolites.
3.2.3 Skeletal stromatolites
Riding (1977) introduced this term to describe "stromatolites in which the

organisms responsible for their formation are commonly preserved as calcified
fossils" (Riding 1977, p. 57). It is important to distinguish now between phys-
icochemical calcification (here termed mineralization) and biochemical
calcification (here termed biomineralization). Skeletal stromatolites refer to
stromatolites formed by biochemical calcification, as was shown on the
original triangular diagram (Fig. 6), and this distinguishes them from tufa
stromatolites which are dominated by mineralization (surficial inorganic
precipitation).
Skeletal stromatolites occur in marine environments in Palaeozoic and
Mesozoic rocks. They are typically constructed by calcified cyanobacteria such
as Girvanella Nicholson and Etheridge and Ortonella Garwood. These fossils
can be classified as Porostromata (Pia 1927, p. 37) and skeletal stromatolites
have also been termed porostromate stromatolites (Monty 1981). However, the
Porostromata is a grouping of heterogeneous fossils and the term skeletal is
preferred here. Some Recent lacustrine stromatolites resemble skeletal
stromatolites. However, freshwater microbial carbonates generally possess an
element of mineralization which places them in the category of tufa stroma-
tolites. Nevertheless, this physicochemical precipitation effect is generally
weaker in lacustrine than in fluviatile freshwater environments because CO 2
sediment trapping
physicochemical biochemical
carbonate precipitation
Fig. 6. Relationships between processes of carbonate deposition in the formation of nonskeletal

stromatolites (NSS), tufa (n. and skeletal stromatolites (SS) (Riding 1977, Fig. I)
evasion is enhanced by water movement. As a result, it is possible for lacustrine

stromatolites to approach skeletal stromatolites in appearance.
The term cyanoid, describing oncoids constructed by calcified cyano-
bacteria (Riding 1979, 1983), is applicable to both tufa oncoids and skeletal
oncoids.
The time distribution of skeletal stromatolites in marine environments
parallels that of other biomineralized microbial carbonate deposits such as
dendrolites (and also thrombolites containing calcified microfossils). These
appear to be restricted to the Palaeozoic and Mesozoic, probably in response to
an environmental control such as sea-water chemistry (Riding 1982).
3.2.4 Subaerial Stromatolites
Research on highly calcareous soils and subaerially exposed limestone surfaces

has resulted in progressive refinement of understanding of calcrete and caliche
deposits. The importance of micro-organisms, particularly prokaryotes, fungi ,
lichens and green algae, in subaerial laminated crusts has been increasingly
recognized (Krumbein 1968; Kahle 1977; Klappa 1979a) with the result thatthe
distinction between calcretes and stromatolites (Read 1976) has become
blurred. As a result, terms such as lichen stromatolite (Kia ppa 1979b) and desert
stromatolite (Krumbein and Giele 1979) have been applied, and it is now
necessary to recognize laminated microbial calcretes as varieties ofstromatolite
(Wright 1981).
34 R. Riding
Subaerial stromatolites form in environments dominated by desiccation

and vadose diagenesis. They are highly variable, but features similar to those of
subaqueous and tidal flat stromatolites are conspicuous. These include well-
developed lamination, fenestrae and also coated grains resembling oncoids (but
usually referred to as vadose pisoids, e.g. Dunham 1969; Esteban and Pray
1983). In addition, pedogenic fabrics, such as glaebules, alveolar texture,
circumgranular cracks and micrite bridges are usually present (see Esteban and
Klappa 1983). Intense desiccation, as well as biogenic processes, result in early
lithification of subaerial stromatolites, and carbonate precipitation and re-
placement dominate. Trapping of particles appears to be of relatively minor
importance. There are similarities between subaerial stromatolites and
travertines: both form in environments strongly influenced by desiccation and
carbonate precipitation. Thus, subaerial stromatolites combine features of
subaqueous stromatolites with distinctive pedogenic fabrics. Their recognition
is further assisted by their intimate association with weathering profiles.
3.3 Dendrolites
This term was introduced by Riding (1988) for non-laminated biomineralized

microbial deposits with a dominant dendritic macrofabric. Dendrolites formed
by genera such as Epiphyton, Renalcis and Angulocellularia are conspicuous in
Cambrian and Lower Ordovician reefs (Fig. 7), commonly in association with
Fig. 7. Dendrolite. Dendritic fabric created

by ERA (Epiphyton, Renalcis and/or An·
gulocel!ularia) microfossils. apparently pen-
dant in reef cavity. Serie Schisteuse. Lower
Cambrian. Assads, Anti-Atlas Mountains,
Morocco. Specimen provided by Andrew
Latham. Width of view 38 mm
sponge-like organisms such as archaeocyaths. They reappear in Upper

Devonian reefs where the dendritic fossils are again Epiphyton and Renalcis
Vologdin, or close relatives of these genera, and the principal metazoans with
which they are associated are stromatoporoids. The "bacterial shrubs" which
occur in some travertines (see Chafetz and Folk 1984, Fig. 1) have dendrolitic
fabrics, but are arranged in layers. There is also a link with skeletal stromatolites,
which differ from dendrolites principally in the presence oflamination, and with
thrombolites which may show intergradation with dendrolites.
Epiphyton-Renalcis-Angulocellularia (ERA) microfossils are common
constituents of Cambrian reefs, often completely dominating the rock fabric.
They are frequently associated with early cementation localized between their
filamentous and bushy masses (Fig. 4). Although generally individually
microscopic, these fossils amalgamate to form dendritic fabrics, either vertically
erect or pendant, which are readily visible in hand-specimen. The result of this
growth, calcification, and associated early cementation, was the formation of
microframe reef structures which provided hard substrates for attached me-
tazoans, notably archaeocyaths. This substrate-attachment relationship could
be reversed, with ERA microfossils clustered upon archaeocyaths. Thus,
Cambrian reefs typically show two basic skeletal end-members: ERA den-
drolite and archaeocyaths.
Although the precise affinities of most ERA microfossils are uncertain; their
probable cyanobacterial affinities (see Riding, Calcareous cyanobacteria and
algae, this Vol.) would ensure their placement in microbolites. Their lack of
lamination, combined with biomineralization, ally them with thrombolites,
although more remains to be understood concerning the origins of the latter.
The diversification of metazoan reef-builders in the Middle Ordovician
modified reef composition considerably, and substantially reduced the im-
portance of ERA micro-organisms in reef ecosystems. Consequently, ERA
dendrolites are most conspicuous at times when the importance of metazoans
was reduced: (1) Cambrian to Lower Ordovician, prior to the advent of the
stromatoporoid-tabulate-bryozoan association; (2) Upper Devonian, during
the decline of stromatoporoids; (3) Lower Carboniferous, following the ex-
tinction of most Palaeozoic stromatoporoids.
Further documentation of ERA and other dendrolites will clarify their
relationships to both thrombolite, skeletal stromatolite, and travertine.
3.4 Thrombolites
Aitken (1967) introduced the term thrombolite for non-laminated cryptalgal

(i.e. microbial) carbonates with a clotted fabric. In the field thrombolites
commonly possess domical morphologies resembling stromatolites, but they
lack lamination. The clotted fabrics ofthrombolites are quite variable and may
alternate with stromatolitic lamination within a single dome. Aitken (1967)
noted that thrombolites are common in the Cambrian and Ordovician and
subsequent studies have confirmed this: they have been reported from the
36 R. Riding
Cambrian, Ordovician, Silurian, Carboniferous and Permian, but are especially

abundant in the Upper Cambrian and Lower Ordovician (Kennard and James
1986, Fig. 8). Despite recognition for more than twenty years, the origins of
thrombolites remain uncertain. Walter and Heys (1985, p. 164) considered that
thrombolites are disturbed stromatolites in which the activity of animals and
algae has disrupted and destroyed the lamination. However, Kennard and
James (1986, p. 494) attributed the clotted fabrics ofthrombolites to calcification
of coccoid microbial communities. Latham and Riding (1988) recognized
Renalcis Yologdin and Tarthinia Drosdova in thrombolites in the Lie de Yin
Series of Morocco. These calcareous microfossils create the irregular clotted
fabrics of the thrombolites (Fig. 8). Their precise origins are uncertain but they
probably represent cyanobacteria. However, it remains quite likely that there is
a variety of origins for the deposits which have been termed thrombolites. For
example, digitate plumose algal-foraminiferal colonies with a blotchy appear-
ance, termed "leopard rock" in the Lower Permian of New Mexico (Toomey
and Cys 1979, p. 66, Figs. 3b and 4c) resemble thrombolites. At present any
deposit that appears to be microbial, and which is unlaminated and clotted, can
be termed a thrombolite . Future investigations should aim to discriminate
between different types of thrombolites and their various origins.
Fig. 8. Thrombolite. Clotted macrofabric in a domical thrombolite created by clusters of botryoidal

microfossils (Renalcis and Tarthinia) (Latham and Riding 1988) and enhanced by partial
dolomiti za tion of intervening matrix. Upper part of Serie Lie de Vin, Lower Cambrian, Tiout,
Anti-Atlas Mountains. Morocco. Width of view 30 mm
3.5 Travertine
Travertine is named from the old name for Tivoli, east of Rome, where
carbonates deposited from hot springs have been mined for thousands of years.
Travertine is used here to describe a crudely layered carbonate rock, commonly
with a bushy fabric. The latter has been attributed to bacterial activity (see
Chafetz and Folk 1984). The terminology of travertine, tufa and sinter has not
been standardized. Here I regard travertine as a product of warm carbonate
springs where the elevated temperatures, together with the dissolved materials
present in these warm waters, exclude most eukaryotic organisms. The result is
that any mediation of carbonate precipitation in these environments is by
prokaryotes. Consequently the bushy fabrics which are conspicuous in
travertines may be bacterial and/or cyanobacterial in origin, and the shrubs
described as bacterial by Chafetz and Folk (1984, Fig. I) also resemble A n-
gulocellularia, although further work is required to determine precisely what
organisms are present and exactly which processes result in the formation of
travertine.
In fact, at least two types of shrub appear to be involved in travertine
formation: (I) the micritic shrubs, such as those described by Chafetz and Folk
(1984) which may be bacterial or cyanobacterial, and (2) needle shrubs de-
scribed by Casanova (1986, p. 204 and Fig. 4) as bacterial from pool rims of hot
springs in Kenya (see also Tiercelin et al. 1987, plates 32 and 33). Despite
assertion of the microbial nature of these latter needle shrubs, they resemble
inorganic precipitates.
A parallel can be drawn between crudely laminated travertines with bushy
micro fabrics and skeletal stromatolites. I prefer to maintain a distinction
between travertine and skeletal stromatolites, until further work can show how
similar (or distinct) the processes of formation of travertine are. In the absence
of banding and lamination, the bushy fabrics of travertines closely resemble
dendrolite.
3.6 Cryptic Microbial Carbonates
Stromatolites, and the other types of microbial carbonates described above,

generally have quite distinctive macroscale fabrics and features which assist in
their recognition. However, there is a large volume of carbonate rock, especially
in reefal facies, which may be microbial in origin but whose origins are
nevertheless obscure and for which descriptive criteria are also difficult to
establish. These are here termed cryptic microbial carbonates. They lack dis-
tinctive macrofabrics, but typically possess micritic, clotted or peloidal
microfabrics, in which there may also be traces of filaments. Some cement-like
spar fabrics and dolomite crystals have also been interpreted as microbial.
Many of these microfabrics are also present in stromatolites, dendrolites,
thrombolites and travertines. However, these latter also possess characteristic
macrofabrics which are lacking in the case of cryptic microbial carbonates.
38 R. Riding
Consequently, cryptic microbial carbonates have to be interpreted purely on the

basis of micro fabrics, and it is the uncertainties inherent in this which create the
particular difficulties which surround them. It is difficult to establish criteria
which allow confident interpretation of a microbial origin for these micritic,
peloidal and other microfabrics. It is sometimes just as difficult to decide
whether they are in place or allochthonous.
Nevertheless, in recent years a number of workers appear to have had an
innate sense of the presence and importance of these deposits and, despite the
obstacles, there has been a growing attempt to deal with the problem. Much
current work on "microbial carbonates" is, in fact, aimed at this cryptic side of
the subject (Maurin 1987). The origins of this research can be seen in early work
such as that by Drew (1911). Part of its revival can be attributed to the need to
understand "cryptalgal" fabrics (e.g. Monty 1976) and also to detailed scrutiny
of the organisms and the deposits within modern microbial mats (e.g. Krumbein
and Cohen 1977); these were developments ofthe stromatolite work ofthe 1960s
and early 1970s. At the same time, a few workers were starting to report
bacterium-like objects in ancient limestones (e.g. Maurin and Noel 1977).
Subsequent work has involved spars (Monty 1982), Devonian reefs (Monty et
al. 1982), mud mounds (Maurin et al. 1981; Tsien 1985b), Stromatactis (Tsien
1985a), ooids (Monty and van Laer 1984; Castanier et al. 1989), peloids (Chafetz
1986), and dolomites (Monty 1986; Castanier et al. 1988).
Yet problems of establishing organic versus inorganic, and in place versus
allochthonous, origins remain. It is particularly difficult to discriminate between
cause and effect: are microbes associated with a crystal of calcium carbonate
because they have caused its precipitation, or because it is a convenient substrate
to live on, or both, or neither? The way forward most likely lies in laboratory
studies (e.g. Castanier 1987). But apart from these fundamental problems of
microbial biomineralization, a descriptive vocabulary for the microfabrics (and
nanofabrics) of these cryptic microbial carbonates also remains to be estab-
lished. This should allow improved comparison to be made between ancient
examples and modern analogues which, in combination with experimental
work, will probably provide the key to understanding these enigmatic, but
potentially very important, deposits.
4 Composition and Fabrics
Calcareous benthic microbial deposits consist, basically, of one or more of the

following: trapped particulate sediment, biomineralized skeletons, and miner-
alized crust, either surficial or as a void-fill. These may be laminated and/or
dendritic, or clotted in appearance (Fig. 9), and they may possess fenestrae.
These variations in composition and fabric are controlled both by the organisms
involved and by the environment of formation of the microbial carbonate.
Particulate sediment is predominant in agglutinated stromatolites, calcareous
skeletons in skeletal stromatolites, dendrolites and thrombolites, and miner-
alized crusts in tufa stromatolites and travertine. Lamination is good in
a dendrolite
vertical
d tufa stromatolite
fabric
travertine
0- skeletal stromatolite
thrombolite agglutinated and subaerial

stromatolites
I
horizontal fabric
Fig. 9. Preponderance of vertical (and radial) versus horizontal (laminated/layered) fabrics in

benthic microbial carbonates
fine-grained agglutinated stromatolites, and crude in coarse-grained agglu-

tinated, skeletal and tufa stromatolites, and also in travertine. Dendritic and
clotted fabrics are characteristic of dendrolites and thrombolites respectively,
both of which are non-laminated. Fenestrae are especially present in tufa
stromatolites, skeletal stromatolites and coarse-grained agglutinated stroma-
tolites. In all of these they can be produced in subaqueous environments. They
also form in tidal fiat stromatolites.
The interrelationships of lamination, mineralized crusts and biominer-
alized skeletons are shown in Fig. 10 and Table 2. Certain microbial carbonate
types are closely related and therefore may show intergradation. This is par-
ticularly true of (l) skeletal stromatolites and travertine; (2) skeletal stroma-
tolites and dendrolites; (3) skeletal stromatolites and tufa stromatolites; (4)
dendrolites and thrombolite; and (5) tufa stromatolites and travertine.
5 Distribution
There are six main stages in the geological history of stromatolites, dendrolites,
and thrombolites. The following outline applies principally to marine microbial
carbonates. The time distribution of non-marine benthic microbial carbonates
is less-well recorded, although it must be pointed out that it is often not clear
whether Precambrian stromatolites occur in marine or lacustrine environments.
It is likely that there is a direct relationship between the organisms and processes
available for the formation of benthic microbial deposits, and the overall
temporal sequence that they show (Fig.l).
40 R. Riding
Fig. 10. Types of microbial carbonate deposits plotted according to development of lamination,
mineralization and biomineralization. Note that potentially close intergradations occur between
skeletal stromatolites and travertine, skeletal stromatolites and dendrolites, dendrolites and
thrombolites, tufa stromatolites and travertine, subaerial stromatolites and travertine, and tufa
stromatolites and skeletal stromatolites. Note that it is likely that not all thrombolites are dominated
by biomineralization
Table 2. Fabrics and principal components typical of major microbial carbonate types"
Principal components
Calcified Particulate
organism sediment
good fine
Lamination Skeletal stromatolites Agglutinated
poor Tufa stromatolites coarse stromatolites
Travertine
Dendritic Dendrolites
Clotted Thrombolites Thrombolites m

aSubaerial stromatolites have good lamination but appear to be dominated by mineralization
products rather than calcified organisms.
5.1 Archaean
The oldest definite record of stromatolites is from the Warrawoona Group (3.5
Ga) of Western Australia (Dunlop et al. 1978). These occur in nearshore
environments dominated by volcaniclastic sediments. Silicified filamentous
microfossils in cherts from these stromatolites could be either bacterial or
cyanobacterial (Awramik et al. 1983; Schopf and Walter 1983, pp. 232-237).
The Archaean stromatolite record is reviewed by Walter (1983).
5.2 Proterozoic to End-Riphean
Stromatolite abundance increases rapidly during the Early Proterozoic, pos-

sibly in response to both the evolution of cyanobacteria, whose definite fossil
record appears to begin in the Fortescue Group (2.7 Ga) of Western Australia
(Schopf and Walter 1982), and to the progressive development of continental
cratons. Proterozoic stromatolites are characteristically, although not exclu-
sively, fine-grained and well-laminated (A wramik and Riding 1986). It has often
been stated that CaC03 precipitation was significant in their formation (e.g.
Gebelein 1976), although this is not proven and often they resemble agglu-
tinated stromatolites. Nevertheless, microdigitate ministromatolites with a
radial fibrous fabric are a feature ofthe Early and Middle Proterozoic (900-2500
Ma) and have been compared with calcified Rivularia and Phormidium mats
(Walter 1988; Grotzinger 1989). The Riphean (800-1650 Ma) was the climax of
development of branching and conical stromatolites, and these have been
applied to stratigraphical correlation following the lead of Soviet geologists
(Semikhatov 1976).
5.3 Sturtian and Vendian
A wramik (1971) recognized a major decline in both the abundance and diversity
of stromatolites following the Riphean, which he attributed to the appearance
of metazoans. Subsequent studies have confirmed this decline (Walter and Heys
1985). The deleterious effects of metazoans are likely to have been various:
grazing, burrowing, competition for space. To these were added the production
of coarse bioclastic sediment following the onset of biomineralization close to
the base of the Cambrian (Pratt 1982).
5.4 Cambrian-Lower Ordovician
Fine-grained agglutinated stromatolites never recovered from the impact of

metazoan evolution. However, cyanobacterial biomineralization, which ap-
peared as dramatically as that of metazoans near the base of the Tommotian
(Riding and Voronova 1984), resulted in the formation of dendrolites and
thrombolites which are both particularly common in the Cambrian and Lower
Ordovician. Skeletal stromatolites also make their appearance at this time.
5.5 Middle Ordovician to Middle Cretaceous
Calcified cyanobacterial dendrolites, thrombolites and skeletal stromatolites

are sporadically prominent during this long period of overall decline in cal-
careous microbial deposits in marine environments. Dendrolites and throm-
bolites appear to be virtually restricted to the Palaeozoic. Decline is likely to be
due to the increasing success, and therefore, competition, predation and dis-
42 R. Riding
turbance, of metazoans. And, particularly in the case of dendrolites and skeletal

stromatolites, it may also be due to a general long-term reduction in CaCOa
precipitation in the world oceans. Only in tidal flat environments did stroma-
tolites (represented by fine-grained agglutinated forms) retain their overall
success; and this appears to clearly relate to the inhibition of metazoans by
strongly fluctuating physical and chemical parameters in these environments
(Garrett 1970).
5.6 Mid-Cretaceous to Recent
Calcified cyanobacteria disappear, or virtually disappear, from normal subtidal

marine environments sometime during the Mesozoic-Cenozoic transition
(Riding 1982). Probably the final occurrence offairly common marine calcified
cyanobacteria was during the Middle Cretaceous, but this remains to be
confirmed. During the Cenozoic stromatolites have a fine record in lacustrine
environments, for example in the Middle Eocene Green River Formation of
Wyoming (see Surdam and Wray 1976), but the principal marine occurrence of
stromatolites appears to have been the tidal-flat environment, and this can be
seen as a refuge for fine-grained cyanobacterial-dominated stromatolites
(Garrett 1970; Awramik and Riding 1988). Undoubtedly, the conspicuous
survival of this type of stromatolite in this environment led to the "intertidal
dogma" of stromatolitology (Hoffman 1973, p. 181) following Black's (l933)
work in Andros. However, a significant new development has been the origin of
the eualgal-cyanobacterial stromatolite-forming community which character-
istically produces coarse-grained stromatolites (Awramik and Riding 1988).
These are exemplified by the Recent well-known subtidal Shark Bay and Lee
Stocking columnar and domical forms. Both diatoms and filamentous green
algae are conspicuous in the formation of these distinctive stromatolites which
have widely, but probably mistakenly, been regarded as analogues of
Precambrian stromatolites. It remains to be seen whether eualgal-cyanobac-
terial stromatolites date from the time of appearance of diatoms, in the Cre-
taceous, or earlier. So far no closely similar fossil stromatolites have been
identified.
6 Outline History of Research
The history of work on microbial carbonate deposits is dominated by research

on stromatolites. This has been reviewed a number of times (Maslov 1960;
Hoffman 1973; Hofmann 1973; Monty 1977; Krumbein 1983).
Twentieth-century progress in stromatolite studies can be characterized as
a succession of dogmas (Monty 1977). Walcott's (1914) views concerning
Proterozoic Belt Supergroup stromatolites in Montana led to a general fresh-
water interpretation until Black's (1933) study of modern Bahamian stroma-
tolites redirected attention to tidal flats and also introduced the important
tra pping and binding concept of forma tion. During the 1960s and 1970s, in terest
in morphological classifications of stromatolites was coupled with the
development of Proterozoic stromatolite biostratigraphy. Two results of this
have been the production of a fairly complex Linnean-style nomenclatural
system, especially for branching stromatolites (for discussion see Cloud and
Semikhatov 1969; Walter 1972, pp. 14-27; Semikhatov 1976), which still needs
to be revised and consolidated into a readily applicable scheme and, on the other
hand, a rather over-simple morphological descriptive terminology (Logan et al.
1964) which awaits refinement.
Simultaneously, the search has gone on for modern analogues of ancient
stromatolites. Much has been learned concerning the biology of modern
microbial mats, particularly the complex physiological adaptations which are
linked to the vertical stratification of various bacterial and cyanobacterial
communities within the mats (see papers in Cohen et al. 1984). However, the
lack of obvious information preserved in most ancient stromatolites has hin-
dered progress in extrapolating results from the modern to the ancient.
Nevertheless, it does seem likely that petrographic evidence in ancient
stromatolites still awaits interpretation. Most studies of Precambrian stroma-
tolites, for example, have been made by palaeobiologists and stratigra phers who
may have overlooked useful, if somewhat cryptic, information within the
carbonate microfabrics. Recognition that the modern coarse-grained, crudely
laminated subtidal Shark Bay stromatolites are really not closely comparable
with the ancient fine-grained well-laminated fossils with which they have
generally been compared (Awramik and Riding 1988) illustrates the type of
comparative approach which may lead to significant new insights into
stromatolites.
Stromatolites can form in hot springs (Doemel and Brock 1974) and work
on hot spring sinters (geyserites) has been related to stromatolite research by
Walter (l976b). However, in general, travertine and geyserite studies have
developed separately, and more slowly, than those on stromatolites, and have
been more closely linked with work on tufa. Travertines and tufas are both
non-marine deposits, and there has been terminological confusion between
them, usually with fluviatile tufas being termed travertine (e.g. Love and
Chafetz 1988), rather than the other way round. Work on tufa, especially, has
often been concentrated on attempting to discriminate between organic and
inorganic origins (e.g. Emeis et al. 1987). Tufa dams in rivers have been termed
phytoherms (Pedley 1987).
Fluviatile and lacustine oncoids have long attracted attention (see Fritsch
1959, p.868, and references therein), but their in-place counterparts, tufa
stromatolites, are probably among the most neglected of all the microbial
deposits considered here, possibly because of their intergradation to, and
incorporation with, more diverse tufa deposits.
Stromatolites and other microbial carbonates reviewed here possess
macroscopically distinctive fabrics. However, research has also been moving
towards studies of less distinctive, and often obscure, fabrics which may be
microbial in origin and are here termed cryptic microbial carbonates. These
44 R.Riding
include an array of micritic, clotted, peloidal and spongiostrome microfabrics

(see Maurin and Noel 1977; Monty et al. 1982; Chafetz 1986). In hindsight, we
can see that research on some of these was begun by earlier workers (Giirich
1906; Pia 1927; Maslov 1960).
Introduction of a microbolite, rather than specifically stromatolite, orien-
tated approach (Burne and Moore 1987) to this field will no doubt assist
comparative studies. Kennard and James (1986) emphasized that thrombolites
and stromatolites are two classes of microbial structure. This present paper
emphasizes dendrolite as yet another type. Dendrolites, although not hitherto
dignified by a specific name, have been obvious fabrics within Early Palaeozoic
reefs for many years. Thrombolites have themselves been named for more than
twenty years. By drawing these deposits, along with travertine, into a broad
category of calcareous benthic microbial deposits we can now begin to see the
similarities which they share, alongside their differences, and thereby more
precisely interpret their origins, distribution and development.
7 Conclusions
Riding (1977, Fig. 1) used a triangular diagram (Fig. 6) to show the relationships
between the processes of carbonate deposition (sediment trapping, physico-
chemical precipitation, biochemical trapping) and the products: non-skeletal
stromatolites (which here are termed agglutinated stromatolites), tufa, and
skeletal stromatolites. The same diagram can be used to express the array of
microbial carbonates described here (Fig. 11).
Agglutinated stromatolites are the classic stromatolites of the stromatolite
literature, referred to as 'normal' or nonskeletal stromatolites by Riding (1977,
p. 57). It is precisely because they have been recognized as the most common
type of stromatolite that they have not been given a more specific name until
introduction of the term agglutinated stromatolite here. Fine-grained agglu-
tinated stromatolites are probably dominantly prokaryotic in origin and range
from the Archaean to Recent. However, there appear to be no good modern
analogues for the complex branching forms which they produced during the
climax of their development during the Riphean. The Shark Bay and Lee
Stocking subtidal stromatolites not only differ in morphology, but are coarse
grained and crudely laminated: features which can be attributed to the
prominent eualgal component of their mat community. The marine refuge of
fine-grained agglutinated stromatolites is the tidal flat environment, which is
somewhat similar to the Precambrian in exhibiting reduced metazoan com-
petition and abundant fine-grained sediment, but in which the stromatolites
characteristically are flat and crinkly laminated due to the exposure character-
istic of this environment. It is likely that modern lacustrine environments
possess better, although nevertheless not precise. analogues for subtidal Pre-
cambrian stromatolites.
Although it is probable that the carbonate microfabrics of fossil agglutin-
ated stromatolites possess valuable information concerning their origins which
TRAPPING
"'.....1'0 II'
~~
'9..... ~
SUBAERIAL ~;;
STROMATOLITES DENDROLITES ~'(
TUFA STROMATOLITES THROMBOLITES '"
SURFICIAL MINERALIZATION BIOMINERALIZATION
Fig.H. Triangular diagram adapted from Riding (1977, Fig. I; see Fig. 6) showing major microbial
carbonate types according to processes offormation. Note that it is likely that not all thrombolites
are dominated by biomineralization. Skeletal stromatolites overlap with thrombolites and den-
drolites on this diagram. The lamination which distinguishes stroinatolites is due to episodic
accretion and/or prostrate orientation of organisms. Biomineralization here is regarded as
precipitation within organisms, or within organic tissue, during life. Surficial mineralization is
external to organisms and their tissues, but does not here preclude the possibility of organic inft uence
on this precipitation
remains to be exploited, they nevertheless retain relatively little information

concerning the organisms responsible for their formation. This is not the case
with biomineralized and mineralized microbial deposits. Skeletal stromatolites,
tufa stromatolites, dendrolites, some thrombolites and travertine all possess
fabrics which reflect the size, shape and orientation of individuals, or clusters of
individuals, of mat-forming organisms to varying extents. Much, however,
remains to be learned about all these categories of calcareous microbolite. In the
case of mineralized varieties (tufa stromatolites, subaerial stromatolites,
travertine) superficial crusts preserve, but also complicate the morphology of the
organisms, whereas biomineralized fabrics within skeletal stromatolites, den-
drolites and some thrombolites probably retain clearer details. Of these latter
three deposits, skeletal stromatolites possess at least poor lamination, and this
distinguishes them from dendrolites and thrombolites. If they are mineralized,
then skeletal stromatolites equal tufa stromatolites. If they are unlaminated,
then they may equal dendrolites. Whether or not all varieties of thrombolite
contain an important component of calcified microfossils remains to be
elucidated; it is still likely that non~skeletal thrombolites will be distinguished.
46 R.Riding
Ifunlami~ated, travertine may equal dendrolite. But travertine can also be

viewed as intermediate between tufa and calcareous sinter, both with respect to
lamination (tufa unlaminated, travertine crudely laminated, sinter well
laminated) and porosity (tufa, high porosity; travertine, medium to low poro-
sity; sinter, zero porosity). As a rough generalization, tufa is formed by
precipitation ofCaC03 from cold water onto cyanobacteria, algae, moss, grass,
etc. Travertine is formed by precipitation from warm water mainly onto
cyanobacteria and bacteria, although diatoms may also be present; sinter is
either precipitated from hot water (at the Earth's surface) in the presence of
bacteria and subsidiary cyanobacteria, or from cold water in caves where
bacteria may be present. Elevated temperatures act to progressively exclude
eukaryotes.
Cryptic microbial carbonates possess micritic clotted, peloidal or sparitic
microfabrics, similar to those of evident benthic microbial carbonates such as
stromatolites, etc., but they lack distinctive macrofabrics. They are probably
widespread and important in Phanerozoic reeffacies. Criteria are still required
to distinguish them from particulate or cement fabrics of inorganic origin, and
to elucidate the particular micro-organisms and processes responsible for their
formation.
Since benthic microbial carbonates vary in time and space, and also
according to the organisms and precipitating fluids involved, there are notable
contrasts between the different types of deposits which have been reviewed here.
In hindsight, therefore, it is not surprising that difficulties have been encoun-
tered in the past in attempting to draw too close analogies between, for example,
freshwater and marine, and modern and ancient microbial carbonates. The
present is only a key to the past if the locks have not been changed.
Appendix
Summary of terms used here
Agglutinated stromatolite (this paper), produced by trapping/binding of par-

ticulate sediment.
Biomineralization, mineral precipitation within organisms, or within organic
tissue, during life.
Cryptic microbial carbonate (this pa per), benthic microbial deposit with micritic,
clotted, peloidal or sparitic microfabrics, but lacking distinctive
macrofabrics.
Cyanoid (Riding 1979, 1982), oncoid constructed by calcified cyanobacteria.
Dendrolite(Riding 1988), non-laminated biomineralized microbial deposit with
a dominant dendritic macrofabric.
Microbial carbonate, carbonate deposits produced or localized by benthic
microbial communities.
Microbolite (Burne and Moore 1987, name emended here), benthic microbial
deposit.
Mineralization, mineral precipitation on inorganic surfaces and on external

surfaces of organic tissue.
Nonskeletal stromatolite (Riding 1977), stromatolite in which the microorgan-
isms responsible for its formation are not calcified. Equivalent to agglu-
tinated stromatolite.
Oncoid (from oncolith, Pia 1927), nodular, originally loose, stromatolite.
Oncolite (from oncolith, Pia 1927), lithified deposit of oncoids.
Sinter, dense, laminated deposit, precipitated often as an incrustation upon
rock.
Skeletal stromatolite (Riding 1977), stromatolite in which the organisms re-
sponsible for its formation are commonly preserved as calcified fossils.
Stromatolite (from stromatolith, Kalkowsky 1908), laminated benthic microbial
deposit.
Subaerial stromatolite, microbial calcrete, formed at subaerial surfaces and
dominated by desiccation which promotes rapid lithification; characterized
by pedogenic fabrics.
Thrombolite (Aitken 1967), non-laminated benthic microbial deposit with a
dominant clotted macrofabric.
Travertine (from the ancient name for Tivoli, Italy; tivertino, tiburtinus),
layered deposit of CaC0 3 with moderate to low primary porosity, com-
monly with a dendritic macrofabric, produced by precipitation associated
with warm springs.
Tufa (an old Italian word from the La tin name tofus for a porous rock), originally
porous, laminated to massive deposit mainly produced by precipitation of
minerals on organic substrates in freshwater.
Tufa stromatolite (this paper), stromatolite produced by precipitation of min-
erals on (rather than within) organic tissue.
Acknowledgements. My interest in tufas was stimulated by Ernst Ott and supported by an Alex-
ander von Humboldt-Stiftung research fellowship at the Technical University, Munich. Anna
Gandin introduced me to travertines in Tuscany and TOTAL, through the encouragement of Andre
Maurin. supported their study. Discussions with Stanley Awramik, facilitated by NATO Research
Grant 84/0176. were a fertile source of insights into stromatolites past and present; nota few of these
views are included here, and I am indebted to Stan for the stimulus. and the fun. which our debates
created. A visit to Lee Stocking Island was supported byThe Royal Society. Constructive comments
by Stanley Awramik and V. Paul Wright improved the manuscript.
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II Major Groups
Chapter 3
Calcified Cyanobacteria
R. RIDING l
Abstract
Calcification affects the sheath of some coccoid and filamentous cyanobacteria during their life,
under suitable environmental conditions. There is probably a link between sheath character, the
environment and nucleation ofCaC01 within or upon the sheath. Calcification is thus extracellular
and is influenced, but not fully controlled, by the cyanobacterium. It is promoted by thick sheaths,
which provide favourable sites for CaCO" nucleation, and by conditions favouring the physico-
chemical precipitation ofCaCO,.
In the Recent, calcification is widespread in freshwater lakes and streams in limestone areas,
but is virtually unknown in marine environments. However, calcification occurred quite widely in
marine cyanobacteria during the Palaeozoic and Mesozoic. This could reflect temporal changes in
oceanic carbonate chemistry.
Calcification results in (I) discrete skeletons; (2) skeletal stromatolites; (3) dendrolites and
thrombolites; and (4) tufa stromatolites. Where calcification is uncommon the mucilaginous sheaths
and felted masses of cyanobacterial mats trap particulate sediment to form stromatolites which may
be calcareous in composition, but which do not consist of calcified cyanobacteria in the sense of
precipitation ofCaCO" on or within the sheath.
The main groups offossil calcified cyanobacteria may be exemplified by the following genera:
Angulocellularia, Epiphyton, Girvanella, Hedstroemia, Renalcis and Garwoodial Mitcheldeania,
although some of these are not definitely known to by cyanobacteria. In addition there is a variety
of fossil calcareous microproblematica which have been widely attributed to the cyanobacteria.
Well-defined and diverse calcified cyanobacteria first appear near the Precambrian-Cambrian
boundary. Dendrolites and thrombolites composed of probable cyanobacteria are common in the
Cambrian and Lower Ordovician. Subsequently, calcified cyanobacteria are generally less common
in marine environments, although they reappear in relative abundance in the Upper Devonian and
Lower Carboniferous, and in the Middle Triassic to mid-Cretaceous. A number of these fossils may
be compared with modern analogues in freshwater lakes and streams, but the precise affinities of
others are still uncertain.
1 Introduction
Cyanobacteria, also known as blue-green algae and cyanophytes, are prokar-

yotes. They are thus more closely related to bacteria than to eukaryotic algae.
Cyanobacteria are responsible for a variety of calcified microfossils, such as
Girvanella Nicholson and Etheridge and Ortonella Garwood, and are also
significantly involved in stromatolites formed by trapping of sedimentary
particles.
Calcified cyanobacteria are individually normally microscopic, but the
deposits which they construct, such as oncoids, stromatolites, tufas and reefal
I Department of Geology, University of Wales College of Cardiff, Cardiff CFl 3YE, UK

56 R. Riding
fabrics, are macroscopic and conspicuous. Thus, they may occur either as very
small individual skeletal fossils, or be amalgamated to form stromatolitic and
reefal deposits. In both cases they have a range in marine environments from the
earliest Cambrian to late Cretaceous or early Cenozoic. Oncoids and stroma-
tolites are also formed by non-calcified cyanobacteria. Consequently, it is
necessary to check, by microscopic examination, whether stromatolite fabrics
are composed mainly by agglutinated sediment particles or by calcified cyano-
bacteria. In the Recent, calcified cyanobacteria are often conspicuous in
freshwater calcareous rivers and lakes where analogues for marine fossils exist
and provide information concerning these simple, but important, fossils.
Calcified cyanobacterial sheaths are commonly irregular bushy masses, or
tubes arranged in radial fans or tangled masses. This simplicity, coupled with
their small size, creates rather than reduces systematic problems. Gradation
between taxa makes it difficult to establish diagnoses, due to lack of distin-
guishing features and variation in sheath shape. The tubular construction of
filamentous sheaths causes some of them to be confused with some small green
and red algae and invertebrates. A wide range of fossils of uncertain affinities,
which should properly be regarded as microproblematica until their relation-
ships are better understood, have been referred to the cyanobacteria. Some
calcified cyanobacteria, particularly forms such as Cayeuxia Frollo and Or-
tonella, have also been confused with udoteacean green algae.
Calcified cyanobacteria have a distinctive distribution as fossils in marine
environments. They are relatively common in the Palaeozoic and Mesozoic, but
are rare or absent in the Precambrian and Cenozoic. Recent marine calcified
cyanobacteria are virtually unknown, although freshwater tufas and oncoids
commonly contain them in abundance. This long-term pattern: Precambrian
scarcity, Palaeozoic-Mesozoic presence, followed again by Cenozoic scarcity of
marine calcified cyanobacteria contrasts with the geological range of non-
mineralized cyanobacteria. The group as a whole possesses the longest range of
any class of organism, apart from bacteria. Cyanobacteria are known from the
late Archaean onwards. An explanation for the appearance and disappearance
of calcification in marine cyanobacteria may lie in a change, probably subtle but
nonetheless significant, in sea-water chemistry (Riding 1982), although it is also
likely that evolutionary events, possibly involving specificity for calcification,
have interacted with this environmental control (Riding 1989). The calcification
site in living cyanobacteria appears to be essentially extracellular and lies in the
mucilagenous protective sheath. This accounts for both the environmental
dependence of cyanobacterial calcification, and also for the morphological
simplicity of the resulting fossils.
2 Calcification
Cyanobacterial calcifica tion is reviewed by Pen tecost and Riding (1986) and has
been described in detail from freshwater environments by Pentecos~ (1987,
1988). Two key features are: (1) the site of calcification is the enveloping sheath.
Calcified Cyanobacteria 57
rather than the cells, of the organisms, and (2) calcification appears to proceed
only under conditions where CaC03 precipitation is thermodynamically
favoured. The result of calcification is the formation of discrete fossils whose
biomineralized sheaths rank as skeletons (see Discussion), albeit very small
ones, if they function for support and protection of the organism. Numerous
individual calcified cyanobacteria may be amalgamated into dendrolites,
thrombolites and other large reefal masses or, if they possess lamination, into
skeletal stromatolites and skeletal oncoids (cyanoids, Riding 1983).
2.1 Processes
Cyanobacterial calcification is sited in or on the mucilaginous sheath. The

sheath itself is secreted by the cell for which it forms a protective envelope (Fig.
I). Coccoid cells are rounded and form clusters within a mass of sheath material.
Filamentous forms consist of hairlike trichomes which are strands of cells
attached end-to-end. One, several, or a number of trichomes may share a
common tubular sheath . Trichomes and sheath together are then termed a
filament. If the trichomes branch, or are arrayed in a fan-shape , then the
resulting sheath may have a bush-like or radial form.
Sheath production is a prerequisite for calcification, but under conditions
where there is little need for protection from water-movement, sediment
particles, sunlight, desiccation or grazing organisms the sheath may develop
only weakly or not at all. Cyanobacteria under laboratory culture commonly
produce abundant soluble polysaccharides, but do not usually develop sheaths
(Rippka et al. 1979, pp. 23-24). They consequently differ from specimens in
natural environments, not only in the absence of the sheath but also in features
which may be sheath-related, such as branching-style and colony form .
The presence of a sheath, and also its consistency (whether dense or
diffluent), influence calcification because the crystals ofCaC0 3 nucleate within
Fig. ia,b. Morphology of a filamentous.

and b coccoid. cyanobacteria.
The sheath is the normal site of
calcification
58 R. Riding
or upon it, possibly as a result of the attraction of calcium ions by uronic acids.
Trapping of sediment grains by cyanobacteria, forming agglutinated stroma-
tolites, is also primarily effected by the sheath; the grains may then be bound into
a felted mass of filaments by subsequent cyanobacterial growth and movement.
Calcification results in either impregnation or encrustation of the sheath, or a
combination of these (Fig. 2). In impregnation, CaC03 crystals nucleate within
the sheath. They are generally small, bladed, and may show orientation in
response to the sheath structure. In encrustation, CaCOa crystals nucleate upon
the sheath and generally form larger crystals. They range in form from simple
rhombs to complex dendritic crystallites (Pentecost and Riding 1986).
Of equal importance to the presence of a sheath for calcification, is a
calcareous environment. Cyanobacterial metabolism may stimulate CaC03
precipitation by uptake of CO 2 or HCO; but, so far as is presently known, the
physico-chemical conditions must also favour calcification. This is readily
achieved in calcifying freshwater streams where turbulence promotes CO 2
evasion and leads to tufa deposition. However, in marine environments at the
present-day, even where early cementation is occurring, cyanobacterial
calcification appears at best to be rare, although it was common during the
Palaeozoic and Mesozo ic.
CALCIFICATION: FORMATION OF CALCIFIED MICROFOSSILS
Internal and external

crystal growth
external Internal
encru,tatlon Impregnation
Trichome
agglutination
of ,edlment no calcification
grains no agglutination
AGGLUTINATED NO CALCAREOUS
STROMATOLITE FORMATION FOSSIL PRESERVATION
Fig. 2. Filamentous cyanobacterium in transverse section to show sites and styles of calcification
(upper part of diagram) and sediment agglutination (lower left part of diagram). Calcification may
be either by external encrustation (upper left) , or internal impregna tion (upper right) of the sheath,
or by a c om bination of these (upper centre). Calcification will result in the formation of calcified
microfossils. Agglutina tion of particulate sediment will result in the formation of agglutinated
stromatolites
2.2 Products
Individuals or colonies of calcified cyanobacteria can occur as discrete skeletons,

or they may be arrayed in a variety of massive deposits, which can be divided on
the basis of macroscopic fabrics into skeletal stromatolites and oncoids
(laminated), thrombolites (clotted), dendrolites (dendritic), and closely spaced
microbush fabrics, as well as less distinctive reefal masses (Fig. 3). Where these
deposits are marine, they are often associated with skeletal invertebrates. In
freshwater they are often associated with enhanced physicochemical precipi-
tation which promotes external encrustation, and then form tufa. Travertine
deposits formed by warm springs may contain cyanobacteria (Copeland 1936),
but they have been reported to be dominated by bacteria (Chafetz and Folk
1984). They require further study.
2.2.1 Discrete Skeletons
Calcification of the sheaths of individual filaments or of filamentous colonies

results in morphologies based on tubes or bush-shaped masses (Fig. 4). The
1. INDIVIDUAL MICROFOSSILS
a~
2 . LAMINATED
~~~ PROSTRATE
-===- --- ~
skeletal
stromatolite
thrombolite
microbushes
dendrollte
3 . MASSIVE
Fig. 3. Modes of occurrence of calcified cyanobacteria. 1 Discrete individual microfossils (e.g. a
Girvanella; b A ngu/ocellu/aria; c Ortonella). 2 and 3 Amalgamated masses, which may be either 2,
laminated (skeletal stromatolites) or 3, massive (microbushes, thrombolite, dendrolite). Skeletal
stromatolites may, within their layers, have either erect or prostrate fabrics , depending upon the
orientation of the calcified microfossils. Examples of massive amalgamated masses include. for
microbushes, Lower Ordovician Angu/ocellll/aria ("Epiphyton , Rena/cis", Riding and Toomey
1972). and Upper Devonian "fenestral renalcid micrite" (Mountjoy and Riding 1981); for
thrombolite, Lower Cambrian Rena/cis and Tarlhinia thrombolites (Latham and Riding 1988); and
for dendrolite, Lower Cambrian Epiphylon.Rena/cis·Angu/ocellu/aria fabrics (Riding, this Vol.)
60 R. Riding
UNORIENTATED BRANCHED
100.,"'' '" Y'"""'"

TUBE OBRUCHEVELLA
~Olled I BOTOMAELL Af ORTONE LLA

tangl~
GIRVANELLA ~ allgned
cab l a-lika ~
HEDSTROEMIA / CA YEUXIA
SUBTIFLORIA
MICRITIC
,..TH'." V GORDONOPHYTON
•• ",.,," ~ACC" IZHELLA

cJi~
~
~ et c.
~ GEMMA
T ANINIA
I
Fig. 4. Simplified outline of the main morphologies of fossil calcified cyanobacteria together with
possible cyanobacteria of problematic affinity. Exa mples shown are believed to be impregnated
shea ths (rather than encrusted sheaths, see Fig. 2). Three principal form s (tubes solid micritic, and
hollow ) ex hibit organiza tions ranging from unorientated to erect and branched. Two of the tube
forms (Obruchevella , S ub/if/oria) show coiling or alignment without obvious substrate attachment.
Named genera a re a selection of principal examples. See a lso Ridingand Voronova (1985) fordetails
of this type of morphologica l approach to Cambrian calcareous "algae", many of which may be
cyanobacteria l
tubes are generally either erect radial clusters (e.g. Ortonella, Cayeuxia) or are
prostrate felts (e.g. Girvanella) or bundles (e.g. Subtifioria Maslov). The bush-
shaped masses, such aSAngulocellularia Vologdin, usually appear to be densely
micritic, but Recent examples formed by Schizothrix calcicola (Agardh)
Gomont show tubular traces of trichomes within the bush (Riding and Vo-
ronova 1982a). A dendritic appearance, but with a dense rod-like rather than
tubiform structure, is also shown in members of the Epiphyton-group . It is
possible that Epiphyton Bornemann and its allies represent calcified filamentous
cyanobacteria: their size and morphology are consistent with this, and
Tubomorphophyton Korde, which appears to be related to Epiphyton, has been
interpreted as a cyanobacterium (Riding and Voronova 1982b). Thus,
filamentous forms are known in the Recent to produce tubiform and bush-like
calcified sheaths, and it is possible that fossil rod-like dendritic forms are also
produced by filamentous cyanobacteria. Members of the Epiphyton-group are
only known for certain from the Palaeozoic and, in the absence of Recent
analogues, their exact affinities remain uncertain.
Calcified coccoid forms are less well-known. They can be expected to occur
as either micro-spherulites, where small clusters of individual cells are calcified,
or as botroyoidal and reniform masses (which may show vertical elongation)
where the mucilaginous sheath of larger coccoid colonies are peripherally
calcified. Possible examples of these are the spherulites in Pleistocene tufas of
the Dead Sea area (Buchbinder 1981), and the Renalcis-group fossils which are
common in the Cambrian and range into the Carboniferous (Hofmann 1975;
Pratt 1984; Fig. 4). Pratt (1984) has also suggested that Epiphyton is a coccoid
cyanobacterium. But the fact remains that because Recent calcified coccoid
forms are poorly known, their ancient counterparts are difficult to interpret.
2.2.2 Stromatolites
Stromatolites formed by calcified cyanobacteria have been referred to as

skeletal stromatolites (Riding 1977a) because they are distinguished from
agglutinated stromatolites. by possessing calcified microfossils which both
construct them and show how they originated. Agglutinated stromatolites
(Riding, this Vol.), in contrast, are essentially sediment grains trapped by a
microbial mat which has since been destroyed and whose precise original
component organisms are usually unclear.
Calcification of cyanobacteria hardens the otherwise sticky sheath and
inhibits agglutination of particles. However, erect filaments are preserved in
growth posi tion by calcifica tion and this promotes tra pping of particles between
the calcified tufts. Thus, growth rates may be faster for skeletal than for
agglutinated stromatolites, and the erect growth form of many filamentous
cyanobacteria results in the formation of relatively large irregular fenestrae
between the filament clusters which are constructional in origin and not related
to desiccation or gas bubbling. At the' same time, lamination is cruder than in
most agglutinated stromatolites and may disappear entirely. When lamination
is altogether absent then the deposit must be regarded as non-stromatolitic (see
Reefs, below).
Nodular, unattached stromatolites are termed oncoids, and skeletal oncoids
formed by calcified cyanobacteria ha ve been termed cyanoids (Riding 1983). As
in skeletal stromatolites, the lamination is made cruder by the irregularities
created by calcification of the cyanobacteria, and cyanoids may acquire an
additional radial fabric due to the preservation of erect filaments (Fig. 5).
Cyanoids are a type of coated grain; they possess a nucleus, usually a rock or
62 R. Riding
LAMINA with rad ial

ij2~~~~~- Interna l fab ri c
NUCLEUS
Fig. 5. Typical skeletal oncoid produced by cyanobacterial calcification. Note that the laminae
thicken towards the upper surface. Episodic, usually seasonal, growth results in lamination, whereas
the erect growth of individual filaments may impart an additional radial fabric
shell fragment (or a piece of grass or reed in freshwater environments), and a

layered cortex composed of calcified sheaths. Recent cyanoids are best
developed in calcareous freshwater lakes and streams where the most common
constituent cyanobacteria are species of Calothrix Agardh ex Bornet and
Flahault, Phormidium Ktitzing and Schizothrix Ktitzing. In lakes cyanoids are
generally small, typically a few centimetres in size, but they may form spheroids
up to several decimetres in size in fast-flowing rivers where turbulence promotes
CaC03 precipitation. However, in addition to sheath impregnation, fluviatile
freshwater oncoids and stromatolites both commonly possess surficial CaC03
sheath encrustation, similar to the crusts which form tufa deposits, and they
grade into tufa stromatolites (see Tufa, below). Consequently, freshwater
examples of cyanoids and skeletal stromatolites are only partial analogues of
fossil marine examples in which calcification during life is more strictly limited
to the sheath. Neither skeletal stromatolites nor cyanoids are known in Recent
marine environments.
2.2.3 Reefs
The hard substrates created by skeletal stromatolites and cyanoids (and den-
drolites and thrombolites, see below) are attractive to both encrusting organisms
and to endoliths. In restricted marine environments, and in freshwater, en-
crusters will generally be limited to molluscs and worms, but in ancient normal
marine waters the diversity of encrusters may result in complex fabrics whose
stromatolitic, dendrolitic, or thrombolitic origin is obscured by an inverte-
brate-dominated reef structure. Calcified cyanobacteria are particularly con-
spicuous in Cambrian, Lower Ordovician, Upper Devonian, and Middle to
Upper Triassic reefs (see Geological History below).
2.2.4 Dendrolites, Thrombolites, and Microbushes
Calcified erect bushes and radial tufts of filamentous cyanobacteria, especially

such as members of the Epiphyton , Angulocellularia and other dendritic groups
(see Table I in Riding and Voronova 1985), if arranged en echelon in vertical
section, rather than being in layers, form unlaminated masses which are not
stromatolitic. These may form dendrolites, defined as: microbial deposits with a
dominant dendritic macrofabric (Riding 1988), or closely spaced microbush
fabrics. No Recent marine examples of these are known but they are locally
important components of Palaeozoic reefs, particularly in association with
archaeocyaths and stromatoporoids. The absence of lamination, presumably
due to lack of well-defined periodicity in their growth, is important in distin-
guishing dendrolites, thrombolites and micro bush fabrics from skeletal
stromatolites (Fig. 3).
Dendrolites are especially prominent in Cambrian reefs, but following the
diversification of middle Palaeozoic reef-builders in the Ordovician they may be
obscured, as are skeletal stromatolites, by associated invertebrates. Microbush
fabrics are nevertheless well-seen in many Lower Ordovician (Riding and
Toomey 1972) and Upper Devonian (e.g. Mountjoy and Riding 1981) reefs.
Thrombolites (Aitken 1967) are non-laminated microbial carbonates with
a clotted fabric. They appear to be preferentially developed in the Cambrian
and Ordovician (Kennard and James 1986). Thrombolites are similar to den-
drolites in that they may be formed by calcified microbes, but thrombolites may
form domes which resemble stromatolites, whereas dendrolites typically form
reefal masses. Uncertainty exists concerning the range of structures which may
be included within thrombolites. The nature of the thrombolites originally
described by Aitken still needs to be established in detail. Walter and Heys
(1985; p. 164) have suggested that thrombolites may represent burrowed
stromatolites. Nevertheless, domical thrombolites from the Lower Cambrian of
Morocco have well-developed clotted fabrics produced by masses of the cal-
cified botryoidal genera Renalcis and Tarthinia (Latham and Riding 1988),
which may be cyanobacteria.
2.2.5 Tufa
Cyanobacteria are common in calcareous streams and lakes where rapid

carbonate deposition on plant surfaces creates porous tufa. Some cyanobac-
teria, such as Calothrix and Phormidium, possess some degree of specificity for
carbonate precipitation which results in impregnation of their mucilaginous
sheaths by CaCOa crystals (see Pentecost and Riding 1986). In other cases,
cyanobacteria, together with green algae and mosses, merely provide a substrate
upon which essentially inorganic surficial mineral encrustation occurs. Where
the substrate is occupied by a variety of these organisms the interaction of
different sizes, shapes, and growth rates results in a thick and generally un-
structured deposit. However, where cyanobacteria are the dominant organisms
their small size and coordinated seasonal growth result in a layered deposit
which may be termed tufa stromatolites and tufa oncoids (Riding, this Vol.).
These occur commonly as oncoids in temperate freshwater calcareous streams
and rivers, but they may also create domical biohermal structures in lacustrine
environments (Eggleston and Dean 1976; Braithwaite, Casanova, Frevert and
Whitton 1989; Winsborough et al. in prep.).
64 R. Riding
3 Classification
Classifications of calcified cyanobacteria have been included in Pia (1927),

Maslov (1956), Johnson (1961), Vo10gdin (1962), Korde et al. (1963), Roux
(1985), and Chuvashov et al. (1987). In addition, broad groupings of Palaeozoic
calcareous algae and cyanobacteria have been outlined by Chuvashov and
Riding (1984). However, no stable classification has yet emerged. Pia's (1927)
concept of the Porostromata has been useful in generally defining small
tubiform fossils, some of which form microfabrics within stromatolites, but this
has included genera such as Rothpletzella Wood, which probably are not
cyanobacteria. On the other hand the Porostromata, despite Pias' (1927) in-
clusion of Epiphyton in it, does not readily encompass solid (as opposed to
tubiform) fossils such as Angulocellularia which are likely to be cyanobacterial
in origin. Consequently, the Porostromata is a heterogeneous group which,
while encompassing some non-cyanobacterial microprob1ematica, neverthe-
less, cannot incorporate botryoidal (such as Renalcis) and solid calcified
cyanobacteria.
At generic level the principal problems for classification concern the status
oftaxa such as: (1) Garwoodia Wood/ Mitcheldeania Nicholson, which have also
been regarded as chlorophytes; (2) Rothpletzella and Wetheredella Wood,
which have at times been assigned to the cyanobacteria; (3) Renalcis and
Epiphyton, which are abundant in the Cambrian and have relatives in the
Devonian, but for which no good Recent analogues have yet been clearly
demonstrated; and (4) Tubiphytes, an important component of some Permian
and Triassic reefal facies which has at times been referred to the cyanobacteria.
With these reservations, which are examined further below, the following
principal groups are recognized here: Angulocellularia and Epiphyton groups,
which are respectively bushy and dendritic in general form; Girvanella group,
filamentous and unbranched; the Hedstroemia group, filamentous and
branched; and the Renalcis group, which is botryoidal to bushy in external
morphology. In addition, there is the Garwoodia/ Mitcheldeania group whose
members are similar to the Hedstroemia group in being filamentous and
branched, but generally larger, and an assortment of microproblematica which
have often been attributed to the cyanobacteria. It should be noted that the
Epiphyton, Girvanella and Hedstroemia groups are quite diverse and could
readily be further subdivided.
3.1 Angulocellularia Group
Features: Micritic, solid, bush-like form, erect or pendant growth.
Example: Angulocellularia Vologdin.
Affinities: Calcification of the sheath of Schizothrix calcicala (Oscillatoriaceae)

produces the micritic, bush-like dendritic skeleton known as Angulocellularia
(Riding and Voronova 1982a; Fig. 6). The divergent trichomes of the cyano-
Fig. 6. A ngulocellularia Vologdin. Recent.

Squaw Island, Conandaigua Lake, New York
State. xl15
bacterium within a common sheath produce the bush-like form, and im-
pregnation of the sheath by precipitation of micritic crystallites, apparently
during life, results in its calcification. The narrow tubes originally occupied by
the trichomes become obscured, possibly due to infilling by micritic cement.
Occurrence: As individual~ and also as massive dendrolites in association with

archaeocyaths and sponges in Cambrian and Lower Ordovician reefs (see
references in Riding and Voronova 1982a). Angulocellularia-like dendritic
clusters are also present in cryptic environments in some Cretaceous shallow
marine carbonate sediments (e.g. see crusts inside dasycladalean algae figured
by Barattolo 1983, Plate 5), and it is likely that more will be reported that have
hitherto been overlooked as being simply irregular micritic crusts. Frutexites
Maslov is an iron-impregnated fossil similar to Angulocellularia (see Riding and
Voronova 1982a, p. 108). Frutexites was originally described from the Or-
dovician (Maslov 1960, pp. 60-61) and is also known from the Devonian
(Playford et al. 1976) and Jurassic (Szulczewski 1963). A Proterozoic example
(Walter and Awramik 1979) is silicified. The only Recent report of An-
gulocellularia is the freshwater lacustrine oncoid occurrence described by
Riding and Voronova (l982a).
3.2 Epiphyton Group
Features: Micritic, narrow dendritic filaments which may be rod-like, appa-

rently chambered, or tubular; erect or pendant growth.
Examples: Epiphyton Bornemann, Gordo n ophyton Korde, Tubomorphophyton

Korde.
66 R. Riding
Affinities: Uncertain, due to the lack of recognized Recent analogues; but the
size and morphology are consistent with a cyanobacterial affinity, as Pia (1927,
p. 39) first proposed. A more specific, stigonematalean, affinity has been
proposed for Epiphyton (Luchinina 1975, p. 21) and Tubomorphophyton (Riding
and Voronova 1982b). The view ofKorde (1959) that Epiphyton is a red alga has
not received support in recent years. For a discussion of these problems see
Riding and Voronova (l982b). Pratt (1984) suggested that Epiphyton is a
diagenetically calcified coccoid cyanobacterium.
Occurrence: Epiphyton group genera first appear close to the Precambrian-

Cambrian boundary and are common and widespread in Cambrian shallow
marine reeflimestones. They are closely associa ted with archaeocyaths in Lower
Cambrian reefs, but they also occur as massive dendrolites in their own right.
The group reappears prominently, but briefly, in the Upper Devonian in
stromatoporoid reefs (Fig. 7), although the genera are usually smaller, and
probably different, than in the Cambrian. Epiphyton has been reported in the
Lower Cretaceous (Masse 1979, Plate 1, Fig. 5) but the bush-like dendritic fossil
in this latter occurrence more closely resembles Angulocellularia.
3.3 Girvanella Group
Features : Thin-walled tubes, tangled, coiled or aligned ; prostrate growth.
Examples: Girvanella Nicholson and Etheridge, Obruchevella Reitlinger,

Subtifioria Maslov.
Fig. 7. Epiphyton Bornemann. Upper

Devonian. Mount Haultain, Alberta. x125.
(Specimen courtesy of E. W. Mountjoy)
Affinities: Recent calcified Plectonema (Oscillatoriaceae) closely resembles

Girvanella (Riding 1977b), and it is likely that other Girvanella-group fossils may
also be calcified oscillatoriacean sheaths.
Occurrence: Girvanella-group fossils typically occur as individuals or in small

clusters (Fig. 8). Girvanella, which is one of the most widely known calcified
cyanobacteria in the Palaeozoic (see Danielli 1981), may also occur in oncoids
and in crusts on reef-surfaces. Subtifioria and Obruchevella are only known from
the Cambrian, but Girvanella is present in marine limestones from Cambrian to
Cretaceous, as well as being present in some Recent freshwater environments.
3.4 Hedstroemia Group
Features: Branching clusters of tubes, clusters with fan-like longitudinal sec-

tion; radial erect growth. Adjacent tubes may share common walls or may be
separate.
Examples: Hedstroemia Rothpletz, Botomaella Korde, Cayeuxia Frollo, Or-

tonella Garwood.
Affinities: Hedstroemia, Botomaella and Cayeuxia closely resemble Recent

calcified rivulariacean cyanobacteria. Ortonella resembles some species of
Scytonema. However, comparisons have also been made with codiacean
(udoteacean) green algae (see Sect. 3.6 below).
Fig. 8. Girvanella Nicholson and Etheridge.

Caradoc (Ordovician). Aldon's Quarry, near
Girvan, Scotland, xl15
68 R. Riding
Occurrence: Hedstroemia group fossils are common in marine environments

from the Cambrian (Botomaella, Bija Vologdin), through the Silurian
(Hedstroemia) (Fig. 9) and Carboniferous (Ortonella), to the Jurassic and Lower
Cretaceous (Cayeuxia) , as well as being represented by Recent analogues. In
addition, it is now evident that a number of fossils hitherto included in the
Solenoporaceae should also be placed in the Hedstroemia group.
3.5 Renalcis Group
Features : Clusters of hollow reniform bodies, usually with thick walls which
may be micritic, fibrous or clotted; irregular or erect growth.
Examples : Renalcis Vologdin, Gemma Luchinina, Izhella Antropov (Fig. 10),

Tarthinia Drosdova.
Fig.9. Hedstroemia Rothpletz. Wenlock

(Silurian), Hogklint Formation, Gotland,
Sweden. x46
Fig. 10. Izhella Antropov. Upper Devonian.

Mount Haultain, Alberta. x50. (Specimen
courtesy of E.W. Mountjoy)
WA ll
thlc kn• ••• tructu r.
F IBROUS
PELO IDAL
~ Iz h. lle
MICR ITIC
.Shullurle
~neI CI.
CHAMBER NU MBER ----..
Fig. II . Morphological characters of representative genera in the Renalcis Group, emphasizing

differences in wall structure and variations in wall thickness and chamber number. Also see Riding
and Voronova (1985, Fig. 4 and Table I)
Affinities: Renalcis-group fossils (Fig. II) have been included in the cyano-
bacteria by many authors, on the basis of their general size and morphology,
but in the absence of Recent analogues it is difficult to systematically place them
with confidence. Hofmann (1975) suggested that some Renalcis-group fossils
represent calcification of g~latinous colonies of chroococcalean cyanobacteria.
He interpreted the thick, dark walls in Renalais as the outer, deeply pigmented
portions of the colonies, and the chambers as the lighter inner portions
(Hofmann 1975, p. 1131; also see Poncet 1976 and Pratt 1984). Izhella, common
in the Devonian (where it has often been confused with Renalcis, e.g. R. turbitus
Wray belongs with Izhella) is characterized by V-shaped clefts in the inner side
of the thick micritic wall, which resemble desiccation cracks in calcified sheath.
Gemma (see Fig. 4) also fits this type of interpretation of the presence ofa thick
sheath. Its wall contains peloidal clots of dense micrite, but the clots appear too
large to represent individual cells. Luchinina (in Zhuravleva et at. 1982)
compared Gemma in general size and morphology with Chlorogloea Wille.
Tarthinia appears to have a fibrous wall and is thus quite different in structure
to both Renalcis and Gemma, although it also possesses the external botryoidal
form. The apparently fibrous wall of Tarthinia could suggest a filamentous
organization, and the external similarities of these genera could be a misleading
guide to their precise affinities within the cyanobacteria. Shuguria Antropov,
which together with I zhella is common in the Devonian, has also been confused
with Renalcis (e.g. R. devonicus Wray belongs with Shuguria). Riding and
Brasier (1975) compared Shuguria (misidentified as Renalcis) with foraminifers
(see also Loeblich and Tappan 1964).
Occurrence: Renalcis-group fossils are equally as common as those of the

Epiphyton-group and share a simila r stratigraphic distribution: they first appear
70 R. Riding
close to the Precambrian-Cambrian boundary, are common through to the

Lower Ordovician, and reappear in the Devonian. In addition, Renalcis-group
genera occur rarely in the Silurian (Riding and Watts 1983) and also are present
in the Lower Carboniferous (Vachard 1974; Bless et al. 1976, p. 138). Renalcis
is most common in the Lower Cambrian, and Gemma and Tarthinia are only
known from the Lower Cambrian. Izhella and Shuguria are widespread in the
Upper Devonian. The overall similarity of geological distribution of the Ren-
alcis and Epiphyton groups emphasizes the way in which these fossils fluctuate
in abundance through time; a feature which suggests that temporal variations
in sea-water chemistry may have acted to periodically enhance marine
calcification in cyanobacteria (Riding 1982).
Tarthinia and Renalcis are known to be present in Lower Cambrian
thrombolites (Latham and Riding 1988) but Renalcis-group fossils generally are
most widely seen in reef-associations with archaeocyaths and stromatoporoids,
as well as with Epiphyton-group fossils in dendrolites.
3.5.1 Microproblematica
The following microfossils are commonly regarded as possessing a cyanobac-

terial affinity, but this attribution is not regarded here as being so strong as it is
for the preceding groups. Indeed, some of the following are not, in my opinion,
cyanobacterial, but they have so often been suggested to be so that they are
included here for completeness. Features and examples given for some groups
follow Chuvashov and Riding (1984, Table 1).
3.6 Garwoodia/Mitcheldeania Group
Features: Coarse, thin-walled, radiating to tangles clusters of tubes which are

commonly juxtaposed.
Examples: Garwoodia Wood/ Mitcheldeania Nicholson
A ffinities: These fossils resemble some of those of the H edstroemia group, but
the tubes are larger and their size makes it difficult to consider them as
cyanobacteria. It has been suggested that they are codiacean (i.e. udoteacean),
an attribution also made for several members of the Hedstroemia group
(Johnson 1961, pp. 97-98). This view has been developed, for example, by
Mamet and Roux (1975), Dragastan (1985,1988), and Mamet and Preat (1986).
Possibly it originates in Rothpletz's (1891) suggestion that Girvanella may be a
codiacean (see Riding 1975, p. 174). Calcified halimedaceans, such as Halimeda,
which Johnson (1961) placed together with Garwoodia and Mitcheldeania, are
much larger and morphologically quite different from the latter. Thus, the main
support for the view that some coarsely tubed porostromates may be udo-
teaceans appears to come only from the superficially similar organization of
extant uncalcified green algal genera such as Codium Stackhouse. However, it

must be admitted that modern analogues have also not yet been found for many
of these fossils in calcified cyanobacteria either.
At present it seems best to place Garwoodial Mitcheldeania and allies in a
separate group to await clarification of their systematic position. The question
of whether Garwoodia and Mitcheldeania are synonymous or are separate
genera is a minor but intricate problem stemming from the history of the
erection of these taxa which remains to be clarified.
3.7 Rothpletzella Group
Features : Flat, curved or encrusting sheets of juxtaposed tubes which branch

dichothomously in one plane.
Examples: Rothpletzella Wood (Fig. 12), Flabellia Shuysky.
Affinities: Wood (1948, pp. 17-18) regarded Sphaerocodium gotlandicum,

which he named Rothpletzella gotlandica, as a probable alga and commented:
"If this form is an alga the habit, and especially the absence of septation of the
filaments, would seem to indicate that it belongs to the Myxophyceae
(Cyanophyceae) or possibly to the group Siphonales of the Chlorophyceae."
However, his reasons for considering an algal affinity were not compelling:
"Their walls are composed of "algal dust", their threads branch dichotomously,
and their diameter is below that which would accommodate any multicellular
animal. Algae of Girvanella type are commonly found in association with them,
sometimes building composite nodules, so that the conditions were such that
algae could flourish" (Wood 1948, pp. 17-18). Although these arguments are
inconclusive, Rothpletzella has generally been considered to be a cyanobacte-
rium. However, Wood's effort to clarify the taxonomy of Sphaerocodium
Rothpletz has been less successful. Many subsequent references to the fossil
which he named Rothpletzella have called it Sphaerocodium, mainly due to
Fig. 12. Rothpletzella Wood. Wenlock (Silu-

rian) , Wenlock Limestone, May Hill, Glou-
cestershire, England. x 115
72 R. Riding
Wray's (1967, pp. 36- 37) rejection of Wood's decision, and this point still
requires clarification.
Rothpletzella is common as reefal crusts, and also forming oncoids and
stromatolites, in the middle Palaeozoic (Chuvashov and Riding 1984, Figs. 3, 4).
A notable occurrence is in the construction of fore-reef stromatolites in the
Upper Devonian Canning Basin of Western Australia (Playford et al. 1976,
p. 557, Fig. lib).
3.8 Wetheredella Group
Features: Short encrusting tubes, possibly branched, with hemispherical or

rounded cross-sections; fibrous wall structure sometimes with additional
micritic layer.
Examples: Wetheredella Wood (Fig. 13), Aphralysia Garwood, Asphaltina

Mamet, Sphaeroporella Antropov.
Affinities: In his revision of Silurian "Sphaerocodium", Wood (1948) recog-

nized not only Rothpletzella (see Rothpletzella Group, above) but also Weth-
eredella. He argued that the latter could be a foraminifer, and noted its
resemblance to Nubecularia Defrance (Wood 1948, pp. 18 and 20-21). Loeblich
and Ta ppan (1964, p. C787) suggested the reverse: "doubtfully a foraminifer,
probably algal". Copper's (1976) attribution of a cyanobacterial affinity to
Wetheredella was not accepted (Riding 1977c), but Mamet and Roux (1975,
pp. 156- 170) tentatively referred Aphralysia, Sphaeroporella, Asphaltina,
Wetheredella , and other similar genera to the green algae. Ischenko and
Radionova (1981) went further, and drew comparisons between Wetheredella
Fig. 13. Wetheredella Wood . Wenlock

(Silurian), Wenlock Limestone, May Hill,
Gloucestershire, England. xl15
and Rhabdoporella Stolley (a dasycladalean, according to Pia, 1927, p. 62), and

suggested a transitional series from these genera to Rothpletzella. These inter-
esting observations deepen the systematic problems surrounding these fossils.
3.9 Tubiphytes Group
Features: Erect, irregular, or encrusting skeletons with a smooth exterior and

dense, dark, flocculent internal structure showing concentric bands; central
tubes often present.
Example: Tubiphytes Maslov (Fig. 14).
Affinities: Maslov (1956, p. 82) placed Tubiphytes in Algae Incertae Sed is, with
the suggestion that it might belong to the Schizophyta (cyanobacteria), but
hydrozoan (for details see Babcock 1977, pp. 17-18) and sponge (Ott in Kraus
and Ott 1968, p. 271) affinities have also been proposed. Tubiphytes ranges from
Carboniferous to Cretaceous, and is especially important in reefal environments
during the Permian.
3.10 Lithocodium/Bacinella Group
Features: Encrusting masses of irregular open mlcnttc networks usually

becoming more orderly externally where a thicker surface layer may be pierced
by numerous pores.
Examples: Lithocodium Elliott, Bacinella Radoicic.
Fig. 14. Tubiphytes Maslov. Sakmarian

(Early Permian), Trogkofel Formation, Trog-
kofel, Carnic Alps, Austria. x16. (Specimen
courtesy of David Edwards)
74 R. Riding
Affinities: These genera are often seen juxtaposed and may well represent the
same organism. Elliott (1956) regarded Lithocodium as a codiacean (i.e. udo-
teacean) , but it has been suggested that these irregular meshwork-like crusts and
masses may be microbial in origin (Maurin et al. 1985; Camoin and Maurin
1988). However, similarities also exist with foraminifers, sponges, and hy-
drozoans. Lithocodiuml Bacinella ranges from Upper Permian to Cretaceous
and is particularly common in shallow marine environments in the upper
Jurassic and the Cretaceous.
4 Geological History
Cyanobacteria have the longest well-defined fossil record of all major groups of
organisms. Only bacteria may have a longer record and their remains are at
present much less well-known. Three principal periods can be recognized in the
geological history of cyanobacteria, on the basis of the development of
calcification in marine environments, with the middle period defined by the
common presence of calcified cyanobacteria in shallow marine carbonate
environments from the Early Cambrian (ca. 590 million years) to the late
Mesozoic (ca . 65 million years; Fig. 15). Both before this, during most of the
Precambrian, and afterwards, during the Cenozoic, there is no clear evidence of
common cyanobacterial calcification in the sea.
CENOZOIC o
85
MESOZOIC 590 to 65
248 } heavily calcified
cyanobacteria
PALAEOZOIC common
590
(.)
o 2000 to 500
stromatolites
N
o common
II:
W
t-
O
II:
a..
2500
2800, earliest
cyanobacteria-like
z fossils
0(
w Fig. 15. Distribution of calcified
:z:
(.)
3500
cyanobacteria and stromatolites in
II: earliest marine environments through
0( stromatolites geological time. Scale in millions
of years from Harland et al. (1982)
4.1 Precambrian
The earliest known occurrence of cyanobacteria-like fossils is in the 2800-

million-year-old Fortescue Group, Western Australia (Schopf and Walter
1983; Fig. 15). These are present in cherty stromatolitic limestones and also
occur in similar younger deposits such as the well-known Gunflint Chert
Formation (ca. 2000 million years) of Ontario (Tyler and Barghoorn 1954) and
Bitter Springs Formation (ca. 900 million years) of Australia (Schopf 1968).
Precambrian stromatolites are associated with extensive, apparently marine,
dolomite and limestone deposits in the Proterozoic (ca. 2500-550 million years)
and reach a peak of diversity and importance near 800 million years (Awramik
1971) which has never since been equalled. Nevertheless, there is considerable
uncertainty concerning their processes of formation. They are obviously cal-
careous deposits, but it is important to know whether they formed by precipi-
tation of calcium carbonate on or within microorganisms, or by trapping and
binding of particulate sediment, or by a combination of these processes.
Gebelein (1976, p. 505-506) has suggested that precipitation is the principal
process involved, on the basis of (I) lack of preserved detrital carbonate in most
Precambrian stromatolites; and (2) the presence offorms with more than 10 cm
of relief which would, it is argued, require cementation for support. However,
despite intensive examination of Proterozoic stromatolites by many workers,
few tubiform or bush-like microfabrics characteristic of calcified filamentous
cyanobacteria have been found, other than poor coarse 'vermiform' micro-
structures (Bertrand-Sarfati 1976, p. 255), although there is now evidence of
better preserved "tussocky" fabrics resembling calcified rivulariaceans from the
latest Proterozoic (Bertrand-Sarfati and Pentecost 1989). Consequently, it has
been suggested that precipitation of calcium carbonate on or within cyano-
bacterial sheath material is only a minor aspect of stromatolite formation
before the late Precambrian (Riding and Voronova 1982c). This in turn implies
that trapping and binding of particulate sediment was the principal process
involved.
It is important to check this apparent absence of calcified cyanobacteria in
the Precambrian. It may simply be due to lack of detailed searching. However,
if it can be established, it will then imply either an inability of sheaths at that time
to provide suitable crystal nucleation sites, or lower CaC03 precipitation rates
relative to the Palaeozoic (see Discussion, below), or both.
4.2 Cambrian-Cretaceous
4.2.1 Cambrian-Lower Ordovician
Heavily calcified cyanobacteria appear dramatically near the Precambrian-

Cambrian boundary (Riding and Voronova 1984). The presence of facies
change from restricted to open marine conditions at this level in key Siberian
sections containing carbonate rocks makes it possible that the sudden appear-
76 R. Riding
ance of calcified cyanobacteria (and other skeletal fossils too, for that matter)
is artificially accentuated. Nevertheless, the overall change from the Vendian to
the Cambrian is striking. Re-evaluation of the Cambrian marine calcified flora
suggests that most Lower Cambrian "algae" are probably cyanobacterial
(Riding, this VoL). Certainly the following important genera are cyanobacteria:
Angulocellularia, Botomaella, Girvanella, Hedstroemia, and Subtiflora. These
occur as individuals, dendrolites, reefal crusts and as oncoids. Even more
common than these are members of the Epiphyton and Renalcis groups which
are prominent in dendrolites and reefs. They may well be cyanobacteria,
although their precise affinities are still unclear.
The abundance and diversity of these fossils continues virtually un-
diminished into the Lower Ordovician, where sponge-cyanobacterial reef as-
sociations (Riding and Toomey 1972) are very similar to those of archaeocyaths
and cyanobacteria in the Lower Cambrian. This Early Palaeozoic phase
represents not only the beginning, but also the height, of the development of
calcified cyanobacteria in the Phanerozoic. At this time cyanobacteria are as
conspicuous in reef formation as are sponges and metazoans.
4.2.2 Middle Ordovician to Permian
The Early Palaeozoic importance of calcified cyanobacteria was not sustained

through the middle Palaeozoic, but various groups reappear sporadically in the
Silurian and Devonian. Hedstroemia, in association with probable cyanobac-
teria previously assigned to Solenopora, constructs thin but massive crests on
Lower Silurian patch reefs in Gotland (Riding and Watts 1981). Epiphyton and
Renalcis-group genera reappear abundantly in Upper Devonian reefs (Wray
1967), although they differ in detail from those present in the Cambrian. Both
groups disappear by the end of the Lower Carboniferous.
Girvanella is common as individual masses, reefal crusts, and in oncoids
throughout the middle Palaeozoic, but other prostrate tubiform genera, such as
Subtifloria are not known from post-Cambian rocks. The type specimen of
Girvanella, from the Ordovician of south-west Scotland, occurs as very small
tangled masses which may have been semi-planktonic rather than benthic.
Recent Plectonema which is analogous with Girvanella also occurs as floating
clusters of filaments (Riding 1977b, p. 36).
Microproblematica which have been attributed to the cyanobacteria, such
as Rothpletzella and Wetheredella, are common throughout the middle
Palaeozoic as reefal crusts and in oncoids (Chuvashov and Riding 1984,
Figs. 3, 5).
During the Carboniferous Ortonella, Bevocastria and Garwoodial
Mitcheldeania are locally important in oncoid and stromatolite formation
in marginal marine environments (Garwood 1931) but generally the Carbon-
iferous is a period of decline in the importance of calcified marine
cyano bacteria.
There is no clear evidence to indicate that this situation alters significantly

in the Permian, but assessment of the importance of cyanobacteria is com-
plicated by the inclusion of micro problematic a, notably Tubiphytes Maslov, in
this group. Tubiphytes is common in Permian reefs, both as an encrusting
organism and as a primary framebuilder. Whatever the nature of Tubiphytes
(see Microproblematica, above) it does not seem to resemble any extant calcified
cyanobacteria, nor does it appear to be algal.
Hedstroemia is present in the upper part of the Capitan reef complex (lower
Upper Permian) ofthe south-western United States, and Girvanella is present in
the Lower Permian of the Carnic Alps together with occasional Ortonella and
Garwoodia (Table 2 in Flugel 1977), but in general calcified cyanobacteria are
not conspicuous in the Permian.
4.2.3 Triassic to Mid-Cretaceous
Calcified cyanobacteria become much more prominent in reef and back-

reef/lagoon facies in the Middle and Upper Triassic (Flugel 1975), and this
situation continues until the mid-Cretaceous. In the Alps, the Dolomites and in
Greece, Middle to Late Triassic sponge and coral reefs have crusts and bushy
growths of Bacinella, Bevocastria, Cladogirvanella, Garwoodial Mitcheldeania,
Girvanella, Hedstroemia, and Ortonella, while nodules formed by Apophoretella
Elliott, Cayeuxia, Polytrichella Schafer and Senowbari-Daryan, and Zonotri-
chites are common in lagoonal environments (Fois and Gaetani 1980; Schafer
and Senowbari-Daryan 1983; Fois 1981). Tubiphytes is also present in Triassic
and some Jurassic reefs.
Cayeuxia and its relatives continue to be common through the Jurassic
(Flugel1979) and into the Cretaceous. Dragastan (1985) has correctly compared
these with Rivularia, although it would probably be best to retain the names
Cayeuxia, Zonotrichites, etc. to distinguish these calcified skeletons from extant
cyanobacteria whose diagnoses are based on details of soft organic tissues and
not on hard parts.
Micritic crusts, possibly of cyanobacterial origin, are present in Middle
Triassic (Fois and Gaetani 1984) and Middle Jurassic (Gwinner 1976) sponge
reefs. More dendritic, but also micritic, crusts resembling Cambrian An-
gulocellularia occur in cavities in Lower Cretaceous grainstones (see Masse
1979).
The problematic porous micritic crusts known as Lithocodium Elliott and
Bacinella RadioCic are locally important in reefs from the mid-Triassic to Lower
Cretaceous. They have not generally been regarded as being cyanobacterial,
and they do not resemble extant calcified cyanobacteria, although a microbial
origin as well as various other suggestions have been made concerning their
affinity (see Microproblematica, above).
78 R. Riding
4.3 Late Mesozoic and Cenozoic Decline
The decline in calcified marine cyanobacteria probably began in the Upper

Cretaceous. By the early Cenozoic there were few or none present in the shallow
car.bonate environments in which they had hitherto been conspicuous during
most periods since the early Palaeozoic, and this scarcity continues to the present
day. The few reports of calcified cyanobacteria, as opposed to those which trap
and bind sediment, in Recent shallow marine carbonate environments include
(1) calcified sheath material associated with partly cemented ooids from Great
Bahama Bank (Winland and Matthews 1974, Fig. 3c); (2) a circular structure,
which may be a cross-section of a similar filament from Red Sea reefs (Friedman
et al. 1974, Fig. 15); and (3) Rivularia mesenterica and R. polyotis with rounded
carbonate inclusions from the Bahamas, Adriatic Sea, and Shark Bay (Golubic
and Campbell 1981). The latter Rivularia specimens, however, are from in-
tertidallocalities which do not appear to ha ve normal marine salinities. Brackish
water Rivularia has been reported from the Pleistocene of Greece (Richter et al.
1979). However, calcified cyanobacteria continue to be common in freshwater
carbonate environments in the Recent where they are present in lacustrine and
fluviatile oncoids, travertines and freshwater marshes.
5 Discussion
5.1 Analogues
Recent analogues are of crucial importance in establishing the identities offossil

calcified cyanobacteria. Despite the fact that calcified cyanobacteria are now
virtually absent in marine environments, they are common in freshwater
calcareous lakes and streams and, provided that they are not too heavily
encrusted (as in tufa deposits), they are convincing analogues for Palaeozoic and
Mesozoic marine fossils. Consequently, examination of Recent calcified
freshwater cyanobacteria has confirmed that fossils such as Girvanella Riding
(l977b), Angulocellularia (Riding and Voronova 1982a), and Hedstroemia are
calcified sheaths of filamentous cyanobacteria. Further comparative studies are
needed to extend this systematic work.
One limitation of this approach which can be anticipated is that a variety
of Recent taxa may produce similar sheaths which, when calcified, are so alike
that they come to constitute a single fossil taxon. This is almost certainly true
in some cases, but, surprisingly, there are examples which also show the
opposite. Hedstroemia and Botomaella, for example, are distinct fossil genera,
but they both possess analogues in extant calcified rivulariacean sheaths
(Riding, unpubl.). These sheaths thus exhibit greater morphological varia-
tion than do the trichomes which they enclose and by which they are
produced. And these differences in sheath character are preserved by calci-
fication. As a result, a Recent taxon may represent several fossil taxa, rather
than the reverse.
Variability in sheath character is the main feature reflected in fossil cyano-

bacteria. Ironically, culturing of these organisms in an attempt by workers on
modern cyanobacteria to stabilize and simplify taxonomy (e.g. Rippka et al.
1979) often leads to absence of the sheath altogether. As a result there is a
tendency by these workers to undervalue sheath characters in taxonomy which
may make it more difficult to relate modern cyanobacteria to fossil calcified
sheath material. However, it is relatively easy to study sheath variability in
Recent oncoids. The occurrence of the same filamentous cyanobacterium on
both the upper and lower surfaces of a stabilized oncoid results in a gradient of
sheath morphologies. On the upper surfaces filaments grow erect, elongate, and
in open tufts, whereas on the lower surface they are compressed and compact
due to more constrained growth. This results in a transitional series of sheath
morphotypes ranged between two end-members, one open and elongate, the
other dense and compact; but all apparently originating from the same species
of cyanobacterium. This information is valuable in interpreting the mor-
phological series exhibited by cyanobacteria, for example some of those noted
by Riding and Voronova (1985).
5.2 Timing of Calcification
Study of Recent analogues indicates that cyanobacterial calcification begins

during life (Pentecost and Riding 1986). In the case of filamentous forms this
does not appear to inhibit growth because the ends ofthe filaments remain open.
There is less information concerning coccoid forms but calcification will not
necessarily prevent diffusion of water or light through the enveloping sheath.
Post-mortem calcification has been reported in association with the desiccation
of En tophysa lis mats at Laguna Mormona (Horodyski and Von der Haar 1975)
and Shark Bay, but the general result is that "mineral incorporation obliterates
the biological structure" (Golubic 1983, p. 313). However, these examples are
from evaporative intertidal environments. Post-mortem calcification in subtidal
environments has been reported in the green alga Ostreobium (Lukas 1974;
Kobluk and Risk 1977). Calcification takes place within 2-3 months and
precipitation of micritic calcium carbonate, on both the interior and exterior of
the filaments, preserves the shape of the alga. Similar effects in cyanobacteria
have not been reported, but could contribute to their preservation. Pratt and
James (1982) and Pratt (1984) have suggested that diagenetic calcification of
coccoid cyanobacteria is the main process by which Renalcis and Epiphyton
group microfossils are preserved, and this possibility requires further study.
5.3 Skeletons
The suggestion that stromatolites formed by calcified fossils should be termed

skeletal stromatolites (Riding 1977a) has from time-to-time aroused criticism. It
has been suggested that cyanobacteria do not form skeletons (Monty 1981, p. 4)
80 R.Riding
although Golubic and Campbell (1981, pp. 223-225) described calcification in

Rivularia and came to the conclusion that: "with respect to biological control
over calcification and specific morphology of the resulting concretions,
Rivularia is comparable to Penicillus and Rhipocephalus, and shows more
organismic control than the intercellular aragonite precipitation of Halimeda".
But they did go on to say that "this criterion, however, is not fulfilled in the case
ofthe nonspecific micrite that surrounds the "filaments" of Girvanella (Golubic
and Campbell 1981, p. 225).
Nevertheless, a skeleton or a skeletal element can be regarded as a hard
mineralized layer or component formed on or within an organism during its life.
Cyanobacterial calcification takes place within or on the mucilaginous sheaths
during life. It results in a hard-part which will have offered protection and
support to the organism and which reflects some aspects of its morphology. It is
possible to argue backwards and forwards about the degree of control involved
and whether or not this kind of hard part is comparable with that of algae,
invertebrates and vertebrates. In these circumstances it seems necessary to
accept that there are different types of skeleton related to different biominer-
alization processes. Those who oppose the use of the term skeleton for calcified
cyanobacteria are likely to encounter problems in attempting to distinguish
these hard parts from those of algae such as Halimeda, and probably also from
those of invertebrates such as some foraminifers. It is unlikely that all organic
hard parts are formed in the same way or for the same purpose, but until we have
a system of clearly discriminating between them then it seems reasonable to
refer to them all as skeletons or skeletal elements.
5.4 Seawater Chemistry
The pattern of cyanobacterial calcification in marine environments through

time shows variation on two scales. In the long term, it appears that heavily
calcified cyanobacteria are only present during the Palaeozoic and Mesozoic.
Within this long period of time they are most common during the Cambrian-
Lower Ordovician, Upper Devonian-Lower Carboniferous, and Middle
Triassic-mid-Cretaceous.
Monty (1973, p. 615) has suggested that the lack of calcification in Recent
subtidal marine stromatolites could be due to some unspecified change in
sea-water composition together with a response, on the part of cyanobacteria, to
ecological pressure from calcareous red algae which has led cyanobacteria to
utilize sediment trapping and binding to occupy niches in which direct com-
petition with rhodophytes is avoided. However, Gebelein (1976, p. 512) pointed
out that there is no clear evidence for general competition of this sort between
cyanobacteria and rhodophytes, although he agreed that red algae may limit the
development of stromatolites in reef communities (Gebelein 1976, p. 514).
He went on to develop the idea ofa change in sea-water chemistry by suggesting
that "rapid rise of the calcareous plankton during the Mesozoic ... may have
adjusted the carbonate equilibria so as to make it virtually impossible for
blue-green algae to precipitate carbonate in normal marine waters" (Gebelein

1976, p. 514). With regard to changes near the Precambrian-Cambrian bound-
ary, Monty (1973) and Gebelein (1976) both took the view (questioned here)
that Precambrian stromatolite-building biotas lost the ability to precipitate
carbonate. Monty suggested that this was due to extinction of carbonate
precipitating cyanobacteria, linked to decrease in CO 2 pressure: "such a
decrease would have affected the rates of photosynthesis as well as the abilities
of blue-green algae to precipitate carbonate" (Monty 1973, p. 605). Gebelin
(1976, p. 506) postulated a "lowering of the carbonate super-saturation due to
removal of bicarbonate ion by metaphytes and metazoans".
This view of a two-stage reduction in ability to calcify in marine environ-
ments, first at the end of the Precambrian and then near the end of the Mesozoic
is inconsistent with the appearance of heavily calcified cyanobacteria in the
early Cambrian. The overall, long-term decline in stromatolite abundance and
diversity from the late Precambrian to the present is plausibly explained by the
hypothesis of direct and indirect competition from metazoans (Garrett 1970)
and changes in sedimentation (Pratt 1982). But changes in calcification require
a different explanation. Monty (1973) and Gebelein (1976) invoked factors such
as changes in CO 2 pressure and carbonate equilibria at both the Precam-
brian-Cambrian and the Mesozoic-Cenozoic boundaries to explain what was
perceived at both boundaries to be a loss of calcification in cyanobacteria.
Instead, it is possible to propose changes in the Mg2+ :Ca 2 + ratio of seawater
and/or in atmosphere-hydrosphere pC02 near the Precambrian-Cambrian
boundary stimulating (rather than inhibiting) calcification, to account for the
Palaeozoic-Mesozoic occurrence of calcified marine cyanobacteria, followed by
a reversed change in Mg2 +·:Ca 2+ and/or pC0 2 near the Cretaceous-Tertiary
boundary to account for the disappearance of calcification in Cenozoic marine
cyanobacteria (Riding 1982). The Cretaceous-Tertiary decline coincides with
the change from calcitic to aragonitic ooids reported by Sandberg (1983).
Fluctuations in abundance of calcified cyanobacteria through the Palaeozoic-
Mesozoic show some broad similarities with the aragonite-facilitating and
calcite-facilitating events recognized by Sandberg (with marine cyanobacterial
calcification being most conspicuous during calcite-facilitating periods, i.e.
Cambrian to Devonian and Triassic to mid-Cretaceous). However, the most
generally held view of Proterozoic seawater chemistry is that it would have
readily facilitated carbonate precipitation (Holland 1984, p. 422; Grotzinger
1989). Consequently, it is puzzling that calcified cyanobacteria have not been
more widely reported from Proterozoic carbonates (see Geological History,
Precambrian, above). Some of the assum ptions tha t ha ve been made concerning
Precambrian life and environments may need to be reexamined in this light. It
is possible that the sudden appearance of diverse calcified cyanobacteria in the
earliest Cambrian reflects an evolutionary event for prokaryotes which is
comparable with that observed in metazoans at this time. However, the sub-
sequent history of marine cyanobacterial evolution is overprinted by temporal
variation in carbonate precipitation rates, as is well seen in the long-term
fluctuations in abundance of calcified cyanobacteria during the Phanerozoic.
82 R. Riding
6 Conclusions
I. With the aid of Recent analogues from freshwater environments it is possible

to clarify the affinities of calcareous microfossils which have been suggested to
be cyanobacteria. This shows that the principal filamentous groups involved are
oscillatoriaceans and rivulariaceans, and that the site of calcification is the
protective extracellular mucilaginous sheath surrounding the cells.
2. The main fossil groups cluster around the following typical genera:
Angulocellularia, micritic; bush-like; in the Recent formed by Shizothrix.
Epiphyton, micritic, tubular, or chambered; dendritic; bearing a resemb-
lance to stigonemataleans.
Girvanella, tubiform; tangled, coiled, or aligned; in the Recent formed by
Plectonema.
Hedstroemia, branched, radial tubes; in the Recent formed by
rivulariaceans.
Renalcis, botryoidal clusters of thick-walled chambers; may be calcified
coccoid colonies.
In addition, the Garwoodial Mitcheldeania and Rothpletzella groups have
also commonly been regarded as cyanobacterial, but this is here considered to
be unlikely.
3. In the Recent, calcified cyanobacteria occur in lacustrine and fluvial envi-

ronments as discrete skeletons and tufa oncoids and as tufa stromatolites. But in
ancient marine environments they are most conspicuous as dendrolites,
thrombolites and skeletal stromatolites.
Dendrolites and thrombolites are particularly abundant in the Cambrian
and Lower Ordovician. In contrast, the Proterozoic shows little evidence of
well-calcified cyanobacteria despite the abundance and diversity of stroma-
tolites. Thus, the appearance of calcified rock-forming cyanobacteria near the
Precambrian-Cambrian boundary was an event immediately followed by the
acme of these organisms in the marine environment. Their subsequent Phan-
erozoic history is one of overall decline in importance, albeit interspersed by
periods of relative abundance in the Upper Devonian to Lower Carboniferous
and Middle Triassic to mid-Cretaceous. During the Cenozoic calcified cyano-
bacteria appear to be virtually absent from the sea. The most likely explanation
for this pattern of appearance and disappearance of calcified cyanobacteria in
marine environments through time is that prokaryote evolutionary and ex-
tinction events are superimposed on fluctuations in environmentally deter-
mined calcification (Riding 1989).
Acknowledgements. It is a pleasure to thank Ernst Ott. who encouraged my interest in modern

calcified cyanobacteria both before and during tenure of an Alexander von Humboldt-Stiftung
research fellowship at the Technical University. Munich. Additional support has been given by the
Natural Environment Research Council. I am grateful to Allan Pentecost and Brian Pratt for
providing helpful comments on the manuscript.
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Chapter 4
The Solenoporaceae: A General Point of View
A.F. POIGNANT l
Abstract
The classification of the Solenoporaceae is very difficult to establish, as the criteria which may be
used are scarce and not very reliable. There is a great deal of confusion surrounding generic and
specific allocation and classification systems. Even today, it is still not possible, e.g. between green
algae and red algae, for one to determine a definite assignment for this group.
There are only a few generic or specific determination tests, and even then they seem far from
convincing. The outer structure seems to alter very slightly in relation to time; the inner structure is
more complex: hypothallus (?), perithallus, partitions, walls, pores, and reproductive organs (?) can
be examined.
Fifty years ago, numerous phylogenic theories from Solenoporaceae to Corallinaceae were
suggested, but of course, all these phylogenic ideas must be restudied if Mamet and Roux's (1977)
new classification is to be accepted.
As long as a certain taxonomic order is not established, the stratigraphical distribution is
meaningless. Now is the time to reestablish those taxa which make up one distinct family.
1 Introduction
The Solanoporaceae represents an extinct group of marine organisms which has

its climax during the Jurassic. The organisms are nodular or encrusting and are
formed by closely packed threads of vertically divergent cells. Hypothallus and
reproductive organs are still uncertain; the cells are much longer than those of
the Corallinaceae. The Solenoporaceae were more or less adapted to their
environment and were gradually replaced by the Corallinaceae during the
Cretaceous.
At first, these organisms were classed as animals such as Tabulatae,
Bryozoa, Hydrozoa, and Stromatoporoidae and subsequently as algae due to
resemblance to the Corallinaceae. Pia (1930) was not totally convinced by this
and Vialli (1938) even went as far as to state that it is all a question of a stage
between Chaetetidae and Corallinaceae, or in other words, a transition between
the animal and plant kingdoms. However, Nicholson (1888) stated:
"If evidence can be obtained, proving decisively the existence of a cellular, and
not a tubular structure in So/enopora, then the reference of the genus to the
calcareous algae will follow as a matter of course."
IUniversite Pierre et Marie Curie, Centre d'Etude des Algues Fossiles, Boite 2000-4, Place lussieu,
75252 Paris Cedex 05, France
The Solenoporaceae: A General Point of View 89
Several arguments have been put forward to confirm that these forms
belong to algae: organs claimed to be reproductive, cell size, the presence of
partitions in tubes, the preserved violet coloring, etc.
There is a great deal of confusion surrounding generic and specific al-
location and classification systems. This confusion is due to descriptions,
admittedly genuine, which are, however, too brief, too hurried, uncritical and
without size criteria. This confusion is also created by a lack of comparisons of
species, and a lack of photographs, being only partially compensated for by
sketches which serve only as an explanatory illustration of the facts.
2 Historical Background
One must take note of the important work carried out by Deninger (1906),
Dybowsky (1879), and Haug (1883), who created their own genera. Not so
distant in time, we have Elliott (1963, 1965, 1973), Flugel (1960, 1961, 1972),
Garwood (1945), Johnson (1960a,b, 1961, 1963, 1966, 1969), Maslov (1956,
1961), Peterhans (1928, 1929a,b, 1930), Riding (1977), Rothpletz (1908,1913),
Wood (1944), Wray (1967,1970, 19"77a,b), and Yabe (1912), who put forward
criticisms and syntheses.
Certain research, such as that by Brown (1894), whose first study suggests
that the Solenoporaceae were the ancestral form of the Corallinaceae, by Pia
(1930), Wood (1944), and above all by Mamet and Roux (1977), must not be
overlooked. Mamet and Roux (1977), after many observations, question the
classification generally acknowledged since Pia (1939). The reasoning of the
criticism and the simplicity·of their observations appear obvious. It seems that
the works of the first observers, Deninger (1906), Dybowsky (1879), and Haug
(1883), were poorly read and badly in terpreted. There are more than 200 species,
extending from the Cambrian through the Miocene, corresponding to more
than 300 bibliographical references.
3 Detennining Criteria
There are only a few generic or specific determination tests, and even these seem
far from convincing.
3.1 The Outer Structure
The outer morphology hardly changes from one genus to another, and there is
an even smaller change from one species to another. It seems to alter very slightly
in relation to time. In general, we are concerned with nodular masses, rounded
ovoid, hemisperical or conical, encrusting or strongly branching. It should be
noted that the nodular forms seem to be the oldest and have been replaced by
ramified forms during the Mesozoic.
90 A.F. Poignant
3.2 The Inner Structure is More Complex
Several authors have pointed out the presence of two tissues: Hypothallus and
perithallus with a rightful doubt concerning the former.
1. Hypothallus. The presence of an authentic hypothallus has not been proved

for several reasons: (1) its thinness; (2) its shape does not make it possible to
distinguish it from the perithallus; and (3) it is not calcified. Very few authors
draw attention to its presence and then do not give many details pertaining to it.
2. Perithallus. The fairly thick perithallus composes, in fact, the entire tissue. It
consists of closely packed tubes, closed packed in the sense that the cell threads
are in contact with one another with little or no vacant space between, forming
fan-shaped structures.
3. Partitions. The tubes are, more often than not, divided by horizontal par-
titions, almost always concave, delimiting large cells, which are longer than
100 p.m.
The considerable height of the cells is a unique feature which means they
can be classified at the edge of the plant and animal kingdoms, unless the
genus Marinella (Barattolo and del Re 1984) is included in this family (cells
smaller than 10 p.m). Moreover, it must be remembered that Lithoporella
(Corallinaceae) also has large cells, but these are definitely narrower.
The frequency and positioning of the partitions, which appear to be neither
a result of change nor of diagenesis, provide, in my opinion, some good
diagnostic criteria.
In Mamet and Roux's (1977) classification, two genera do not have hor-
izontal partitions, and this is not a question of a lack of original calcification nor
a consequence of diagenesis. In fact, the horizontal partition could be nothing
more than a trail made by longitudinal partitions; thus, two longitudinal
partitions facing each other would make it appear as if there are horizontal
partitions, equivalent to Peterhans' (1928) meanderings or to the indentations
noted by other authors.
4. Walls. The tubes always contain vertical walls, sometimes sinuous, of which
a thorough investigation has never been made. Every tube possesses walls, but
no wall is common to any two tubes. When cross-sections are taken, the tubes can
be seen to be circular or polygonal in shape. This appearance is meaningless in
systematics and it may even change in the same individual.
5. Pores. Perforations have been alluded to by the earliest authors. For some
time, these pores were given a fair amount of importance since they were used
as determining criteria and as arguments to back up classifications. However. no
explanation has been offered, since most authors prefer to deny their presence.
They could possibly be a question of "primary synapses" and/or "secondary
synapses" or could even be the result of diagenetic phenomena in the cases

where original structures are not being considered.
Other puzzling cell shapes have been pointed out, such as those which are
star-like or resemble miliolids. No explanation has been suggested for them.
6. Indentations. Here, we are talking about a special and original feature in the
internal structure of the tubes. This was acknowledged a long time ago but has
since been forgotten. These indentations were noted e.g. by Nicholson (1888),
Rothpletz (1908), Peterhans (l929a,b), and H~eg (1961).
With regard to these, we owe certain pertinent remarks, which totally
change the determining criteria and systematics, to Mamet and Roux (1977).
The first observations on Solenopora made by Dybowsky (1879) show that
in cross-sections, the wall of the cell is corrugated and affected by septiform
processes (up to six), which are directed inwards as if the wall were invaginating.
From this, Johnson (I 960a) saw a branching of the tubes, whereas H0eg (1961)
for his part, saw longitudinal and incomplete septa located above a bifurcation.
Peterhans (1928, 1929a,b, 1930) pointed out, without elaborating further, that
certain cells show indentations. This interpretation has been very uncertain for
a long time.
Nicholson (1888) recognized implicitly the systematic interest provoked by
these indentations:
"no radiating 'septa' are developed, but the type species exhibits more or less
inwardly directed septiform processes."
In regards to Solenopora compacta, more or less numerous septiform

processes, and in Solenopora? filiformis "the tubes do not exhibit inward
septiform processes. I have some doubts about ... this fossil to the genus."
Mamet and Roux (1977) have explained remarkably well what these inden-
tations mean. It is a question of longitudinal, incomplete, and more or less
developed partitions, which extend the length of the tubes and which are
independent from branching. Thus, the study of cross-sections at random can
give the impression of horizontal septa; however, it is rather a question of
longitudinal septa on which, depending on the genus, an authentic horizontal
septation can superimpose itself.
This apparent or real septation is a determining criterion which will playa
role in the classification system.
7. Reproductive Organs. This is a somewhat controversial subject. Hasty ob-

servations have led to uncertain conclusions; the discovery of actual re-
productive organs would greatly simplify the problem of systematic attributions
and phylogenetic problems.
I stipulated in 1977 the suggestion that the Solenoporaceae had sufficiently
large cells for the reproduction of spores to take place without the need of a
favorable, noncalcified external location.
92 A.F. Poignant
A claim favoring the existence of reproductive organs in Solenoporaceans

was made for the first time by Brown (1894) and then by Rothpletz (1913),
Garwood (1945), Pia (1930, 1940), Opik and Thomson (1933), Peterhans
(I 929b ), Maslov (1956), and Elliott (1965, 1973). Elliott's (1965, 1973) examples
cannot all be accepted: N eosolenopora patrinii was proved to be a bryozoan and
Solenopora sardoa an anthozoan.
There are certain authors who disagree with this theory and some who have
changed their standpoint. On the whole, authors remain doubtful:
Wray (l977a): "unequivocal reproductive organs have not been found In
Paleozoic species."
Elliott (1965): "definite reproductive structures are usually missing."
Garwood and Goodyear (1919): "occasionally small elonga ted fusiform spaces
occur in the thallus, filled with crystalline calcite. Similar spaces have been
described by Rothpletz (1913) as sporangia. They suggested rather due to the
irregular solution of partitions of cell walls."
Garwood (1913): "a case where this is a definite association of these
(pseudosporangia) bodies with growth lines,
their parallelism with the trend of the cell walls;
their regular spacing ... I do not wish to assert that the section here shown
proves conclusively the presence of true sporangia ... "
Elliott (1965): "may be sporangia 1structure ... may well be sporangia, but this
is still uncertain."
Cuif (1969) finally found in the Triassic of Turkey some Solenoporaceae
with areas presenting a remarkable resemblance to reproductive organs; the
cells were created by the differentiation of cells of the thallus.
In short, there is apparently no certainty as to whether a structure can be
seen in cells without any regular shape, similar to that of reproductive organs.
These "rosette", "star-like", or "miliolid" -shaped cells are only vegetative cells.
There is nothing to show reproductive organs in the cases mentioned. The
geometric arrangement in the tissue in no way corresponds to that already
known about the Corallinaceae. The relationship between cellular perithallus
and reproductive organs is not clear. Undoubtedly we are concerned here
with foreign bodies, present in the tissue during its growth, or with traces of
decaying organisms, of various scars, and of pre-, pene-, or post-diagenetic
dissolution.
4 Systematics and Classification
The classification of the Solenoporaceae is very difficult to establish as the

criteria which may be used are scarce and not very reliable. Johnson (196 J)
stated that there are:
"Difficulties in identifying and interpreting Solenoporaceae, both because of

the nature ofthe calcification ofthe original plants and because of the effects of
solution and recrystallization in the course of fossilization."
Indeed, the distinction between hypo thallus and peri thallus is not clear
because, e.g. the presence of reproductive organs is not certain, the pores and the
rosettes cannot be used as criteria which playa part in the classification system,
and the polygonal or circular appearance of the tubes in cross-sections cannot
be used to back up an argument.
Yet, other characteristics, although not very accurate, can be used; Elliott
(1965) stated:
"It should be evident that the Solenoporaceae family does not include any
representatives which have very loosely packed filamentous cells such as are
seen in Pycnoporidium. All the genera of the Solenoporaceae have a compact
tissue."
Indeed, the positioning of partitions form the basis of Pia's (1930)

classification system:
A) Shapes with pores in rows
I. Cells irregularly dispersed, few horizontal partitions
Solenopora = Metasolenopora
II. Cells in rows, horizontal partitions clearly represented
Parachaetetes
B) Shapes with isolated pores, rarely found in rows
I. Cells irregularly dispersed, horizontal partitions clearly represented
Pseudochaetetes = Solenoporella
II. Cells in rows, horizontal partitions clearly represented
Petrophyton
Maslov (1961), as a continuation of Peterhans (1929b), suggested:
A) Pores in concentric rows
cells irregularly arranged
not very many horizontal partitions
Solenopora = Metasolenopora
Cells in concentric rows
clearly defined horizontal partitions, on the one level
Parachaetetes
B) Isolated pores, rarely found in concentric rows
horizontally arranged cells
clearly defined horizontal partitions
Pseudochaetetes - Solenoporella
cells in concentric rows
horizontal partitions on one level
Petrophyton
94 A.F. Poignant
Later, various classifications were suggested, varying from author to author

and even varying with the same authors. There is no point in referring to all the
different theories. The various authors are quick to deny use of pores and cell
shapes in cross-section when making a classification, and only make use of the
arrangement of the partitions. If these horizontal partitions are on the same
level, we are dealing with the genus Parachaetetes; otherwise the genus in
question is Solenopora. Mamet and Roux (1977) have shown the possible
existence in cross-section of pseudosepta (identations); Peterhans (1929b) had
already drawn attention to this fact. It concerns characteristic structural
elements, from which one sees the trail of longitudinal, incomplete partitions,
more or less developed inside the tubes. There may also be some real
bifurcations.
According to Mamet and Roux (1977):
A) Shapes form with longitudinal partitions
without horizontal partition
Solenopora
with horizontal partitions
Parachaetetes
B) Shapes without longitudinal partitions
without horizontal partition
Pseudosolenopora
with rare and irregular horizontal partitions
Pseudochaetetes
with frequent and regular horizontal partitions
Petrophyton
According to Wray (1977): Rhodophyta
Rhodophyceae
.Floridae
Cryptonemiales
Solenoporaceae
Gymnocodiaceae
Squamariaceae
Corallinaceae
4.1 Phylogenetic Theories
For 50 years, numerous phylogenic theories for transition from Solenoporaceae

to Corallinaceae were suggested:
Maslov (1956)
Johnson (l960b): The common ancestor splits into two at the Ordovician-
Silurian boundary:
Sl near Archaeolithothamnium, Mesophyllum, and Lithothamnium;

S2 near other Corallinaceae.
Endo (1961): The ancestral Cambrian form (not named) divides three times:
Solenopora, then Petrophyton, Pycnoporidium, and Nipponophycus;
Parachaetetes;
Pseudochaetetes.
It is unfortunate that this original idea has not been elaborated on since.
Johnson (1960a): There is a separation at the lower part of the Silurian into
Solenopora (Lithoporella, Lithothamnium, Archaeolithothamnium) and Para-
chaetetes (the other Corallinaceae).
Johnson (1963): This theory is different since, from the separation, we see
Solenopora without descendants and Parachaetetes (Mesolithon, Archaeoltho-
thamnium, Mesophyllum, and Lithothamnium). The other Corallinaceae have
a phylogenesis which is independent of the Solenoporaceae.
Wray(1970): On the one hand, the separation of the common ancestor ends in
Solenopora going on to Archaeolithothamnium, Lithothamnium, and Meso-
phyllum and, on the other hand, it ends in Parachaetetes from which more or less
directly all the other Corallinaceae originate.
Wray (1977a): The separation starting from the Silurian shows the continua-
tion on the Solenoporaceae and the derivation of the Squamariaceae and the
Corallinaceae.
Poignant (1976, modified in 1979): Solenopora gave rise to Paleothamnium and
Lithothamnium.
Of course, all these proposed phylogenic ideas must be restudied if Mamet
and Roux's new classification is to be adopted.
Indeed, Solenopora and Parachaetetes seem to be linked by the presence of
indentations, whereas other forms do not have any; for example, indentations
are unknown in the Corallinaceae. Perha ps, one must look for the ancestor ofthe
Corallinaceae in forms which have no indentation (Pseudoso/enopora,
Pseudochaetetes, and Petrophyton).
In short, as long as a certain system is not established, the stratigraphical
distribution is meaningless.
5 Conclusion
Now is the time to identify those genera which make up one distinct family. The
Solenoporaceae have been in existence for 500 million years and, throughout
this length of time, they have been important rock formers.
The apparent fall-off of the Solenoporaceae seems to have begun during
important ecological changes at the beginning of the Mesozoic. The Corallin-
aceae benefitted from this decline in Solenoporaceae and filled biotopes which
are slightly different. Indeed, the almost certain absence ofthe hypothallus gives
the Solenoporaceae certain growth faculties which differ from those of the
96 A.F. Poignant
Corallinaceae. One may also add that the type of reproduction, unknown in the
case of the Solenoporaceae, was without doubt less efficient than that found in
their rivals.
There is now a growing need for the knowledge based on Mamet and Roux's
classification. The microscopic structure of these algae must be examined using
the electron microscope. One must also strengthen the classification system and
put it into some kind of order.
References
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London 56: 147-148, I pi
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23 pi
Johnson JH (I 960b) The Jurassic Algae. Col Sch Mines Q 59, 2: 129,45 pi
lohnson JH (1961) Review of Ordovician algae. Col Sch Mines Q 56. 2-1: 1-101
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Johnson lH (1966) A review of the Cambrian algae. Col Sch Mines Q 61. I: 1-162
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68 pi
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36 pi (French translation BRGM 384)
Nicholson HR (1888) On certain anomalous organisms which are concerned in the formation of
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36:1-8,3 pi
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Peterhans E (l929b) Algues de la famille des Solenoporacees dans Ie Maim du Jura Mlois et
soleurois. Mem Soc Paleontol Suisse I L: 1-15, 7 pi
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23:37-39,2 pi
Pia H (1930) Neue Arbeiten tiber fossile Solenoporaceae und Corallinaceae. N Jahrb Mineral Geol
Palaontol3: 122-147
Pia J (1939) Sammelbericht tiber fossile Algen; Solenoporaceae 1930 bis 1938, mit Nachtragen aus
frtiheren Jahren. N Jahrb Mineral Geol Palaontol3 :731-760
Pia J (1940) A new fossil alga (Solenoporaceae) from the Jurassic rocks of Western Australia. J R
Soc West Aust 26:29-33, 3 pi
Poignant AF (1976) La phylogenie des Algues rouges. C R Acad Sci Paris 282: 1677-1680
Poignant AF (1979) Les Corallinacees mesozoiques et cenozoiques: hypotheses phylogenetiques.
Bull Cent Rech Explor Prod Elf-Aquitaine 3,2:753-755
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pp 202-214
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Vetenskap 43: 1-25,6 pi
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Obersilur Gotlands. Sver Geol Unders 10: 1-57,9 pi
Vialli V (1938) Su Taluni fossili liassici del Monte Peller. Stud Trentini Sci Nat 19: 1,2 pi
Wood V (1944) Organs of reproduction in the Solenoporaceae. Geol Assoc London 55: 107-113,
pi 5-6
W ra y JL (1967) Upper Devonian calcareous algae from the Canning Basin, Western Australia. Prof
Contrib Col Sch Mines 3: 1-76, 11 pi
Wray JL (1970) Algal in reefs through time. Proc N Am Palaeontol Chicago 1969:1359-1373
Wray JL (I 977a) Calcareous algae. Developments in palaeontology and stratigraphy, vol4. Elsevier,
Amsterdam, 185 pp, 170 fig
Wray JL (l977b) Late Paleozoic calcareous red algae. Fossil Algae Spec Vol, pp 167-176
Yabe H (1912) Uber einige gesteinbildende Kalkalgen von Japan und China. Sci Rep Tohoku Imp
Univ Ser 3 Geol Sendai I, 1:1-8,3 pi
Chapter 5
CoraUine Algae: Mineralization, 1axonomy, and
Palaeoecology
D.W.J. BOSENCE1
Abstract
This critical review focusses on three areas of recent research on the crustose coralline algae:
Calcification and Diagenesis

Recent work has clarified the calcification of corallines as being a two-stage process; firstly,
tangential and secondly radial calcite is deposited within the matrix of the cell walls. This internal
calcification of cell walls is unique to the coralline algae and should be used to interpret early
problematic red algae in which other characteristic structures are not present. Recent descriptions
of subcrustal aragonite from corallines show that this form of mineraliza tion is not restricted to the
squamariaceans, as has been generally considered.
Taxonomic Revisions of Genera

New investigations on Recent coralline algae by workers in Australia and Britain is leading to
important revisions of well-known Recent and fossil genera. The time is appropriate to assess the
implications of this work for palaeontology. Important questions are the preservation of taxonomic
characters and the relationships between fossil taxa and groups of Recent taxa. Specific deter-
mination of many fossil corallines suffers from excessive splitting on the basis of too few characters.
The importance of detailed measurements and statistics is emphasized.
Ecology and Palaeoecology of Coralline Algae

The longevity of coralline genera, together with their ecological restrictions and plasticity of growth
forms in relation to environmental parameters, makes the corallines a good group for palaeoen-
vironmental analyses. Recent work has shown the strong relationships between growth form and
hydraulic energy; and the generic composition of floras and water depths. The latter has exciting
potential for erecting quantitative palaeobathymetric zones for the Cenozoic.
1 Introduction
Most of the major advances in corallinology at present are being made by those
studying Recent corallines, particularly in the study of calcification, generic
classification and morphology. An excellent review of this recent work is to be
found in the text by Woelkerling (1988). Palaeontologists, meanwhile, have
made some progress in generic concepts and in palaeoecology. In this post-
Harlan-Johnson era there still remain major problems in the validity of fossil
IDepartment of Geology, Royal Holloway and Bedford New College. University of London.
Egham. Surrey TW20 OEX. UK
Coralline-Algae: Mineralization, Taxonomy, and Palaeoecology 99
coralline species and there is little palaeontological data on which to build a

phylogeny of the coralline algae.
In this review the palaeontological implications of the recent advances on
present day crustose corallines and new developments in ecology and
palaeoecology are discussed under the following headings:
I. Calcification and diagenesis;

2. Taxonomic revisions of coralline taxa;
3. Ecology and palaeoecology.
2 Calcification and Diagenesis
The coralline algae are the most consistently and heavily calcified group of red
algae and as such have recently been elevated to ordinal status (Corallinales:
Silva and Johansen 1986). The calcification involves high magnesium calcite
(and occasionally brucite, Weber and Kaufman 1965) precipitation within most
cell walls (Figs. IA-C, 2A-D). Other calcified red algae are the Solenoporaceae,
the Squamariaceae (Denizot 1968) and some members of the Nemalionales
(Galaxaura, Liagora and Nemalion: Dixon 1973).
Early electron microscope work showed that the calcified cell walls of
coralline algae had a two-layered structure: an inner layer of acicular calcite
parallel to the cell wall was succeeded by radial, inward-growing calcite crystals
(Figs. lA, 2C; Bailey and Bisalputra 1970; Alexandersson 1974, 1977; Flajs
1977a,b; Garbary 1978). Flajs (1977a,b) described two cell-wall types: the
Lithothamnium-type with the two layered structure described above and a
Goniolithon-type with a central noncalcified zone and outer radial crystals. The
Lithothamnium-type has subsequently been described from a wide range of
corallines (Mesophyllum, Lithothamnion, Lithophyllum (Fig. 2C), Phyma-
tolithon, Porolithon, Corallina and Jania) by Cabioch and Giraud (1986) and
Massieux et a1. (1983) and is also seen in the extinct Solenopora (Flajs 1977b).
However Flajs' (1977b) Goniolithon-type has neither been confirmed by other
authors nor has it been found in other genera. More recent work in the 1980s
using SEM and TEM techniques on calcified tissue has significantly advanced
our knowledge of calcification (Massieux et a1. 1983; Walker and Moss 1984;
Cabioch and Giraud 1986). However the physiological processes involved in
calcification within the cell wall remain unclear. Experimental work by Digby
(1979) showed how, in the normally reducing environment of the thallus,
increased alkalinity is brought about by oxidation through strong oxidase,
catalase and carbonic anhydrase activity and the removal of hydrogen ions.
More recently Okazaki et a1. (1982) have shown that alginic acid is associated
with cell wall calcification and that this acid induces calcite deposition and
strongly inhibits aragonite deposition. Cabioch and Giraud (1986) have found
no ultrastructural differences between calcified and uncalcified fioridean cells.
In an ultrastructural study of a range of temperate corallines they have shown
that calcification is a two-stage process which relates to the two calcified zones
100 D.W.J. Bosence
2 STAGE CALCIFICATION CEMENTATION OF CELL CAVITY

A OF CELL WALLS
B
----=-~ ---.-
~ -=---=-=-.- ~---
needle crystals
Epithallu8
Perllhallu8
LOW Mg CALCITE DIAGENESIS

C
Low Mg calcite
cement
Dissolved and repreclpltated

D
cell wall
Conceptacle cement
SUBCRUST AL ARAGONITE BOTRYOIDS

Fig. lA-D. Calcification and diagenesis. A Sketch from electron micrograph (Cabioch and Giraud
1986) illustrating cell wall calcification. Early needle calcite is parallel to cellwalls ; later radial calcite
mineralizes epithallial and perithallial cell walls, x2,OOO. B Sketch indicating two layered coralline
cell wall with later infilling of cell cavities by radial calcite after disintegration of cell contents, x2,OOO.
C Diage nesis of coralline tissue. Dissolution of high magnesium calcite and reprecipitation of low
magnesium calcite in cell wall. Low magnesium calcite cementation of cell cavity, x2,OOO. D
Su bcrustal aragonite botryoids growing in spaces between crusts and early aragonite cements
infilling conceptacles, x200
of the cell wall described above . Whereas Flajs (1977a) considered calcification
to be a relatively late-stage process linked to cell degeneration, Cabioch and
G ira ud (1986) have shown that it occurs around the epithallus which contains
the uppermost cells of the thallus (Fig. lA). The first phase of precipitation
involves the growth of needle-shaped crystals within and parallel to polysac-
charide fibrils between filaments in a position analogous to the middle lamella
of higher plants. With growth, these needle-shaped crystals are replaced or grow
into the platey and needle-shaped crystals of the middle cell wall. Later, bladed
to fibrous calcite crystals grow radially and parallel to an organic matrix to form
the secondary layer at right angles to the primary layer (Figs. lA, 2C). The
mineralogy of these different phases has not been described but earlier work by
Flajs (1977b) and Massieux et al. (1983) using microprobe plotting ofmagne-
sium and calcium concentrations indicates that the secondary radial layer has a
higher magnesium content than the earlier magnesium-poor layer.
When dead tissue is buried within the thallus by outward surface growth,
cells are frequently further infilled by layers of micron-sized magnesium calcite
crystals (Figs. IB, 2F) which may eventually fill the cell cavity (Alexandersson
1974; Bosence 1985). This micrite infilling of cells is patchy and is clearly seen
in Recent dead, and fossil material (Fig. 2F). Alexandersson (1974) suggests that
decaying cell contents may trigger off this later calcification. The supply of
calcium carbonate for crystal growth is a problem as there appears to be no local
dissolution. Therefore, it would appear to have to come from pore waters
passing through the cells and the outer living tissue. An additional or related
problem is that whilst most of the cells are infilled with high magnesium calcite
others are infilled with aragonite (pers. observations and analyses). This may
help to explain earlier reports (Winland 1969; Milliman 1974) of aragonite
replacing coralline grains in shallow tropical environments. In extensive surveys
of coralline skeletons from the Caribbean I have never seen any replacive
textures (e.g. Bosence 1983, 1984, 1985). Oti and Muller(1985) have studied the
fine-scale alteration of the high magnesium calcite skeleton to low magnesium
calcite both experimentally and in Cenozoic fossil corallines. They show that the
process is a small-scale dissolution and reprecipitation replacement of the
original cell walls such that the wall microstructure is lost but the walls are
recognizable from later low magnesium calcite cements in the cell cavities (Fig.
lC).
Extracellular cements are also common in the corallines (Massieux et al.
1983; Bosence 1985) and the Squamariaceae (Denizot 1968). The commonest
are subcrustal botryoids of aragonite (Fig. ID) which have been described from
living, dead and fossil corallines. When first described these were described as
early diagenetic cements (Alexandersson 1974) but recently Walker and Moss
(1984) have shown that the subcrustal aragonite crystals contain concentrically
arranged organic material within the fans which is in intimate association with
the basal layers of the alga. Walker and Moss (1984) consider that the fans may
have a functional importance to the algae in re-attaching loosened crusts to their
substrates. Although similar, but now calcitized, subcrustal fans are known in
fossil material it would be difficult to decide whether they were laid down
extracellularly by the algae or as chemical precipitates. Other, similar, prismatic
aragonite cements are commonly found in growth cavities, vacated concepta-
cles, etc. (Alexandersson 1974; Bosence 1985). A variety of subcrustal aragonite
fans and palisades has been described from squamariacean algae by Denizot
(1968). Recently James et al. (1988) have compared these with the common
calcite crusts formed beneath Palaeozoic phylloid algae. These calcite crusts
102 D.W.J. Bosence
may now also be compared with subcrustal cements found below coralline
algae.
Other palaeontological implications of this work on calcification include
the recognition of coralline algal cells and filaments in fossil material when
differentiated thallus (Fig. 2 a) or conceptacles are absent. The red algae are the
only algal group in which the cell and filament walls are calcified (intracellular
calcification). In addition, the cell walls have a differentiation into primary and
secondary layers (Figs. 1, 2f). It is therefore reasonable, in the absence of other
morphological characters, to use this as an identifying character for fossil
calcified red algae. Thus, the Solenoporaceae have been shown to have calcified
cell and filament walls which are two-layered (Flajs 1977b), and, in the absence
of differentiated tissue and reproductive structures, it seems most appropriate to
view this group as an extinct family of the red algae. Similarly, in the absence of
any other data it is reasonable to assume that Cuneiphycus and Archaeolitho-
phyllum are fossil red algae as has been suggested by Wray (1977). Such
problematic algal groups as the ungdarellids would repay detailed study with
polished and etched sections under the SEM, which should reveal the nature of
the calcified tissue.
3 Taxonomic Revisions of Coralline Taxa
3.1 Morphological Characters
An important aspect of the systematics of the coralline algae is the relationship

between Recent and fossil taxa, the most stable and fundamental taxon in this
context being the genus. In defining genera many of the taxonomically im-
portant characters can be recognized in both fossil and Recent forms. However
in Recent material there are additional morphological features such as
developmental and staining characteristics (Table 1). The commonly used
characters for describing fossil genera according to Lemoine (1977), and
Fig. 2a-f. Coralline algal microstructure. a Vertical section through crust of Porolithon illustrating
basal hypothallus with horizontal filaments and upper perithallus with vertical filaments and with
horizontal row oftrichocytes in centre. Recent, St. Croix, US Virgin Islands, x220. b Vertical section
through crust of Lithophyllum illustrating perithallial tissue and single layered outer layer of
epithallus. Note increase in cell wall calcification with depth in thallus. Recent, leddah, Saudi
Arabia, x500. Coli. L. Montaggioni. c Detail ofb illustrating cell wall calcification in Lithophyllum.
Thin central zone with outer radial cell wall crystals. Note pit connections between contiguous
filaments, xSSO. d Detail of calcified cell wall in vertically arranged perithallial filaments with
primary pit connection between adjacent filaments and cells. Lithophyllum. Recent, leddah, Saudi
Arabia, x2,500. ColI. L. Montaggioni. e Preservation of epithallus beneath overgrowing foramin-
ifera in Recent Lithophyllum. Note patches of cells infilled with micrite. Lithophyllum. Recent,
leddah, Saudi Arabia. General view, left, xl SO, zoom, right x720. Coli. L. Montaggioni. fFractured
surface of Lithothamnion with perithallial filaments arranged lower left to top right. Note secondary
cell fusions (upper left) and micrite infilled cells (right). Recent, Co. Galway, Ireland, x2000
104 D.W.J. Bosence
1llble 1. List ofcharacters used in the description and assignment of Recent and fossil coralline algae
Generic characters Occurrence of characters"

Fossil
Recent
Commonly used Potentially usable
Morphology
General
Crustose
•• • •
Branching
•• •
Columnar
•• •
Mamillate • •
Articulated
• • •
Articulate branching
Epithallus
• 0 0
Thickness
•• 0
•
Cell Shape 0
•
Meristem
Peri thallus
• 0
•
Presence/absence • • •
Filaments/rows
• • •
Secondary cell fusions
• 0
•
Secondary pit connections
•• 0
•
•
••
Double perithallus
Trichocytes
Medullary perithallus
• •
Presence • • •
Thickness • • •
Cell height
Cell shape
•
•
•• •
•
Genicula
Hypothallus
• 0 0
Single layered • • •
Palisade • • •
Thickness • • •
Coaxial/non-coaxial
• • •
Median hypothallus
Asexual Conceptac1es
• • •
Location
• • •
Perforations
• • •
Shape
• • •
Primordia
• 0
•
Tetraspores
Sexual conceptac1es
• 0
•
Location
• • •
Shape
• • •
Spermatangium
• 0
•
Carpogonium
• 0
•
Carpospore • 0
•
Biochemical
Staining of sporangia I
caps
• 0 0
Developmental
Spore cell division
• 0 0
• •
a.
Conceptacle development 0
Yes; 0 no.
Poignant (1984, 1985) are also listed (Table I) and they represent some 64% of
characters available to algologists. However, I argue below that it may be
possible, with more detailed thin section and SEM observations, to increase the
morphological characters available to palaeoalgologists.
Epithallus. It has been generally assumed by palaeoalgologists (Wray 1977;

Lemoine 1978a) that the epithallus is uncalcified and, as such, is not a preserv-
able character. The reason for this may be that until recently all present-day
material had been illustrated from decalcified material. However, with the
publication of SEM micrographs of fractured surfaces of fresh and dried
museum specimens (e.g. Woelkerling et al. 1985, 1988), it is clear that the
epithallus can be well-calcified and it is difficult to see why it should not be
preserved. The most likely circumstance for its preservation is where crusts are
overgrown by later crusts and are therefore protected from bioerosion or
abrasion (Fig. 2e). Such an occurrence has been recently illustrated by Voigt
(1981). Recognition will require careful SEM work, together with a knowledge
of epithallial tissue and location ofthe perithallial meristem. Ifthis tissue can be
differentiated in fossils then it will have important implications because
epithallial characters are used to differentiate between Lithothamnion,
Phymatolithon and Leptophytum (Adey 1966, 1970). These closely related
genera can only be differentiated in the Recent by epithallial cell shape, length
of meristematic cells, and the phosphotungstic haematoxylin staining of the
sporangia I caps. At our present state of knowledge they are all grouped into the
genus Lithothamnion in fossil material.
Secondary Cell Connections. Connections between contiguous cells are clearly

seen in living tissue (Fig. 2) and occur as two types. Secondary pits are small (1-2
p.m) circular pits on opposing sides of cell walls separated by a plug (Figs. 2 c - d).
They are considered by Adey ( 1970), Johansen (1981) and W oelkerling(1987b,
1988) to be of fundamental importance in coralline systematics and of ordinal
significance in the red algae (Pueschel and Cole 1982; Silva and Johansen 1986;
Gabrielson and Garbary 1986). Cell fusions are larger, irregular connections
between rows of cells or along filaments of cells (Fig. 20. The sub familial
classification of Recent Corallinaceae by Johansen (1981) recently modified by
Woelkerling (1987a; Table 2), emphasises the taxonomic importance of these
characters. As far as I am aware, neither of these structures has been described
from fossil corallines although they are clearly seen in fractured Recent
specimens (Fig. 2; Woelkerling et al. 1985) and in diagenetically altered Recent
rhodoliths from the Skagerak (Alexandersson 1974) and the Red Sea (Flajs
1977b). If diagenesis involved small-scale solution and reprecipitation as-
sociated with the change from high to low magnesium calcite, and the cell lumen
filled with a coarser calcite cement, then these cell wall characters should be
visible in fossils under the SEM as they are in diagenetically altered Recent
material.
106 D.W.J. Bosence
Table 2. Outline of subfamily classification of Corallinaceae (after Johanse.n 1981: Woelkerling

1987a)
Thallus Cell fusions Secondary pit Conceptacles;

joining connections tetras pores
contiguous joining conti-
filaments guous cells
Amphiroideae Geniculate with no yes Uniporate;

genicula of one tetraspores
or more tiers lacking plugs
of cells
Corallinoideae Geniculate with yes no Uniporate;
genicula of tetraspores
single tier of lacking plugs
cells
Metagoniolithioideae Geniculate with yes no Uniporate;
genicula not tetraspores
aligned in tiers lacking plugs
Lithophylloideae Non-geniculate no yes Uniporate;
tetraspores
lacking plugs
Mastophoroideae Non-geniculate yes no Uniporate;
tetraspores
lacking plugs
Melobesoideae Non-geniculate yes no Multiporate;
tetras pores
with plugs
Choreonema toideae Non-geniculate no no Uniporate;
tetras pores
with plugs
3.2 Location and Development ofConceptac1es and Spores
The details of conceptac1e development and of tetras pores and carpospores is

only rarely considered in fossil material even though spores often remain in
conceptac1es and become mineralized (e.g. Bosence 1983). Therefore, it is urged
that more detailed descriptions ofthe position and development of conceptac1es
and the nature of tetraspores and carpospores be given in future.
Geniculae. The decalcified joints (genicula) of articulated corallines are not

normally preserved. However, in unusual circumstances articulated corallines
may be preserved in situ by overgrowing crustose corallines (e.g. Bosence 1984)
and the genicula and margins of the intergenicula thus preserved. Although this
is the only case I am aware of, if articulating corallines encrust other colonial
organisms they can potentially be overgrown by their host and preserved intact.
If the above criteria can be considered and fossils carefully examined under
the SEM, then the proportion of preservable characters which might be
recognized in both fossil and Recent material rises from 64% to a respectable
89%. The number offairly broadly defined characters used in identifying genera
rises to 34, which is a substantial figure for any fossil group and should form the
basis for an accurate linkage between fossil and Recent taxa.
3.3 Recent Taxonomic Revisions
At present, corallinologists in Australia, Britain and America are substantially

revising the morphology, types and taxonomy of coralline genera. Many of the
original type specimens have been located and examined under optical
microscopy, and with an SEM on fractured surfaces, and the morphology
redescribed in detail. Much of this work is contained in the excellent new text
by Woelkerling (1988). Whilst it is not possible here to review the large number
of papers which have been published in the last 3 years, the results so far indi-
cate important revisions of coralline genera. I briefly discuss below the major
revisions published to date and palaeontological implications of this work.
Dermatolithon. Woelkerling et al. (1985) suggest that this thin, crustose form
cannot be differentiated from Titanoderma as both have a dorsiventrally ar-
ranged prostrate crust with a single layered palisade hypothallus, secondary pit
connections and unipored conceptacles.
Leptophytum. Woelkerling and Irvine (1986) consider that this genus requires
clarification as a taxon which mayor may not be distinct from the very similar
Phymatolithon.
Litholepis. Woelkerling (1986) shows that this genus is a heterotypic synonym

of Titanoderma as they both have secondary pit connections, unipored tetra-
sporangial conceptacles, single layered palisade hypothallus and dorsiventral
crust organisation. Secondary cell fusions are absent in both genera.
Lithothamnion. (Heydrich 1897) is now conserved against Lithothamnium

Philippi (1837) as Woelkerling (1983, 1985a) has shown that all Philippi's type
material bears unipored conceptacles and is a mixture of species of Goniolithon,
Lithophyllum and Pseudolithophyllum.
Paragoniolithon. Woelkerling (1987b) has recently reexamined this genus and

found it to be a heterotypic synonym of Neogoniolithon, as both have trichocytes
singly or in vertical stacks in an otherwise similar thallus.
Pseudolithophyllum. This genus has had a varied history but present views are
that it should be considered in the sense of Lemoine (1978b) and Mendoza and
Cabioch (1984) rather than that of Adey (1970). The thallus has a single-layered
hypothallus and the peri thallus has rectangular cells arranged in filaments
joined by secondary pit connections.
108 D.W.J. Bosence
Spongites (KUtzing). This genus has been resurrected by Woelkerling (1985b)

to include crustose corallines with a multilayered, non-coaxial hypothallus and
a perithallus with trichocytes occurring singly or in vertical stacks.
Tenarea. Woelkerling et al. (1985) consider that this genus, with its isobilater-
ally arranged two rows of hypothallial cells, is clearly distinct from Titano-
derma. They also suggest a similarity with the isobilaterally arranged fossil genus
Distichoplax.
These taxonomic revisions are particularly important to palaeoalgologists
as the characters which are now being used by algologists are those which are in
the main preservable and are visible in fossils. In addition, genera such as
Litholepis, Paragoniolithon and, probably, Leptophytum which have not to my
knowledge been recognized in fossil material are no longer considered separate
genera. Hopefully this will give confidence to palaeoalgologists in that the
criteria that have traditionally been used are now being used by algologists.
However, when examining fossil material the additional characters should be
looked for (Table 1) which may only be recognizable under the SEM.
There are very large problems concerning the identification of fossil species.
Most specific identifications require careful measurements of crust and branch
thickness, hypothallus thickness, hypothallial and perithallial cell sizes and
conceptacle shapes and sizes. Because these characters are naturally variable the
mean and standard deviation as well as the ranges are required if specimens are
to be compared. Many palaeontologists have follow.ed Johnson (in Johnson and
Adey 1965) who only used size ranges and many species described by Johnson
are from incomplete material. It is therefore very difficult to compare specimens
with published material and much work is required redescribing fossil types.
4 Ecology and Palaeoecology
Work in recent years on the variable morphology of present day coralline algae
and the generic floral associations in different climates and water depths has
provided a good database for palaeoecological and palaeoenvironmental analy-
sis. This information can be confidently applied to ancient examples because
of the generally good preservation of coralline thallus morphology and the
relative ease of identification of corallines to generic level. In addition most
coralline genera are extant and have an average longevity of85 Ma (data from
Wray 1977). This increases the confidence with which comparisons can be made
between present-day coralline ecology and ancient corallines.
4.1 Relationships Between Morphology and Turbulence
The morphology of coralline crusts and branches in present day reefs was
considered by Bosence (1983, 1985). I showed then that high energy (i.e. reef
front or algal ridge) locations are typified by thick crusts which are closely
CRUSTOSE & BRANCHING CORALLINE FRAMEWORKS

A HIGH ENERGY B MODERATE ENERGY
Robust fused framework Delicate framework with
with intraskeletal cement intra skeletal cement
C RHODOLITH FORM
•
Radial Branching Concentric Crusts
~ INCREASING ENERGY AND TURNING

•
o
•
2 em
Fig.3A-D. Relationships between water energy and coralline thallus. A Typical growth forms of
crusts and branches in high energy coralline reefframeworks. B Typical growth forms of crusts and
branches in coralline reefframeworks in moderate energy settings (after Bosence 1985). C Increase
in lateral growth of branches and density of branching ofrhodoliths with increasing turbulence and
development of dense concentric crusts (after Bosence 1976). D Increase in numbers of branching
directions with turning ofrhodoliths (after Bosence 1983)
superposed (Fig. 3A). Branches and columns are thick and perithallial tissue
from neighbouring branches may fuse to further strengthen the framework. The
framework is also made rigid by the extensive extracellular cementation in this
environment. Coralline genera commonly forming such frameworks are
Lithophyllum and Porolithon. Reef frameworks from medium energy reefs and
buildups such as maeri, back-reef mounds, and the deeper water coralligene
(Bosence 1983, 1985) typically have thinner branches and crusts, are more
delicate in their construction (Fig. 3B) and are formed ofleafy crusts which may
leave the substrate and overgrow neighbouring crusts. Adjacent branches do not
fuse together. Corallines found forming such leafy and branching frameworks
are Mesophyllum, Lithophyllum, Titanoderma, Neogoniolithon, Lithothamnion
and Phymatolithon.
ItO D.W.J. Bosence
In a recent discussion of coralline ecology and functional morphology

Steneck (1986) describes similar ranges of coralline morphologies as described
above. He discusses experimental work which suggests that the thicker crusts are
better able to resist physical and biological disturbance. The major biological
disturbance is considered to be herbivory by parrot fish, sea-urchins, limpets and
chi tons (Steneck 1986).
Free-living rhodoliths (rhodolites) can be very sensitive indicators of tur-
bulence in the shallow marine environment. Within anyone species the mor-
phology of free living forms can vary from open-branched through to densely
branched forms (Cabioch 1966; Bosellini and Ginsburg 1971; Bosence 1976
(Fig. 3C). Open-branched forms are found in the quieter water environments
where there is little transport of rhodoliths. If rhodolith transport is very
infrequent in comparison to branch growth rates, then zones of redirected,
upward growth is seen in the rhodoliths after turning (Fig. 3D). The densely
branched forms are typical of rhodoliths growing in higher energy conditions.
The increase in branch density is achieved by lateral branch growth and lateral
growth following apical abrasion during transport. This may be followed by
concentric crustose growth ofthe original algae or other encrusters. Stabilization
results in reversion to branching growth (Bosence and Pedley 1982). In addition
to turning by currents, rhodoliths may also be turned by organisms, as was
suggested for rhodoliths from a wackestone facies in the Miocene of Malta in
which crustacean debris is common (Bosence and Pedley 1982).
4.2 Palaeobathymetry
The establishment of depth/abundance charts for coralline floras from the

Caribbean and Pacific sites by Adey (1979) indicates the great potential that the
coralline algae have for palaeobathymetry. Since present-day crustose genera
have existed from the Miocene, the following bathymetric ranges should be
applicable for the Neogene:
Intertidal-20 m; Neogoniolithon, Porolithon, Lithophyllum and

Hydrolithon.
20-40 m; Neogoniolithon, Lithophyllum, Hydrolithon, Titanoderma and
Mesophyllum.
40-60 m; Mesophyllum dominant with Archaeolithothamnium, Litho-
thamnion and Lithophyllum.
60- 100 (possibly 200 m); Mesophyllum and Lithothamnion with Ar-
chaeolithothamnium and Lithophyllum.
The only way of confirming these depth sequences back through time is to
analyze the distribution and abundance of in situ crusts where the palaeodepth
can be measured. Such sites are on very well exposed reef fronts where
depositional units can be traced down from the reef flat. Such sites should be
investigated to ascertain the depth distributions of Neogene coralline floras.
Coralline-Algae: Mineralization, Taxonomy, and Palaeoecology III
5 Conclusions
This review concludes with two important points for palaeoalgologists. Firstly,
that there has been a history of confusion with respect to the taxonomy of
coralline genera which is only recently being corrected. The characters which are
being used to redefine genera are, in the main, preservable but palaeoalgologists
must use the SEM to obtain additional characters concerning the epithallus and
cell wall perforations to increase confidence in their determinations. Thus, at the
generic level it is possible for algologists and palaeoalgologists to work closely
together to establish evolutionary relationships and higher taxonomic group-
ings. However, at the species level a large amount of work is required on fossil
types to provide good quantitative descriptions and illustrations.
Secondly, with respect to palaeoecology it is now realized that thallus shape
together with the generic associations give very good palaeoenvironmental
information. Thus, coralline thalli are amongst the most sensitive skeletons to
changes in hydraulic energy and should now be used in detailed palaeoenvi-
ronmental analysis. Similarly, generic associations of floras which can be
accurately assessed in fossil sequences give the opportunity for quantitative
palaeobathymetry which has wide-ranging uses in basin analysis.
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Chapter 6
Cyclocrinitids
S.c. BEADLEI
Abstract
The cyclocrinitids are a small group of middle Ordovician to early Silurian macrofossils. They are
usually regarded as an extinct tribe of dasycladacean algae, since they are morphologically very
similar to the living dasycladacean Bornetella. Cyclocrinitids had a central main axis bearing
numerous lateral branches, which radiated outwards in all directions. The branches expanded at
their tips to form thick lateral heads, which were generally hexagonal in outline and usually 1-3 mm
wide. The lateral heads were tightly packed together to form a faceted outer cortex, which was
spherical to claviform in shape and typically 1-4 cm in size. The thallus was encrusted by calcium
carbonate, particularly around the lateral heads. Cyclocrinitid preservation is quite variable, and
this has often confused taxonomists; new descriptions of the European and Asian forms are needed.
Cyclocrinitids are important associates of many benthic invertebrate palaeocommunities. They are
often abundant locally and may be significant rock-builders; several rock units have been informally
named after them. Cyclocrinitids were restricted to shallow, quiet waters at low latitudes. Eco-
phenotypic thallus-size variation within Silurian species has been used to indicate relative
palaeodepths. Ordovician cyc10crinitids have much potential value as palaeoenvironmental
indicators.
The Tribe Cyclocriniteae was named by Pia (1920) and emended by Bassoullet
et al. (1979). It includes the Ordovician and Silurian genera Cyc/ocrinites
Eichwald, Mastopora Eichwald, Coelosphaeridium Roemer, and Apidium
Stolley. It should be noted that the genus name Cyc/ocrinites (not Cyc/ocrinus)
has priority, so the tribe is the Cyclocriniteae (not Cyclocrineae). Furthermore,
Mastopora is now commonly regarded as a junior synonym of Cyclocrinites (as
are Nidulites Salter, Pasceolus Billings, Cerionites Meek and Worthen, and
Lunulites Owen). Nitecki (1970, pp. 73-75) elaborates on these points.
The history of cyclocrinitid research is discussed by Osgood and Fischer
(1960) and Nitecki (1970). Early workers variously regarded cyclocrinitids as
foraminifers, sponges, corals, bryozoans, gastropod eggs, cystoids, crinoids, or
tunicates. The first comprehensive study was that of Stolley (1896), which
reviews all previous work. Stolley showed convincingly that cyclocrinitids
resembled dasycladacean green algae, and this idea soon gained wide accep-
tance in Europe. Pia (1920, 1927) reviewed the cyclocrinitids, and described
them as a dasycladacean tribe. Unfortunately, Stolley'S work had a very limited
distribution in North America, so it was largely overlooked; North American
workers commonly regarded cyclocrinitids as aberrant sponges, rather than as
algae. However, J.H. Johnson and Konishi (1959) and J.H. Johnson (1961)
reviewed the cyclocrinitids and placed them with dasycladaceans. The second
I Dept. of Earth & Planetary Sciences, The Johns Hopkins University, Baltimore. MD 21218. USA
Cyclocrinitids 115
comprehensive study was the monograph of Nitecki (1970). This work is the
foundation of modern research; it has clarified much of the confusion regarding
the morphology, preservation, and taxonomy of cyclocrinitids. The valuable
annotated bibliography of Nitecki et al. (1987) lists all works on cyclocrinitids
to 1980.
The cyclocrinitids are now widely accepted as an extinct dasycladacean
tribe (Beadle 1988), and that interpretation is followed here. However, Nitecki
(1986) suggested that they were problematic a lgae related to receptaculitids.
1 Morphology
Cyclocrinitids had a centra l main axis, from which lateral branches radiated
outward in all directions (Fig. I b). The lateral branches expanded at their distal
tips to form bulbous lateral heads, which were more or less hexagonal in outline.
The lateral heads were tightly packed together to form a faceted outer cortex.
The living dasycladaceans Bornetella Munier-Chalmas (Fig. la) and Neomeris
Lamouroux have very similar morphologies (Stolley 1896; Nitecki 1970; Ni-
tecki and Johnson 1978; Beadle 1988). The resemblances are apparent even at
early growth stages (Elliott 1972). It is possible to reconstruct some cyclocrinitids
in considera ble detail (Pia 1927; Osgood and Fischer 1960; N itecki and Johnson
1978). The standard anatomical terminology is that of Nitecki (1970).
Cyclocrinitid thalli vary from claviform (Fig. 2a) to spherical (Fig. 3b) in
shape. It is often difficult to determine their original shapes, as they were very
vulnerable to compaction (Nitecki and Johnson 1978). The unusual 'bilobate'
thalli described by Lee and Caldwell (1977) may well have been compacted.
Specimens in shales are often completely flattened (Fig. 3c). Cyclocrinitid
Fig. la,b. Schematic longitudinal cross-sec-

tions through a Recent dasycladacean and a
Silurian cyclocrinitid. CA Central axis ; 1 LB
primary lateral branch; 2LB secondary late-
ral branch; LH lateral head ; Ggametangia.
a Recent Bornetella capitata (Harvey). Thal-
lus size = \.0- \.5 cm. After Valet [1968, PI.
10(7), Fig. 5) ; b Lower Silurian Cyclocrinites
dactioloides (Owen). This species had no
secodary branches, but other cyclocrinitids
did . Position of gametangia unknown. Thal-
lus size = 1-4 cm
116 S.c. Beadle
Fig.2a-d. Middle Ordovician Cyciocriniles py riformis (Bassler). a Holotype. Note clavi form shape.
The preserved calcareous material encrusted the proximal surfaces of the lateral heads. USNM
56625. Chambersburg Limestone; Strasburg, Virginia. x1.75 , bSpecimens from the 'Niduliles Bed'.
Thalli are filled with sparry calcite. USN M 97551. Same locality as a x I, c Horizontal cross-section,
showing central main axis, lateral branches. a nd lateral heads. USNM 111806. Ward Cove
Formation; Staffordsville, Virginia. x3.4, dEnlargement of calcareous layer. The lateral heads were
generally hexagonal, and tended to show hexagonal closest packing. USN M 111789. Chambersburg
Limestone; Hagerstown, Ma ryland. x6.25
Cyclocrinitids 117
Fig.3a-d. a-b Lower Silurian Cyclocrinites dactioloides (Owen). Farmers Creek Member, Hop-
kinton Dolomite; Delaware Country, Iowa. a Longitudinal cross-section, showing proximal sur-
faces of lateral heads and remnant of central axis. FMNH UC 59064. x 1.5, b Internal mould. Note
spherical shape. FMNH UC 59458. x1.25, cod Lower Silurian Cyclocrinites favus (Salter). Photo-
graphs courteously provided by Dr. Atle M0rk. c Flattened thallus in shale. PMO 105837. Solvik
Formation ; Bj0rk0ya, Norway. x 1.6. d Cyclocrinitid thalli in shale, probably swept together by
currents. PMO 105839. Solvik Formation; AvI0S, Norway. xO.25
118 S.c. Beadle
preservation is quite variable, even within a single species (Nitecki 1970). Many
specimens have complex diagenetic histories (Osgood and Fischer 1960; Nitecki
and Johnson 1978; M0rk and Worsley 1980). Preservational differences have
often confused taxonomists.
Most cyclocrinitids are only 1-4 cm in length; the largest approach 6 cm.
They probably had a siphonous cellular organization like other dasycladaceans,
and this restricted them to small sizes (Beadle 1988). Certain forms, notably
Apidium, were only 1-10 mm in length (Kummerow 1937; Lee and Caldwell
1977; R.E. Johnson and Sheehan 1985). Some small thalli probably represent
immature individuals (Elliott 1972; Beadle and Johnson 1986, p. 597).
The main axis was rooted to a solid substrate; attachment scars and
'anchoring rhizoids' have been reported (Nitecki 1970; Lee and Caldwell 1977).
The main axis and lateral branches are only rarely preserved; they were
normally very thin, as in other dasycladaceans (Figs. 2c, 3a). In some forms,
nota bly Coelosphaeridium, the axes and branches were thicker, but the increased
width was generally balanced by decreased length. In some, but not all, species,
the lateral branches divided to form secondary branches. However, there was
generally less branching than in Recent dasycladaceans, and so cyclocrinitids
typically had fewer, larger lateral heads (Beadle 1988). The lateral heads are
usually 1-3 mm in width; they increased in size as the thallus grew (Beadle and
Johnson 1986).
The arrangement of the lateral branches is not well established. Pia (1920)
regarded cyclocrinitids as 'aspondylous', with branches randomly arranged.
However, Nitecki (1970) found that Cyclocrinites welleri Nitecki had lateral
branches in whorls. The lateral heads of many species are arranged in a
strikingly regular manner.
Cyc10crinitids were encrusted by surficial deposits of calcium carbonate.
The calcification typically encrusted the proximal or distal surfaces ofthe lateral
heads; these are often the only structures preserved (Fig. 2d). The calcification
was probably aragonitic (Osgood and Fischer 1960; Beadle 1988). Some species
apparently had calcareous external membranes (Nitecki 1970, Fig. 19). The
problematic Amia hexagona Fry may well represent an uncalcified cyc10crinitid
(Fry 1983).
Certain forms had small, radially-arranged ridges or knobs on the proximal
or distal surfaces of the lateral heads (Fig. 4). This unusual ornamentation is
especially variable and well preserved among the Ordovician species of the
Baltic area (Stolley 1896, 1898; N eben and Krueger 1973, 1979). This feature has
not been reported among living dasycladaceans, and so it merits further study.
No definitive cyclocrinitid reproductive structures are known. The 'spo-
rangia' of Currie and Edwards (1942) appear to be inorganic crystals (Elliott
1972). The ovoid 'gametangia' of Osgood and Fischer (1960) resemble peloids
of faecal pellet origin. Nitecki (1972a) reported 'gametocysts' in Cyclocrinites
dactioloides (Owen); these structures apparently occur only in dolomitized
specimens, and may be diagenetic mineral growths. Cyclocrinitids may have
been 'endosporate', with uncalcified reproductive structures that were formed
within the main axis (Pia 1920; Elliott 1972, p. 368).
Cyclocrinitids 119
Fig.4. Reconstruction of Cyciocriniles porosus

Stolley, from the Ordovician of the Baltic area .
The distal surfaces of the lateral heads had com-
plex ornamentation, termed 'Skulptur' by Pia
(1927). Thallus size = 8-10 mm. From Pia
(1927 , Fig. 43b)
2 Geographic Distribution
2.1 North America
Cyclocrinitids occur widely in the USA, Canada, and Greenland. N itecki (1970)
reviews all previous work. Lee and Caldwell (1977) list the 'Arctic Ordovician'
localities. Beadle and Johnson (1986) review all Silurian localities. Other recent
studies include Nitecki (1972b), Nitecki and Johnson (1978), M.E. Johnson and
Campbell (1980), R.E. Johnson and Sheehan (1985), and Frey (1987).
2.2 Europe
Cyclocrinitids are common in southern Norway (St0rmer 1953; H0eg 1961;

Skjeseth 1963 ; M0rk and Worsley 1980; Baarli 1987) and in northern Estonia
(Raymond 1916; Bekker 1924; Roomusoks 1970). They probably also occur in
submarine beds in the northern Baltic Sea (St0rmer 1953, p. 136). Cyclocrinitids
from these deposits have been widely distributed in glacial erratics (Kiesow
1894, 1899; Stolley 1896, 1898; Hucke and Voigt 1967; Neben and Krueger
1973, 1979). Cyclocrinitids occur at a few localities in Scotland and Wales
(Currie and Edwards 1942; Elliott 1972); an additional report, by Spjeldnres
(1955), remains unconfirmed . Aproblematic Shropshire fossil, Sphaerospongia
hospitalis Salter, resembles a cyclocrinitid (Salter 1873). The many Ordovician
species of Europe need review and revision. Silurian forms are reviewed by
Beadle and Johnson (1986). N itecki et al. (1987) list many additional references.
120 S.c. Beadle
2.3 Asia
Ordovician cyclocrinitids occur at several localities in central and southern Asia,

including eastern Kazakhstan (Gnilovskaya 1972), northern India (Reed 1912),
and western China (Mu 1982a,b). The Asian forms most closely resemble those
found in Europe.
3 Stratigraphic Distribution
The oldest reported cyclocrinitid is Cyclocrinites welleri Nitecki, from the lower
middle Ordovician of California (Nitecki 1970). Cyclocrinitids were most
abundant and diverse during the Caradoc, when they were widely distributed in
North America, northern Europe, and central Asia. They are less common in
Ashgill deposits, and they declined throughout the Llandovery (Beadle and
Johnson 1986). They were apparently extinct by the end of Llandovery time.
The radiation and decline of the cyclocrinitids closely paralleled that of
other dasycladaceans. The decline of these warm-water algae at the end of the
Ordovician may reflect the simultaneous cooling and glaciation (Beadle 1988).
4 Palaeoecology
Cyclocrinitid palaeoecology is discussed by N itecki (1970), N itecki and Johnson

(1978), M0rk and Worsley (1980), Beadle and Johnson (1986), and Beadle
(1988). Cyclocrinitids were ecologically very similar to other dasycladaceans.
Cyclocrinitids typically occur with normal marine invertebrates. However,
they have also been found in high-salinity restricted-water environments
(M.E. Johnson and Campbell 1980). Palaeogeographic reconstructions show
that most, and perhaps all, cyclocrinitids lived within 30° of the palaeo-
equator (Beadle and Johnson 1986; Beadle 1988). They are usually found in
carbonate rocks, but they are also common in certain siltstones and shales. Cy-
clocrinitids probably colonized soft substrates by attaching themselves to
small solid objects; living dasycladaceans are often found on soft bottoms
attached to pebbles, shells, or coral fragments.
Cyclocrinitid thalli were relatively fragile; they were largely restricted to
quiet-water environments, either below wave-base or in protected lagoons
(Nitecki 1970; Beadle and Johnson 1986). Cyclocrinitids were probably very
vulnerable to strong currents. Comparison with the anatomically-similar
Recent dasycladacean Bornetella sphaerica (Zanardini) suggests that their thalli
may have been easily torn away from their holdfasts (Nitecki and Johnson
1978). Furthermore, they probably grew on small objects which were readily
transported; living dasycladaceans in Texan lagoons are often moved about in
this manner (Conover 1964, p. 14). Thus, large accumula tions of cyclocrinitids
may have been formed by rare storm-generated currents, which penetrated into
normally quiet waters and swept the thalli together (Nitecki and Johnson 1978).
Cyc10crinitids 121
Cyclocrinitid thalli often show evidence oflocal transport (Osgood and Fischer
1960; Mr6rk and Worsley 1980).
Cyclocrinitids were most common at relatively shallow depths, below
wave-base but within the photic zone. Comparison with living dasycladaceans
suggests that they lived at depths ofless than 100 m (Beadle and Johnson 1986).
Silurian benthic communities are commonly regarded as depth-zoned; cy-
clocrinitids are most common in Cryptothyrella, Pentamerus, and Stricklandia
communities, or their equivalents (Beadle and Johnson 1986). They have also
been found in very shallow protected-water environments (M.E. Johnson and
Campbell 1980). Cyclocrinitids rarely ranged below the Stricklandia com-
munity, but there are a few reports of them in Clorinda (or equivalent) com-
munities (Cocks and Toghilll973; Baarli 1987).
Some Silurian cyclocrinitids show apparent ecophenotypic variation:
certain populations have significantly smaller thalli than stratigraphically ad-
jacent populations. This variation most likely represents differences in ambient
light intensity, and it has been used to indicate relative palaeodepths (Beadle
and Johnson 1986). This approach is of interest to palaeoceanographers, since
it can be used to determine the bathymetric relationships of the associated
invertebrate communities (M.E. Johnson 1987, Fig. 2).
5 Stratigraphic Significance
The distribution of cyclocrinitids is sporadic. However, they are often extremely

abundant locally, and they may be significant rock-builders. Some units could
well be termed 'cyclocrinitid limestones'; for instance, their thalli comprise
25-40% of the rock in parts of the Thumb Mountain Formation of Cornwallis
Island, Canada (Lee and Caldwell 1977). They are also very common in certain
shaly units, such as the Solvik Formation of Norway (Fig. 3d) and the Gasworks
Mudstone of Wales (Beadle and Johnson 1986).
Cyclocrinitids are often used as local guide fossils because of their abun-
dance and distinctive appearance. Several rock units have been informally
named for them, including the 'Cyclocrinites Beds' (or 'Cerionites Beds') of
Iowa (M.E. Johnson 1983), the 'Nidulites Beds' of Virginia (Fig. 2b) (Cooper
and Cooper 1946), the 'Coelosphaeridium Beds' and 'Cyclocrinus Beds' of
Norway (Skjeseth 1963), and the 'Coelosphaeridium Zone' of Estonia (Bekker
1924). Cyclocrinitid-bearing glacial erratics in northern Europe have been
described as 'Coelosphaeridiengesteine' and 'Cyclocrinus-Kalke' (Kiesow
1894; Hucke and Voigt 1967, Table 2).
Some species may be useful for interregional correlation. Cyclocrinites
pyriformis (Bassler), from the Champlainian of the Appalachians, may be
synonymous with Mastopora parva (Nicholson and Etheridge) from the Ca-
radoc of Scotland (Nitecki 1970). Spjeldnres (1955) claimed that Coelosphae-
ridium cyclocrinophilum Roemer occurred in middle Caradoc strata in Wales,
Norway, and Estonia. In both Norway and Estonia, these Coelosphaeridium-
bearing beds are directly overlain by Cyclocrinites-bearing beds (Raymond
122 S.C. 'Beadle
1916, p. 245; St0rmer 1953, p. 136). Cyclocrinites favus (Salter) characterizes

lower Llandovery strata in Norway, Scotland, and Wales (Beadle and Johnson
1986); it may also be synonymous with the contemporary North American
species C. gregarius (Billings).
6 Further Research
Ideally, all cyclocrinitids should be well-defined taxonomically, precisely dated

with respect to a global standard, and placed into a palaeoecological context,
along with the associated invertebrates. Silurian cyclocrinitids approach this
ideal. However, considerable work is needed on the Ordovician forms, which
are much more abundant, widespread, and diverse.
Nitecki (1970) has recently revised all North American species; a similar
review of the European and Asian forms is needed. Many ofthese 'species' may
represent only different states of preservation. Furthermore, many are based
upon specimens from glacial erratics, so their stratigraphic ranges are somewhat
uncertain. New descriptions would be welcome, especially for the Estonian and
Norwegian forms, which are quite important stratigraphically.
Cyclocrinitids may prove useful in reconstructing Ordovician palaeogeo-
graphy and palaeoecology. Cyclocrinitids appear to be reliable indicators of
low palaeolatitudes. They can also be used as relative depth indicators, and
this could help to establish the bathymetric relationships of Ordovician
palaeocommunities.
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Chapter 7
Dasycladalean Algae of the Palaeozoic and Mesozoic
G.F. ELLIOTT!
Abstract
General compilative publications with good bibliographies for the Dasycladesofthe Palaeozoic and
Mesozoic are mentioned, followed by brief discussion of published literature on classification and
changing trends in the characters to be considered for this, and their evaluation. The absence of
available dasycladalean evidence between the Upper Permian and Middle Trias is discussed, and
suggestions made for possible future research to remedy this. The influence of good and bad
preservation on interpretation is illustrated. Examples are given of the usefulness of dasycladaleans
in palaeoecology and palaeogeography. The classification of Deloffre is discussed.
1 Introduction
Dasyclads (the general noun for members of the chlorophyte algal order
Dasycladales) are well represented in the Palaeozoic and Mesozoic. They
appear to have been largely confined to shallow marine warm clearwater facies,
as interpreted by comparison with the habitats ofliving survivors. Nevertheless,
they are common enough at many levels to be useful microfossils for strati-
graphic purposes, and they have been the subject of very numerous studies.
Inevitably the dasyclads of the different parts of this long time span have
received somewhat unequal treatment. Best served are the Jurassic and Cre-
taceous, where the monograph ofBassoullet et al. (1978) offers a ready entry and
detailed guide to dasyclads of these periods. The Trias, in great part the subject
of Julius Pia's earlier pioneer studies (1920, 1927) is not now served by a
comparable contemporary work, though some regional studies such as that of
Bystricky (1964) on the dasyclad limestones of eastern Czechoslovakia, go some
way towards this, and Fliigel's summarized study on the diversity and envi-
ronments of Permian and Triassic Dasycladacean Algae, though not a taxo-
nomic-descriptive study, contains a wealth of useful information and interpre-
tation (FliigelI985). For the Palaeozoic, the summary work ofRoux (1985) deals
with all algae of this age, thus including dasyclads. A mine of information, it can
be supplemented by other studies; one thinks especially of the long series of
regional papers by Mamet and his collaborators, largely but not exclusively on
the Upper Palaeozoic. Finally, the published results of the Groupe fran~ais
d'etude des algues fossiles (Bassoullet et al. 1975), on the different values of
various dasyclad structures in classification, are leading to a revised classifi-
'Department of Palaeontology, British Museum (Natural History), Cromwell Road, London SW7
5BD, UK
126 G.F. Elliott
cation long overdue. It is noteworthy that dasyclads, from the limited nature of
the fossilized remains, do not usually lend themselves to an easy evaluation of
phylogeny. Valet's study ofliving dasyclads (1968, 1969) shows how much is not
normally preserved in the fossil state. The Groupe Frant;ais attempted to group
all dasyclads in tribes, on the basis of their morphological similarities (Bassoullet
et al. 1979). It is interesting that when the time-ranges are compared, a pro-
gression in time, admittedly with much overlap, is readily observed (Bassoullet
et al. 1979, p.431); i.e. the successive range-terminations progress from older to
younger.
With all this in mind, we know that dasyclads appeared in the Lower
Palaeozoic, proliferated in the Permian, and again in the Middle-Upper
Triassic, Upper Jurassic, Lower Cretaceous, surviving in facies-abundance to
the Palaeogene, after which they declined to their present-day "relict-flora"
position. Throughout, they appear to have favoured the same general envi-
ronment, so far as we can tell: shallow warm clear-water marine conditions,
usually coastal, often lagoonal. Beginning as aspondyl (the branches set non-
verticillate on the central axial body), they proceed to the verticillate branch
arrangement, sometimes leading to adult terminal reproductive discs, or other
structures. The branches themselves show an astonishing number of patterns,
which are invaluable taxonomically. Guven'S (1979) has carefully analyzed
those with the metaspondyl type of branch, to show the affinities between the
Palaeozoic and Triassic genera to indicate possible relationships. The basic
trends towards this kind of evolution may have occurred separately in different
stocks and at different times, as recognized by Kochansky-Devide and Gusic
(1971) for dasyclad structures generally. A good example is the Ordovician
Archaeobatophora (Nitecki 1976), with branches comparable to those of the
living Batophora, although contemporaneous with much more primitive gen-
era. Our colleagues the French Group, in a major review of dasyclad clas-
sification characters, inclined towards a morphological-structure taxonomy
(Bassoullet et al. 1975), replacing the older ideas of Julius Pia, who placed a
principal emphasis on the positions of the reproductive structures, whether
within the stem-cell, in swollen primary branches, or in special structures borne
on these branches. Pia wrote before much later work, and the more recent
changes in classification were long overdue. Indeed, there have been indications
recently of different interpretation of reproduction structures, as in Skompski's
(1984) account of the growth and function of Kulikia in the Carboniferous.
Obviously, there is a great deal still to be discovered. What can be done,
however, is to suggest what is desirable in future research, and this fits with the
title wording of this chapter: Palaeozoic and Mesozoic dasyclads. How do the
two fit together?
2 The Palaeozoic/Mesozoic Junction
It is generally accepted that the level at which top Permian underlies bottom
Trias is the record of a time of profound changes for life on Earth. Many and
various explanations, of varying degrees of credibility, have been advanced to
Dasycladalean Algae of the Palaeozoic and Mesozoic 127
account for this. So far as dasyclads are concerned, a rich flora with Mizzia as its
key plant ceases abruptly as the Permian stops. The detailed summarized study
of FlUgel (1985), whilst not primarily on the component taxa of the Upper
Permian/Triassic dasyclad floras, contains much useful information on them.
Above the Permian there comes in most places a lower Triassic of largely
arenaceous rocks with molluscs, but apparently without dasyclads. Then, in the
right facies of the Middle Triassic limestones, dasyclads appear again, as
diplopores in incredible abundance (they were the Halimeda of their time,
ecologically speaking). Whilst it is true that some dip lopores have been de-
scribed from the Upper Palaeozoic, and their possible ancestry and relation-
ships evaluated, the contrast between the two floras is striking. One asks, what
were the dasyclads doing, and where?
This is, I think, a very real gap in our knowledge. To fill it, one needs to find
a Lower Trias in algal-limestone facies. Two possibilities can be suggested. One
is in the Trucial Oman area of Arabia. About 25 years ago, the now defunct Iraq
Petroleum Group, by whom I was then employed, prospected there and carried
out a basic geological survey of the local succession. Both palaeontologically
determined Permian and Triassic occur, in a long succession oflimestones. The
key formation for our problem is the Ghail Limestone, ca. 600 m thick (Hudson
1960). Although I described an alga (not a dasyclad) from the Upper Ghail
(Elliott 1964), the lower part was not intensively sampled, although noted as
algal. Moreover, such samples as were taken never reached me, and it was too
early in my algal career to realize fully the possible importance of these. The
other area, which I have never worked on, is the Western Himalayan/Central
Asian region, where the impact ofthe drifting Indian plate on the Asian mass has
possibly preserved a similar rock.
If one takes the conventional, unexciting but likely view that late-Permian
shallow-marine life suffered from a geologically sudden dearth of suitable
shelf-sea facies, and that this continued in the Lower Trias, then occasional
dasyclad survivors had a wonderful opportunity to explode evolutionary-wise
when good times returned again, and this they seem to have done. If a
dasyclad-yielding Lower Trias exists, then it may hold the key to the dasyclad
transition from Upper Permian to Middle Trias, and it may profoundly affect
thinking regarding the title of this chapter. One would then see which genera
survived into Triassic time, which of them did not flourish, and the nature of
these early Triassic dasyclads in relation to the later floras known to us.
By comparison, the change from Mesozoic to Cenozoic was apparently
much less abrupt for algae. Such dasyclad genera as Cymopolia and Neomeris,
both surviving today, are well represented in both Upper Cretaceous and
Palaeocene.
3 Some Problems
Apart from this major question, the study of new dasyclads (and of better
preserved material of known genera), builds up our knowledge of these fas-
cinating plants. That it sets us new problems as well is part of the attraction of
128 G.F. Elliott
these fossils. Skompski (1987), working on exceptionally well-preserved ma-

terials, has re-examined and discussed the problematic organisms known as
Palaeoberessellae, which have been referred by different workers to the dasy-
clad algae, to the Foraminiferidae and to the extinct Ischyrospongia
(summarized in Skompski 1987). He concludes that they are dasyclads, and
discusses their position within the order Dasycladales. Preservation after
diagenesis plays a great part in the appearance of certain structures, and the
necessity for allowing for this has not always been obvious to investigators. We
look forward to extending our knowledge from well-preserved known taxa, as
well as from new taxa.
It will be seen from the above that our knowledge of the Dasyclads of the
Palaeozoic and Mesozoic is in an exceptional state of flux. Much is unknown,
with insufficient certainty to avoid controversy in an overall view. But the
progress of new researchers should, we hope, remedy this.
4 ~aeogeography,~aeoecology
As mentioned above, dasyclads are useful fossils for palaeoecology (and

palaeogeography). Their occurrence globally in Mesozoic Tethyan sediments
has been used by Elliott, not merely to show the probable origins oftheir current
distribution (Elliott 1981), but also their occurrence and distribution in seas
marginal to the Tethys in what is now northwestern Europe (Elliott 1977, 1984,
1986). In the Palaeozoic, Poncet (1987) has studied the palaeogeography of
Vermipore/la in the Middle and Upper Ordovician.
5 Deloffre's Classification, 1988
The dasyclad classification of Deloffre (1988) is an important landmark in

dasyclad studies. He divided the Dasycladales into five families: Beresellaceae,
Seletonellaceae, Diploporaceae, Dasycladaceae and Acetabulariaceae, cover-
ing about 180 genera arranged in 30 tribes. The two new families are welcome,
as recognizing the special characters of their component genera. Beresellaceae
form a group somewhat set apart from other dasyclads, and the character given
as distinctive for the family is "Tubular algae with or without septation". It will
be interesting to see if this definition is widely accepted or not. In contrast, the
criteria for separation of Diploporaceae from Dasycladaceae - metaspondyle
thallus, and euspondyle thallus with verticils similar, respectively - is in accord
with recent acceptance of primary branch form as significant in classification. It
should be borne in mind that whilst logically one uses the same uniform criteria
for classification in different related taxa, the extent of evolution varies in these
characters in different groups, necessitating some modification if classification
is to reflect the evolution that has taken place. However. the separation of
Diploporaceae from Dasycladaceae is a useful step towards the understanding
of dasyclad evolution of the Upper Palaeozoic/ Lower Mesozoic. The characters
Dasycladalean Algae of the Palaeozoic and Mesozoic 129
used for classification are selected-morphological, and follow naturally from the
work of the Groupe Fran'Sais.
Many students ofthe dasyclads will find genera with whose tribal allocation
by Deloffre they will not agree; they will from their personal knowledge place
them elsewhere, or at any rate query them. This does not detract from the value
of his attempt, in view of our incomplete knowledge of so many genera. He
recognizes the possibility of an eventual phylogenetic classification: the most
likely family to show this is the Acetabulariaceae, whose component genera may
well be of different ancestries, so making the family polyphyletic, but we do not
know this for a fact.
It is usual for Dasyc/adus, which gives its name to the Order, Family and
subtribe, to give its name to the appropriate tribe also, rather than TriploporeUa,
even if this is a well-calcified fossil genus.
Deloffre's work is a useful reference paper for all workers on dasyclads, and
should be a useful tool for future efforts as knowledge grows.
References
Bassoullet JP, Bernier P, Deloffre R, Genot P, Jaffrezo M, Poignant AF, Segonzac G (1975)
Refiexions sur la systematique des Dasycladales fossiles. Etude critique de la terminologie et
importance relative des criteres de classification. Geobios 8, 4:259~290
Bassoullet JP, Bernier P, Conrad MA, Deloffre R, Jaffrezo M (1978) Les algues Dasycladales du
Jurassique et du Cretace. Geobios Mem Spec 2:330 pp
Bassoullet JP, Bernier P, Deloffre R, Genot P, Jaffrezo M, Vachard D (1979) Essai de classification
des Dasycladales en tribus. Bull Cent Rech Explor Prod Elf-Aquitaine 3, 2:429~442
Bystricky J (1964) Siovensky kras Stratigrafia a Dasycladaceae mesozoika Siovenskeho krasu.
Ustredny Ustav Geol, Bratislava, 204 pp
Deloffre R (1988) Nouvelle taxonomie des algues dasycladales. Bull Cent Rech Explor Prod
Elf-Aquitaine 12:165~217
Elliott GF (1964) Zonotrichites (calcareous algae) from the Arabian Triassic. Eclogia Geol Helv 57
2:567~570
Elliott GF (1977) Inferred isocrymal distribution of Jurassic dasycladacean Algae in Europe, north
Africa and southwestern Asia. J Geol Soc London 133:363~373
Elliott GF (1981) The Tethyan dispersal of some chlorophyte algae subsequent to the Palaeozoic.
Palaeogeogr Palaeoclimatol PalaeoecoI32:341~358
Elliott GF (1984) Climatic tolerance in some aragonitic green algae of the post-Palaeozoic.
Palaeogeogr Palaeoclimatol Palaeoecol48: 163~ 169
Elliott GF (1986) English Jurassic dasyclads: seaweed indicators of marine climate. Geol Today 2,
1:20~23
Fliigel E (1985) Diversity and environments of Permian and Triassic dasycladacean Algae. In:
Springer, Berlin Heidelberg New York Tokyo, pp 344~351
GiivenlS T (1979) Dasycladacees metaspondyles du Paleozoique superieur et du Trias. Bull Cent
Rech Explor Prod Elf-Aquitaine 3, 2:625~637
Hudson RGS (1960) The Permian and Trias of the Oman Peninsula, Arabia. Geol Mag 97:299~308
Kochansky-Devide V, GuS'ic I (1971) Evolutionstendenzen der Dasycladaceen mit besonderer
Beriicksichtigung neuer Fiinde in Jugoslavien. Palaontol Z 45, I~2:82~91
Nitecki MH (1976) Ordovician Batophoreae (Dasycladales) from Michigan. Fieldiana Geol 35,
4:29~40
Pia J (1920) Die Siphoneae Verticillatae yom Karbon bis zur Kreide. Abh Zool Bot Ges Wien II,
2:263 pp
130 O.F. Elliott: Dasycladalean Algae of the Palaeozoic and Mesozoic
Pia J (1927) Thallophyta I. Abteilung. In: Hirmer M (ed) Handbuch der Paliiobotanik. Oldenbourg
Munich Berlin
Poncet J (1987) Paleobiogeographie du genre Vermiporella (a!gue verte calcaire) a l'Ordovicien
moyen et superieur. Ann Soc Oeol Nord 106 3:279-283
Roux A (1985) Introduction a l'etude des algues fossiles paleozoiques (de la bacterie a la tectonique
des plaques). Bull Cent Rech Explor Prod Elf-Aquitaine 9, 2:465-699
Skompski S (1984) The functional morphology of the Carboniferous dasycladacean genus Kulikia.
N Jahrb Oeol Paliiontol Mh 7:427-436
Skompski S (1987) The dasycladacean nature oflate Paleozoic palaeoberesellid algae. Act Oeol Pol
37,1-2:21-31
Valet 0 (1968) Contribution a l'etude des Dasycladales. l. Morphogenese. N Hedwigia 16:21-:82
ViLlet 0 (1969) Contribution a l'etude des Dasycladales. 2. Cytologie et reproduction. 3. Revision
systematigue. N Hedwigia 17:551-644
Chapter 8
Cenozoic and Recent Dasycladales
Abstract
Cenozoic Dasycladales include about 43 genera and 200 species. The number of living genera is
reduced to 8, with about 40 species.
Calcification encloses sterile and fertile organs to a varying degree, according to genera,
species, and individuals. Consequently, preservation offossils mainly depends on the extent of the
initial calcification around the thallus.
Main characters commonly used to define taxa include the type and the position of the
reproductive organs, absence or presence of articulation, and division of branches.
A large number of species, belonging to the genera Neomeris, Cymopolia, and Acicularia, is
known in Palaeogene sediments. The richest cenozoic assemblages have been found in Sardinia
(Palaeocene) and in the Paris basin (Palaeocene and Eocene).
Living representatives have a discontinuous geographical distribution. They are confined to
shallow warm marine or brackish waters.
1 Morphology
The thallus of Cenozoic and living Dasycladales (Table 1) is usually cylindrical

or club-shaped, undivided or dichotomously branched. Other species are
spherical or appear like a disc at the top of an erect axis. In some genera, the
thallus is composed of alternate calcified segments and uncalcified joints. The
whole plant is attached to the substrate by means of un calcified rhizoids that are
never preserved in fossils.
Primary branches are arranged in regular whorls along the tubular axis
(Figs. 3,10-11). Most Cenozoic and Recent species bear two orders of branches :
each primary branch produces one (M ontiella) to eight (Bornetella, Cymopo/ia)
second order sterile branches. Dilated tips of primaries (Zittelina) or second-
aries (Cymopolia, Neomeris, Bornetella) may form a faceted outer surface (Fig.
5). Some genera bear more than two orders of branches, exceptionally seven
(Table 2).
In Dasycladaceae, fertile organs are spherical or ovoid vesicles usually
located at the top or on the sides of primary sterile branches. They are considered
as modified branches, assigned to the reproductive function. Each ampulla
contains one to several cysts or produces gametes directly (Table 3).
In Acetabulariaceae, fertile ampullae are elongated and radially arranged
in specialized verticils. Many cysts occur inside reproductive chambers (Tables
4-6) (Valet 1969; Tappan 1980).
1 Laboratoire de Biogeologie et Biostratigraphie, Universite de Nantes. 44072 Nantes Cedex, France

132 P. Genot
Table 1. Shape of thallus in Cenozoic and Recent Dasycladaceae
r\
I
(7 ~
Q Vb)J
spherical
c
club - shaped to cylindrical
non - divided
~d V
Idivided articulated
.... e
Photographs Fig.9 Figs .7-8 Figs.1-2 Figs.5-6

Dasycladus
RECENT Bornetella Cymopol ia
<t: ~
o:::(/)
Neomeris
Q)
We.
~
Dactylopora Dissocladella Belzungia
z§ u
~
0
Jodotello Broeckella
w~ N
Neomeris Cymopol ia
~ 0
<.D z:
W
u Trinocladus Uteria
Zittelina
- . - - - .. -.-- .. - -------.- _._--------

----------------..-
Fig. 1. Neomeris annula/a Dickie, Recent. Rose Island. Bahamas. x7.5. External view of the thallus.
Fig. 2. Neomeris [ragilis (Defrance), Lutetian, Grignon, Paris Basin. x35. Transversal section
showing axial calcification Fig. 3. Neomeris annu/a/a Dickie. Recent. Rose Island. Bahamas. x35.
Central axis (righ/), primary branches and fertile ampullae (Ie/i) Fig. 4. Neomeris sp .. Lutetian.
Campbon, Brittany. x95. Calcification offertile ampullae Fig. 5. Crmopolia barbata (Linne). Recent,
Andros Island. Bahamas. x25. Segments of the thallus Fig. 6. Cvmopolia sp .. Stampian. Ormoy-
la-Riviere, Paris Basin. x40. Isolated segment Fig. 7. Borne/ella oligospora Solms-Laubach. Recent.
Cauda. Vietnam. x6. External view of the club-shaped thallus Fig. 8. Daur/opora (ylindracea
Lamarck. Auversian. Le Faye!. Paris Basin. x II. External view of the club-shaped thallus Fig. 9.
Borne/ella sphaerica (Zanardini). Recent. Oahu. Hawaii. x7. Central axis. branches and fertile
ampullae (dry specimen) Figs. 10-11. Neomeris auversiensis (Marellet). Auversian. Auvers- sur-
Oise. Paris Basin. Calcification of the central axis at the top (Fig. 10. xllO) and at the base (Fig. II
x190) of the thallus
Cenozoic and Recent Oasyc1adales 133
134 P. Genot
t-<b k
Table 2. Division of branches in Cenozoic and Recent Dasycladaceae
a
I---- C ~d :f
<
<
':
no division 1 division 2 divisions more(uD to 6)
Photogrophs Flg.12 Figs.13,16,17 Fi g.14
Bornete II a Dasycladus
RECENT CymODol ia BatoDhora
c:( ~
cr::V)
-
Q)
Neome ris
Dacty loDOra Broeckella Tr inac ladus Belzun91a
-
We.
2:6 u Zl ttellna CymODolla Thyr sODore II a
lJ..J ~ 0
~~
N
0
z:
Dissoc ladella
W
u Jodotel la
Neomeris
Fig. 12. Zillelina dactyloporoides (Morellet), Auversian, Baron, Paris Basin. x130. Primary branch
Fig. 13. Cymopolia elongata (Defrance), Lutetian, Coislin, Brittany. x250. Primary and secondary
branches Fig. 14. Belzungia terquemi Morellet, Lutetian, Campbon, Brittany. x220. Numerous
divisions of the branches Fig. 15. Bornetella nilida (Harvey), Recent, Palawan, Philippines. x170.
External view of the upper faces of two secondary branches Fig. 16. Cymopo/ia barbata (Linne),
Recent, Andros Island, Bahamas. x190. Primary branch, secondary branches and fertile ampulla
Fig. 17. Cymopo/ia zilleli Morellet, Auversian, Le Fayel, Paris Basin. x125. Primary branches,
secondary branches and fertile ampullae Fig. 18. Neomeris larvarioides (Morellet), Lutetian, Le
Bois-Gouet, Brittany. xiIO. Primary branch, secondary branches (partially preserved) and fertile
ampulla Fig. 19. Neomeris herouvalensis Steinmann, Cuisian, Herouval, Paris Basin. x90. Secondllry
branches and fertile ampullae (primary branches are not calcified) Fig. 20. Balophora oerstedii
Agardh, Recent,. Key West, Florida. x45. Fertile ampulla Fig. 21. Parkerella sp., Montian, Mons,
Belgium. x 190. Fertile ampuUae arranged in pairs Fig. 22.Carpenlerella jonesi Morellet, Lutetian,
Campbon , Brittany. x250. Tuft of fertile ampullae
Cenozoic and Recent Dasycladales 135
136 P. Genot
Table 3. Fertile organs in Cenozoic and Recent Dasycladaceae

A - Fert il e roo
organs at
the top of
branches ~-----o r----@ r---~ rSQ..r:
. --. .
.. ... . .
~
. \,:#
0 b c d
one ampulla on each prima ry b rancn nume r?us
ampul ae
no cyst one Cyst cysts and Cysts
Phot ographs Figs.16-17 Flgs.3, 18, 19 Fig .20
<l:
c::: RECENT Cymopol la
Dasyclodus
Neomeri s Ch lorocladus
(Cyst Batophoro
lJ.J
:z
w Cymopol ia unknown
<.!l CENOZOIC Mont iel la in fossils)
B - Fertl Ie
organs around
the sides of ~<!.i~ro_ e
~ __cm___ f
branches
Numerous Cysts Tufts of organs wi thout cyst
Photographs Figs.9, 23 Figs .21-22
<l:
0::: RECENT Bornetel la
w
z:
lJ.J CENOZOIC Zi ttellna (inc luding Carpenterel la, Jodotella
<.!l Maupasia , Dlgitel la) Parkerella
Q.
C - Fertile
organs inside
branches
-
9
~~:
~_I' h o
~•. i
Ampul lae and Cysts Cysts Non-calcif ied organs ?
Photographs Flg .26 Fig.27 Fig .28
CENOZOIC GEN . Uterl a Broeckella Thyrsopore Iia
D - Particular
cases
Grouped or
scatte red cysts
around branches
....
~• • •
-"-"--j
Isolated
ampu l lae 0 k
Photographs Fig .24 Fig.25
CENOZOIC GEN. Dactylopora Terquemel la
Freder ico
Fig. 23-32. (see pp 138 for caption)

Cenozoic and Recent Oasycladales 137
\38 P. Genot
Fig. 23. Zillelina dacly/oporoides (Morellet), Auversian, Baron, Paris Basin. x115. Fertile ampullae
around a primary branch Fig. 24. Dacly/opora cylindracea Lamarck, Auversian, Le Fayel, Paris
Basin. x70. Group of cysts inside ovoid area of the wall Fig. 25. Terquemel/a dissimilis Morellet,
Lutetian, Campbon, Brittany. xIOO. Section of an isolated fertile ampulla Fig. 26. Uleria brocchii
Morellet, Montian, Mons, Belgium. x165. Section of two fertile ampullae showing the location of
cysts Fig. 27. Broeckel/a be/gica Morellet, Montian, Mons, Belgium . x90. Primary branch and short
secondary branches. Cysts are not preserved in this specimen Fig. 28. Thyrsoporel/a cancel/ala
Guembel, Lutetian, Fercourt, Paris Basin. x250. Division of branches inside the wall. Very wide
primary branch Fig. 29. C/ypeina digilala (Parker and Jones), Lutetian, Montjavoult, Paris Basin.
x 12. Exceptionally preserved specimen with numerous fertile verticils Fig. 30. C/ypeina sp., Lutetian,
Montjavoult, Paris Basin. x45. Terminal verticil Fig. 31. C/ypeina sp., Lutetian, Chambors, Paris
Basin. x50. Section ofa terminal verticil Fig. 32. Halicoryne morelleli(Pokorny), Sarmatian, Podivin,
Czechoslovakia. x70. Group of calcified cysts
...
Table 4. Shape of thallus in Cenozoic and Recent Acetabulariaceae
f\
-1
C - :"'E=:::>
J J.
-"
~
~ ~
>;
-1
a b c
umbrella-shaped cylindrica l to ovoid
Photographs Figs.33 , 40 Fig .29
<1:
c::
RECENT Acetabuiarla Hal icoryne
i.LJ
z
i.LJ Acicularia Clype!na Rostroporella
<.!) CENOZOIC Or! opore ll a Ha l !cQryne
Cenozoic and Recent Oasycladales 139
Table 5. Position ofverticils in Cenozoic and Recent Acetabulariaceae
Fertile (vf) and sterile (vs ) vertlclls Fertile

non alternate 01 ternate vertlclls
Fertile Fertile only
verticil verticlls known
f:: f::
at the top along the axis
l: vI
- ,-.;;
'"
0'~-9
vI
.,
0 b c d
Photographs Fl g.33 Flg.29 Flgs.37,40
«
ex: RECENT Acetabularla Hallcoryne
lLJ
Clypelna Aclcularla
CENOZOIC (Aclcularla) Rostroporella
:z
lLJ
<.!)
? Orioporella
Table 6. Fertile organs in Cenozoic and Recent Acetabulariaceae
...~
... ~
..
I.~.::: j ~
'"
·: ' ~b
.. -";;,;~
o~"~
.. _."c.'t1-W d
': :.- Y
0 c d
Ampul lae and Cysts Ampullae
without Cysts
Non - Calcified Cysts (non-calcified
calcified strongly I partially Cysts ?)
Photographs Flg .35 Figs .32, 36,39 FIgs .42,43 Fig.31
RECENT Acetabularla
Hallcoryne
CENOZOIC Acicularia Orioporella Clypeina
140 P. Genol
Cenozoic and Recent Dasycladales 141
When fertile organs are unknown in fossils, thick primary branches are
usually presumed to be reproductive structures.
In some particular cases the calcified wall may contain groups of cysts or
scattered cysts along the branches (Table 3D).
2 Calcification
Aragonitic calcification covers the outer layer of the organs. In fossil species,
aragonite may be diagenetically replaced by calcite.
Calcification encloses organs to a varying degree; for example, in Neomeris,
calcium carbonate impregnates different parts of the plant (Figs. 2-4, 1O-11):
almost the entire thallus in N. larvarioides, only reproductive organs in N.
pseudo-eruca (Genot 1987).
Some living Dasycladales are never calcified (Batophora).
3 Definition of Taxa
The main characters commonly used to define Cenozoic and Recent sub-
families, tribes, and genera are: type of reproductive organs (single or clustered
ampullae without cysts, or containing one or several cysts); position of re-
prod uctive organs; type of thallus (a bsence or presence of articula tion); division
of branches. Taxonomic importance ofthe shape of the thallus and aspect of the
outer surface is variable according to authors (specific or generic level).
4 Stratigraphical and Geographical Distributions
A few genera that arose during the Mesozoic (Acicularia, A croporella, Clypeina,
Cymopolia, Dissocladella, Neomeris, Pra turlon ella , Trinocladus and Triplopo-
rella) have Cenozoic representatives. Two of them are known in present seas:
...
Fig. 33. Acetabu/aria crenu/ata Lamouroux, Recent, Barranquilla, Colombia. x6. Axis and re-
productive cap Figs. 34-35. Acetabu/aria acetabulum (Lamouroux), Mediterranean coast, France.
Upper corona in the central part of the cap (Fig. 34, x40) and internal view of two rays showing the
cysts (Fig. 35, x140) Fig. 36. Acicu/aria munieri Morellet, Lutetian, Grignon, Paris Basin. Internal
view of a ray showing the location of the cysts Fig. 37. Acicu/aria cf. munieri Morellet, Lutetian,
Villiers-Saint-Frederic, Paris Basin. x50. External view of two rays Fig. 38. Acicu/aria pavantina
d'Archiac, Auversian, Le Guepelle, Paris Basin. x90. Transverse section of a ray showing the
location of cysts Fig. 39. Acetabu/aria schenckii Moebius, Recent, Key Largo, Florida. x 130. Group
of calcified cysts Figs. 40-42. Orioporella villallae Segonzac, Thanetian, Carla-de-Roquefort,
Pyrenees. External view ofa reproductive cap (Fig. 40. xl3,5). Longitudinal section (Fig.4l, x22,5).
Transverse sections of two rays (Fig. 42, x32,5). Fig. 43. Orioporella bonieri Morellet, Montian,
Mons, Belgium. x580. Detail of the internal view of a ray showing the location of a cyst
142 P. Genol
Cymopolia and Neomeris. About 25 genera are restricted to the Cenozoic, each
one including very few species (Deloffre and Genot 1982).
Acicularia, Cymopolia and Neomeris have the most numerous species,
particularly during Palaeocene and Middle Eocene (Tables 7-8).
The number of species greatly decreases after the Eocene. Some genera are
short-lived, such as Broeckella, which is widespread in the Tethyan Palaeocene.
Richest Cenozoic assemblages (Table 9) have been discovered in the
Palaeocene of Belgium (Mons basin), France (Aquitaine - Pyrenees area and
Paris basin), Italy (Sardinia), USSR (Ukraine), Yugoslavia (Slovenia), Cze-
choslovakia (Carpathian Mountains), Middle East (Iraq, Iran) and China
(Tibet), in the Eocene of France (Paris basin, Brittany, Cotentin), southern
England (Sussex), Hungary and China (Xizang), and in the Miocene of
Romania and Poland. .
Only eight genera are living in modern oceans and seas, including about 40
species, mainly of Neomeris and Acetabularia (Table 7). They have a discon-
tinuous geographical distribution in the Indo-Pacific, Atlantic (tropical Ame-
rica and West Indies) and Mediterranean areas.
5 Environmental Distribution
Most living Dasycladales are growing in tropical and subtropical marine waters.
Few species occur in warm-temperate seas (Valet 1979). They are confined to
shallow environments, usually less than 5 metres depth. Maximum abundances
occur in quiet areas sheltered from wave action. Individuals are attached to
rocks, small stones, shell or coral fragments, mangrove roots, etc. Some species,
such as Halicoryne spicata and Acetabularia mediterranea, tolerate substantial
salinity fluctuations. Batophora oerstedii is found in marine and brackish
environments, even sometimes in freshwater.
Ecological requirements of Cenozoic Dasycladales seem to be the same as
those of living representatives (Elliott 1984). Richest assemblages have been
found in sediments and with other organisms that suggest shallow warm marine
environments: lagoonal waters behind reefs, coastal bays, etc. [Palaeocene-
Lower Eocene of the Middle-East; Lutetian of the Paris basin (Genot 1987);
Palaeocene of Sardinia (Dieni et al. 1985)]. No fossil species is known in
freshwater deposits.
(J
Table 7. Proportional representation of species number in Cenozoic and Recent DasycIadales genera
:>
'"o
~
DAS Y LAD
C ACE A E ACE TAB U L A R IACE A E n-
Ol
:>
Q.
;:0
()
Hoilcoryne '"
~Chl~ .-~....:..- . ~ '"
::'.
><:
~ ,
~ ~
".,
()
z:
UJ Q.
~ Cymopollo Ol
"'
u ;;-
UJ
'"
a::
---..:ce'ObUIor 10 ~
Bornetello
IW~
T
Rostroporelio
inc
(21I.oth gen.erc
3Serspecies)
~. ~HallCOryne
u ~J(
;....
o JOdote~ =:J ;loP;~ rrA
N
o
~ . "
z:
UJ
Dissociodeiia
u
lil~r\
.j:>.
w
144 P. Genot
Table 8. Main Dasycladales genera: evolution of the number of species during the CenozoIc
PLIOCENE
MIOCENE
.1,
Iinc ludlng
Lorvor 10
Lemolnello ()
?
-- 10 specIes
12mm wIde
= 'specIes)
Memlnello)
CymoDollo Aclculorlo
?
OLIGOCENE
ZI ttellno
I including
Ii
~ MoUDosio
0 PRIABONIAN DIgltellol
? /
u.J~ BARTONIA~
20 0
()
u.J
u
0
s LUTETIAN I
0
u.J
;:
I(
YPRESIAN
S Clypelno
'" \.
( \
?
...;
0
THA~ETIAN
0
u.J
«
-' I
""
<>. DANO -MONTI ~.N
\ J
DASYClADACEA E AC E TABULARIACEA E
Acknowledgements. I am very grateful to M. Hauray and A. Barreau for scanning electron

micrographs and A. Cossard for drawings (University of Nantes).
References
Deloffre R. Genot P (1982) Les Algues Dasycladales du Cenozoique. Mem Cent Rech Explor Prod
Elf-Aquitaine 4:247 pp. 20 pI
Dieni I. Massari F. RadoiCic R (1985) Palaeocene dasycladalean algae from Orosei (eastern
Sardinia). Mem Sc Geol Inst Geol Miner Univ Padova. XXXVIII. 22 pI
Elliott G F (1984) Climatic tolerance in some aragonitic green algae of the Post-Palaeozoic.
Palaeogeogr Palaeoclimatol Palaeoecol 48: 163- I69
Genol P (1987) Les Chlorophycees calcaires du Paleogene d'Europe Nord-Occidentale (Bassin de
Paris, Bretagne, Cotentin, Bassin de Mons). These Doctorat d'Etat, Nantes, 518 pp, 48 pi
Tappan H (1980) Order Dasycladales. In: Freeman W (ed) The paleobiology of plant protists.
Freeman San Francisco. pp 860-912
Valet G (1969) Contribution a l'etude des Dasycladales, pt2,3. N Hedwigia 17 :551 - 644, 133- 162 tab
Valet G (1979) Approche paleoecologique du monde des Dasycladales a partir de I'ecologie des
formes actuelles. Bull Cent Rech Explor Prod Elf-Aquitaine 3, 2:859-866
Table 9. Dasycladales: main discoveries of Palaeogene species in the world ("')
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Chapter 9
Fossil Udoteaceae and Gymnocodiaceae
Abstract
The Gymnocodiaceae and fossil Udoteaceae (erect) are similar to each other both in growth form
and in vegetative structure. They differ in that the Gymnocodiaceae contain internal reproductive
organs which are usually absent in the Udoteaceae. It is generally accepted that the Gymnocodiaceae
belong to the red algae and the Udoteaceae to the green algae. Well-preserved silicified Gym-
nocodiaceae can reveal some details of cell filaments, reproductive organs and parasitism. However,
the systematic position of this family still remains to be resolved. The published records of fossil
Udoteaceae are more abundant, but their biological characters are less well known and much
discrepancy exists in their nomenclature, definition and content. In view of the fact that fossil
material can only provide limited data on the anatomy of the algae and no information is available
at present about pigments, storage products, chemical characters of cell wall or reproduction which
are critical to the classification ofliving algae, and that parallelism is frequently found in different
groups of living algae, the Gymnocodiaceae and fossil Udoteaceae are considered heterogeneous
and in practice it is often difficult to distinguish between them. In order to advance our under-
standing of these important groups, search for better preserved permineralized fossil material and
close cooperation between students of fossil and living algae are badly needed. .
1 Introduction
Udoteaceae and Gymnocodiaceae are important groups of calcareous algae.

The former are green algae, represented by both fossil and Recent forms widely
distributed in tropical (or subtropical) shallow seas, and their history can be
traced back to the Ordovician; the latter is an extinct family, which is generally
considered to belong to the red algae, ranging from the late Paleozoic to the
Early Cenozoic(?) with an ecological distribution roughly similar to that of the
Udoteaceae.
Gymnocodiaceae are characterized by an erect, branched or unbranched,
segmented or nonsegmented, thallus composed of medullary and cortical zones
with reproductive organs located in the peripheral part. The erect Udoteaceae,
which predominate in the family, have much in common with Gymnocodiaceae
except that the reproductive organs are usually not preserved in the former and
in practice it is often difficult to distinguish between them. For this reason they
are treated here together.
1Nanjing Institute of Geology and Palaeontology, Academia Sinica, Chi-Ming-Ssu, Nanjing,

People's Republic of China
Fossil Udoteaceae and Gymnocodiaceae 147
2 Fossil Udoteaceae
Although the history of studies on fossil U doteaceae (long called Codiaceae) can
be traced back more than a hundred years, it was not until 1926-1927 , when Pia
incorporated Boueina and Palaeoporella together with Gymnocodium into the
Codiaceae and pointed out their importance as rock builders, that the concept of
a group offossil Udoteaceae became rooted in the literature. Since then the fossil
records of the family have steadily increased with new genera and species being
added, until now more than 30 genera ha ve been described as U doteaceae; some
of which, however, may not belong to the family. Konishi (1961) first sum-
marized Palaeozoic Udoteaceae. Recently, Roux (1985) has made a synthesis
for Palaeozoic algae in which fossil Udoteaceae are included. A comprehensive
summary of fossil U doteaceae (erect) ranging from Palaeozoic to Cenozoic has
been compiled by Bassoullet et al. (1983). The latter is an important source,
although it is not exhaustive and some genera included may not belong to the
Udoteaceae.
Morphologically, the fossil Udoteaceae are naturally represented by three
categories: filamentous (nodular), phylloid, and erect. The following will centre
on the last group, with only a brief account offilamentous forms when necessary.
3 Morphology and Anatomy
The morphology and anatomy of Recent U doteaceae vary greatly, ranging from
simple filamentous to erect segmented forms with tissue differentiated into
medulla and cortex. The erect segmented forms are exemplified by Halimeda
although some forms of Halimeda have a prostrate habit which is represented
by both Recent and fossil taxa, and its morphology and ana tom y ha ve been used
to make comparisons by analogy with other fossil forms. Hillis (1959) detailed
the morphology and anatomy ofliving Halimeda, and Bassoullet et al. (1983)
made a generalized schematic diagram of fossil erect Udoteaceae.
4 Growth Form
The growth forms offossil erect Udoteaceae which also resemble those of the
Gymnocodiaceae, may be branched (dichotomously or trichotomously) or un-
branched, and segmented or non-segmented. Segments are variously shaped
and may be cylindrical, oval, spherical, etc. The node (Hillis 1959), an important
feature in living Halimeda, has not yet been recorded in fossil Udoteaceae.
5 Anatomy
The thallus is generally composed of a medulla and cortex (Fig. la). The
medulla may be monoaxial or multiaxial. The medullary filaments are arranged
148 X.Mu
a d
c f
Fig. la-f. Convergence of internal organization in some living algae. a Halimeda (Udoteaceae,
Chlorophyta); b Penicillus (Udoteaceae, Chlorophyta); c Codium (Codiaceae, Chlorophyta); d
Cumagloia (Helminthocladiaceae, Nemalionales, Rhodophyta) ; e Galaxaura (Chaetangiaceae,
Nemalionales, Rhodophyta); f Furcellaria (Furcellariaceae, Gigartinnales, Rhodophyta). a after
Hillis-Colinva ux (1980); b after van den Hoek (1981) ; c after Tseng et al. (1962) ; d, f after Dixon,
1973 ; e after Svedelius (1953)
parallel to the axis of the thallus or interwoven or tangled, and bifurcate or

trifurcate to give rise to lateral cortical filaments. The greatest variation in fossil
Udoteaceae is represented by the cortical filaments, which are straight, swollen,
or constricted, or show a combina tion of these (Fig. 2). They rna y branch several
times before reaching the surface. The termination of lateral filaments may be
represented by small utricles (swellings).
Fig.2a-h. Representations of filament and utricle structure in various udoteacean genera, mostly
based on materials in the collections of the British Museum (Natural History): a Dimorphosiphon
(Ordovician); b Aphroditicodium (Permian ; BM(NH) Dept. Palaeont. reg. no. V.59461) ; c Tau-
ridium (Permian) ; d Arabicodium (Jurassic-Cretaceous); e Boueina (Triassic-Cretaceous); f
Halimeda incrassata (Ellis) Lamx (Recent) (Hillis 1959); g Leckhamptonella lIewellyae Elliott
(Jurassic); h diagrammatic growth plan of serial-segmented udoteacean. See also comparisons for
some other Palaeozoic genera in Obrhel (1968. Fig. I) and Guilbault and Mamet (1976, Fig. 2) (after
Elliott 1982)
6 Reproductive Structures
The reproductive structures in Recent Udoteaceae are both internal and ex-
ternal (Hillis-Colinvaux 1984), and the latter have no potential for fossilization.
In most fossil Udoteaceae no reproductive structure can be seen , However,
rounded cavities located in the calcareous wall of the thallus, which are
interpreted as reproductive organs, have been reported from a few Palaeozoic
150 X.Mu
genera such as Orthriosiphon (Johnson and Konishi 1956), Orthriosiphonoi-

des (Petryk and Mamet 1972) Litanaia (Maslov 1956), Uva? (Maslov 1956).
They also exist in some Mesozoic forms, e.g. Arabicodium elongatus (Dragastan
1971 P1.7, 3,4,7), A. texana (Johnson 1969, P1.2, 5,7), and possibly Halimeda
elliotti (Conrad and Rioult 1977, Pl.l, 7,9,10, etc.). Since all the so-called
"reproductive structures" in fossil material show no remaining cytological
features and it is impossible to determine their biological nature, further
comparison of them with Recent algae cannot be made at present.
7 Criteria for Classification
W ra y (1977) proposed three criteria for generic classifica tion of the erect as well
as phylloid Udoteaceae, which I have modified as follows:
I. Growth form: branched, unbranched, segmented, non-segmented, and

bladed.
2. Vegetative structure: size, shape, number, arrangement and branching
mode of med ullary and cortical filaments.
3. Reproductive structure: shape, size and location.
8 Calcification
The calcification of living Udoteaceae, e.g. Halimeda, has been well

documented. It is characterized by precipitation of aragonite crystals,
predominately needle-shaped, on the surface offilament walls within the thallus
(Flajs 1977a,b; Hillis-Colinvaux 1980). In most of the fossil Udoteaceae their
calcareous skeletons are composed of calcite which are the replacement
products of the original aragonite. Two kinds of calcite may be recognized in
fossil Udoteaceae: the more common is sparite, the other is dark micrite, which
is only seen in a few genera, e.g. Lowvillia and Dimorphosiphonoides (Guilbault
and Mamet 1976).
In fossil Udoteaceae, calcification mainly occurs in the cortex and the
medulla is usually poorly calcified. As a result, the fossil is mostly represented
by a calcareous tube perforated by many canals (pores). In this case, they are
easily confused with Gymnocodiaceae and sometimes it is also difficult to
distinguish them from fossil Dasycladaceae. Conrad and Rioult (1977) dis-
cussed the micritization, sparitization and silicification of Halimeda elliotti
which, however, may not belong to the Udoteaceae (see below).
9 Systematic Position
Although the published records of fossil Udoteaceae are much more abundant
than those of the Gymnocodiaceae, the biology of the material is less well known
and their taxonomic assignments, at least for some of genera, have not been
resolved with certainty. The reproductive structures deciphered in certain

forms, as mentioned above, throws doubt upon their udoteacean affinities. They
are more similar in shape, size and preservation to those commonly seen in the
Gymnocodiaceae, rather than to Recent Udoteaceae. They resemble to some
extent the gametangia of Codium (Codiaceae) and Pseudocodium
(Pseudocodiaceae, Hillis-Colinvaux 1984) in being internal, but do not open to
the surface of the thallus. This kind of reproductive organ is common in a
number of Recent red algae, so the possibility that these fossil forms may not
belong to the Udoteaceae cannot be ruled out.
The major arguments for udoteacean affinities of fossil forms lies in their
gross morphology and coenocytic nature. Since the information obtained from
fossil material is limited and parallelism is a common phenomenon among
different algal groups (Fig. 1), it is likely that the assignment of a fossil form to
the Udoteaceae may some times exceed the evidence available in the fossil state.
10 On the Classiffication of Fossil Udoteaceae
The classification offossil Udoteaceae is a matter of controversy (see Table 1).

In the 1920s, Pia referred erect forms, such as Boueina and Palaeoporella,
together with Gymnocodium, to the Codiaceae; and filamentous (or nodular)
forms such as Ortonella, Mitcheldeania, and Hedstoemia, were assigned to his
artificial group Porostromata in the Cyanophyta (Pia 1927). However, some
authors have suggested codiacean affinities for the latter and a new subfamily
Mitcheldeanieae was created by Pia for them (Pia 1937). After Pia's work, some
authors accepted the codiacean affinity (Elliott 1956; Johnson 1961; Konishi
1961; Bourque et al. 1981, etc.), buta new subfamily name Garwoodioideae was
proposed to replace Mitcheldeanieae (Elliott 1956; Konishi 1961). Others still
preferred a cyanophyte affinity (Maslov 1956, 1963; Shuysky 1973; Emberger
1976, 1978, 1979; Wray 1977, etc.); or considered that some belonged to the
Cyanophyta and others to Chlorophyta. Riding (1977) suggested that the
informal taxonomic name Porostromata be retained to cover these filamentous
forms together with Girvanella. Subsequently Dragastan (1985) distributed
between the Cyanophyta and Codiaceae. It should be noted that the family
name "Codiaceae" used by the above authors is synonymous with the current
usage of Udoteaceae. It is evident that the filamentous (or nodular) and erect
groups are not homogeneous, and hence should belong to different algal
divisions. For example, there are good reasons to compare Ortonella with the
blue-green alga Scytonema (Monty 1967), whereas Paragarwoodia (Poncet
1974) may be related to the Udoteaceae. Considering that parallelism also
occurs in filamentous forms among different divisions of Recent algae, caution
must be made in determining their systematic position.
Konishi (1961) tried to establish a system of classification to incorporate
both Recent and fossil Udoteaceae. Because the classification of Recent ex-
amples relies heavily on cytology and reproduction, details of which are rarely
available under fossil conditions, this approach is limited. The same problems
apply to the approach ofRoux (1985), in which filamentous forms are assigned
152 X.Mu
Table 1. Classification schemes of filamentous or nodular "U doteaceae" (boxed)
Johnson 1961;
Bourque, Mamet &
Pia 1927 Maslov 1963 Wray 1977 Pia 1937 Roux 1981
Cyanophyta Cyanophyta Codiaceae Codiaceae
BedstToemia BedstTaemia
Garwoodia Garwoodia
OTlonella OTtonella
Cayeuxia Cayeuxia Bevocastria
Rothpletzella Bevocastrza DimoTphosiphon
PalaeopoTella
Chlorophyceae
Codiaceae Codiaceae Codiaceae
Halimeda Boueina PalaeopoTella
Boueina Calci[olium DimoTphosiphon
Gymnocodium Abacella Eugonophyllum
PalaeopoTella Litanaia Anchicodium
Ovulites Lancicula Calci[olium
Uva Ivanovia
Dragastan 1985 Roux 1985 Riding 1977
Cyanophyta Codiaceae Porostromata

Rivulariaceae filamentous Codiaceae (blue green + green algae)
Ortonella (Proparte) Hedstroemia Girvanella
Cayauxia Garwoodia OTtonella
RivulaTia Mitcheldeania HedstToemia
Ortonella Garwoodia
Scytonemataceae Gaspesiella Cayeuxia
OTtonella Bevocas!nQ
ParaoTtonella
Mitcheldeaniaceae Udoteaceae
I Mitcheldeania erect U doteaceae
Chlorophyta Palaeoporella
Codiaceae Dimorphosiphon
Litanaia
Lowvillia
Bevocastria Succodium
Arabicodium Mas/ovina
to the Codiaceae, and erect ones to the Udoteaceae. Without information on

cytology and reproduction it is very difficult to distinguish between fossil
Codiaceae and Udoteaceae.
Studies of fossil Udoteaceae owe a great deal to the advances in our
knowledge of Recent Siphonales. They suffer from these as well. Several
decades ago when Codium and Halimeda were grouped into the same family
Codiaceae (e.g. Fritsch 1935) it was convenient for palaeophycologists to follow
this classification. Recently it has been realized that essential differences exist in
the biological nature of Halimeda and Codium; they have been assigned to
different families in different suborders (Hillis-Colinvaux 1984), and fossil
Codiaceae, following Halimeda, have been transfered to the Udoteaceae or
Halimedaceae. Incidentally, it has been shown recently (Hillis-Colinvaux 1984)
that the family name Halimedaceae has priority over Caulerpaceae and
Udoteaceae (Silva 1980). For reasons mentioned above these changes com-
plicate the subdivision of fossil Udoteaceae. The most probable difference
between the two families which could be preserved in the fossil state is
calcification. The known fossil Udoteaceae are calcified, whereas the fossil
Codiaceae are not. However, calcification seems less important than has been
suggested by some authors (Poncet 1982; Hillis-Colinvaux 1984) because
calcareous and non-calcareous forms can be found in the same family, e.g. in the
Dasycladaceae.1t has also been suggested that a Recent non-calcareous group
may have calcareous ancestors and vice-versa (Elliott 1978). Therefore, it is very
difficult, if not impossible, to distinguish between fossil Codiaceae and
Udoteaceae.
11 Geological Importance and Evolution
Recent members of the Udoteaceae (e.g. Halimeda, Penicillus) are among the
most important carbonate producers and bioherm builders in warm seawater
(Milliman 1974; Neumann and Land 1975; Hillis-Colinvaux 1980; Drew 1983;
Hudson 1985; Drew and Abel 1988; Hine eta!' 1988; Marshall and Davies 1988;
Multer 1988; Orme and Salama 1988; Payri 1988; Phipps and Roberts 1988;
Roberts et al. 1988; Hillis-Colinvaux, this Vol.). Their fossil representatives
could be important limestone or bioherm builders too (e.g. H0eg 1927; Johnson
1961; Jux 1966; Segonzac 1986; Fluge11988; Mankiewicz 1988).
Recent Udoteaceaens are a major source ofCaC03 -mud from post-mortem
disintegration of their skeletons. It has been speculated that many Palaeozoic
carbonate rocks rich in mud may be formed by fossil Udoteaceae (Wray 1977).
In the study of Ordovician algae from Canada, Mamet et al. (1984) pointed out
the role of the Udoteaceae in sedimentation of carbonate: Dimorphosiphon
contributes to carbonate production and Lowvillia to buffiestones. Recent
Udoteaceae are distributed in tropical (or subtropical) seas with low to moderate
energy. Although most ofthem are shallow-water inhabitants, they may extend
to 140 m with relatively high density and diversity (Hillis-Colinvaux 1986).
They are important constituents of the chlorozoan assemblage (Lees and Buller
1972), commonly in association with Dasycladaceae and hermatypic corals.
However, the lower limit of their depth distribution is greater than the latter
two.
It is believed the same is true for their ancestors. The palaeoecology offossil
Udoteaceae is discussed in a general way by Elliott (1978, 1979, 1984) and he
(1981, 1984) demonstrated their Tethyan distribution during Mesozoic and
Cenozoic time. Poncet (1982) used Devonian occurrences of fossil Udoteaceae
to locate palaeolatitudes of continents. Information about the characteristic
representatives and their palaeoenvironmental distributions in Palaeozoic time
can be found in the summary by Roux (1985).
Little is known of the evolution of Udoteaceae. A level of organization
similar to that of Recent Halimeda was achieved in Dimorphosiphon in Or-
dovician times. It is suggested that Halimida arose by the hybridization of
Boueina and A rabicodium in the Early Mesozoic (Elliott 1965). In a review ofthe
154 X.Mu
evolution of the "serial segmented" Udoteaceae from Palaeozoic to Recent,

Elliott (1982) considered that swollen cortical branches is a more advanced
character than that of straight ones.
12 Gymnocodiaceae
Gymnocodium, the type genus of the Gymnocodiaceae Elliott, was established

by Pia on the basis of Gyroporella bellerophontis Rothpletz 1894 (Pia 1920),
which was previously referred by him to the genus Macroporella, a dasyclad
acean (Pia 1912). Afterwards, he compared it with Halimeda and Boueina,
and transferred it to the Codiaceae (Pia 1920, 1927). Having discovered re-
productive structures in some specimens, he compared it with Galaxaura, a
Recent red alga of the family Chaetangiaceae, and transferred it once more to
the Chaetangiaceae (Pia 1937). In 1955, Elliott created the Family Gym-
nocodiaceae, and in doing so he made a further comparison with Galaxaura
which supported that of Pia, and established a new genus Permocalculus. Since
Elliott's work the red algal affinities ofthe Gymnocodiaceae have been generally
accepted although some authors still consider them as Codiaceae (Bilgiitay
1959; Konishi 1961; Maslov 1963, etc.).
During the last three decades, studies of the Gymnocodiaceae have ex-
panded both geographically and geologically: from Europe to Asia, America
and Africa; from the Permian to Jurassic and Cretaceous and Early Tertiary?
The following works of regional floras are some of the publications concerned,
to differing degrees, with the family: Accordi (1956), Bilgiitay (1959), Elliott
(1955, 1958), Giiven~ (1966), Herak and Kochansky-Devide (1960, 1963),
Johnson (1951, 1968), Kochansky-Devide (1954), Konishi (1954a), Korde
(1965), Mu (1981,1982), Ogilvie-Gordon (1927), Rao and Varma (1953), Rezak
(1959), Srivastava (1982), Termier et al. (1977), Vachard (1980). Emberger
(1976) compiled bibliographies of the Carboniferous and Permian algae, in
which both Gymnocodiaceae and Udoteaceae are included.
In the study of Permian Gymnocodiaceae from SW China, I have found
beautifully preserved silicified material. Using orientated thin-sections of
isolated specimens, which reveal some details of the anatomy of the algae I
established a new subgenus Pyrulites in Permocalculus (Mu 1981), and for the
first time discovered fossil fungi parasitic in Gymnocodiaceae (Mu 1977). Mu
and Riding (1983) used the SEM to study acid-extracted silicified material from
the Permian of China, and established a new genus Nanjinophycus, which was
compared with Galaxaura and tentatively placed in the Gymnocodiaceae.
13 Morphology and Anatomy
1. Growth Form
The growth form of the Gymnocodiaceae is erect, branched (dichotomously or
trichotomously) or unbranached, segmented or non-segmented. The shape of
the segments is cylindrical, oval, spherical of pyriform. No uncalcified node has

been found between segments.
2. Anatomy
The thallus is composed of filaments arranged in central (medullary) and
peripheral (cortical) zones. The medullary filaments can rarely be seen because
of poor calcification in this part of the thallus (PI. 1,6). They may be straight, or
swollen and constricted. They run longitudinally along the axis, at tiIl1e~
bifurcating and giving rise to lateral filaments at right or acute angles. Lateral
filaments are generally similar to medullary ones in shape, and branch several
times to reach the surface of the thallus where they form polygonal structures.
The diameters of the lateral filaments may be thinner than those of the
medullary ones, but in some forms they terminate in the form ofa funnel or bowl
with diameters which may exceed those of the medulla.
The filaments are usually preserved in limestone in the form of canals or
pores perforating calcareous walls, which are generally filled with dark matrix
(micrite). None of the soft parts (e.g. cell wall) can been seen and hence no
cross-partitions have been found. The filaments seems to be coenocytic. This will
be discussed in some detail below.
14 Reproductive Structures
In the cortex, or the area between cortex and medulla, oval or spherical cavities
are frequently found which are filled with matrix (micrite) or cement (spar). The
pattern of their preservation falls into two types:
l. Rounded cavities scattered in the calcareous wall filled with dark matrix
(PI. 1,6);
2. Rounded or oval bodies each with a thin dark membrane (PI. 1,1,3). In the
best specimens, it can seen that they are each located on a short lateral filament
with the terminal part tapering into a thin tube which is open on the surface of
the thallus. Sometimes they are arranged in layer along the peripheral part ofthe
thallus (PI. 1, 1,3,5,6; Mu 1981, pI. I, 11; pI. II, 2,3,6).
The reproductive organs of the Gymnocodiaceae have been compared
with conceptacles of the Chaetangiaceae (Elliott 1956; Mu and Riding 1983) or
tetrasporangia of Corallinaceae (Mu 1981). In the study of Permocalculus
ampulancea, Elliott (1961) recognized asexual and sexual (male and female)
reproductive organs, which were suggested to represent asexual and sexual
generations comparable with those in living Galaxaura. However, it was found
out later that the "male" plant was another alga (Elliott, personal commun-
ication). Korde (1965) elaborated a number of reconstructions of reproductive
organs of Gymnocodiaceae, which are based only on a few thin-sections. It
seems to me that the preservation of her material does not allow such
imaginative reconstructions.
The reproductive structures seen in the Gymnocodiaceae are the strongest
argument for red algal affinities, which led to the transfer of Gymnocodium from
156 X. Mu
Plate I
the green to the red algae (Pia 1937). Subsequent work seems to support this
interpretation. However, the systematic position of Gymnocodiaceae is not
finally resolved. This will be discussed below.
15 Criteria for Classification
The following criteria can be useful for generic classification in the family:
1. Growth form: branched, unbranched, segmented, non-segmented,
pyriform.
2. Vegetative structure: size, shape, number and arrangement of medullary and
cortical filaments; and the ratio of diameter of medulla to outer diameter of the
thallus.
3. Reproductive structure: shape, size and location.
16 Calcification and Silicification
The most commonly found skeletons of the Gymnocodiaceae are calcite. Since
the family is compared with Recent Galaxaura or Liagora, the calcareous
skeletons of which are aragonitic (Flajs 1977a,b), it is suggested by analogy that
the same is true for the Gymnocodiaceae (Mu and Riding 1983). Calcification
usually takes place in the cortical zone; the medullary zone is generally not
calcified. During diagenesis their aragonitic skeletons are transformed into
calcitic ones.
Silicification is common in the Gymnocodiaceae. In the early diagenetic
stage, the cell wall of the algae may be preserved (see Mu 1977, 1981). In late
silicification, which is common in carbonate rocks and frequently characterized
by the selective silicification of fossil skeletons, no soft parts of the algae are
preserved; Mu and Riding (1988) studied the pa ttem and process of silicifica tion
of Nanjinophycus, a possible Gymnocodiacean, and proposed a model for the
silicification process. They suggested that water insoluble organic material in the
matrix controls the selective silicification, by preventing the calcite in the matrix
from silicifying.
Plate I. 1-5. Permocalculus (Pyrulites) sinicus Mu. Changhsingian, Upper Permian; W. Guizhou,
China (Mu 1981).1 A sporangium terminal on a short lateral filament (x50); 2 lateral view ofa solid
silicified specimen (xl); 3 longitudinal section of the same specimen (2) (x8); 4 transverse section of
the initial part of the same specimen (2); (xI5); 5 surface view of epidermal cells ofthe cortex in an
isolated silicified specimen. Note the larger openings (arrows) representing the outlets of discharge
tubes of sporangia (x45); 6 Gymnocodium bellerophontis (Rhothpletz). Longitudinal section; Be!-
lerophon Formation, Upper Permian, Alleghe area (Belluno), W. Dolomites, Italy. Note oval
sporangia with terminal discharge tubes (x20; Fois 1977) 7 Dimorphosiphon rectangulare H0eg. A
longitudinal section and some transverse sections; Middle Ordovician, Lake Mj0sa, Norway (x20;
Elliott 1972)
158 X.Mu
17 Systematic Position
The growth form, internal structure with differentiation into medulla and
cortex, together with the shape; size and location of reproductive organs, all
suggest that the Gymnocodiaceae are more similar to red algae than to any
others. However, their red algal nature has not conclusively been confirmed. An
important difficulty which remains is that until now no cross partitions have
been found within the filaments of Gymnocodidium and Permocalculus, which
are typical ofthe family, even in the best materials where the cell wall ofthe algae
probably is preserved (see Mu 1977). If the cell filaments are really non-septate,
the algae are more likely to be green algae. It may be, however, as I have
suggested (Mu 1981) that the cross septa were entirely represented by pit-plugs
(septa plugs) which are characteristic of red algae. In Recent red algae the septal
plug is chiefly composed of protein (Pueschel 1981) which is less resistant and
cannot survive fossilization. Considering that no direct evidence is available at
present, assignment of the Gymnocodiaceae to the red algae is not conclusive.
Certain genera relevant to the Gymnocodiceae need discussion of their
taxonomic assignment. Nipponophycus Yabe et Toyama was originally referred
to the Furcellariaceae (red algae) (Yabe and Toyama 1928). Since then it has
been assigned to the Solenoporaceae (Endo 1961) and Udoteaceae (= Codia-
ceae) (Johnson 1964, 1969; Bassoullet et al. 1983). Tappan (1980) retains
it in the Furcellariaceae. I showed the presence of reprod uctive structures like
those of the Gymnocodiceae and tentatively transferred it to the family (Mu
1982). Although its precise systematic position remains to be resolved, it seems
more reasonable to ascribe it to the red algae than to any other division.
Succodium Konishi is generally considered as belonging to the Udoteaceae.
However, the presence of structures interpreted as reproductive organs led
Korde (1965) to transfer it to the Gymnocodiaceae. This was followed by
Vachard (1980) and Vachard and lVIontenat (1981), but rejected by some others
(Bassoullet et al. 1983). Mu and Riding (1983) suggested that if its internal
reproductive organs can be substantiated, Succodium Konishi should be as-
signed to the Gymnocodiaceae like Nanjinophycus Mu and Riding. The sys-
tematic position of Succodium is not conclusive at present. Diversocallis
Dragastan was erected as a gymnocodiacean alga (Dragastan 1967). However,
its irregular form and compact internal structure suggest that it is very likely to
be a synonym of Pseudolithothamnium Pfender (Squamariaceae). Oligoplagia
Herak was initially referred to calcisponges (Herak 1944) and subsequently to
microproblematica (Ott 1967) and then Gymnocodiaceae (FlU gel 1971 ). Owing
to poor preservation of its internal structure, and presence of cross partitions it
cannot be assigned to the Gymnocodiaceae and its systematic position remains
to be clarified. Abatea Senobari-Daryan and Schafer was classed as a genus of
the Gymnocodiaceae (Senowbari-Daryan and Schafer 1980). But the existence
of cross septa and "sporangia" in the medullary zone, which are quite different
from the characters of the Gymnocodiaceae, make this assignment doubtful.
18 Geological Importance of Gymnocodiaceae
The records of Gymnocodiaceae range from the Permian to Tertiary? with the
most abundant occurrence in Permian rocks. They are also abundant at some
levels in the Cretaceous. The Triassic genera Abatea and Oligoplagia, as men-
tioned above, migh t not belong to the family. Gymnocodiaceans are widespread
within the Tethys and indicative of warm, shallow sea waters with low-moderate
energy. They are important carbonate producers, and gymnocodiacean
limestones have been widely found in Tethyan Permian and some Cretaceous
rocks. During Permian time, their importance geologically and biologically,
includes: baffle sedimentation, carbonate production, and habitats for inverte-
brates such as foraminifers, ostrocodes, bivalves and gastropods. They also have
stratigraphical value. The Permocalculus (Pyrulites) zone, which is roughly
the equivalent of Palaeofusulina zone, indicative of the Changhsing stage of
the Upper Permian, can be recognized in a vast area of S. China, (M u, 1981).
19 General Remarks
The biological features of soft tissue of algae are rarely preserved in the fossil
state. For most Gymnocodiaceae and Udoteaceae, even the cell wall, which is
the most resistant part, usually could not survive fossilization. What we are left
with are only their altered skeletons, which are in the form of moulds. The
original filaments and reproductive organs of the algae are replaced by cavities
filled with matrix or cement (micrite or sparite).ln such material it is impossible
to identify the nature of reproductive organs and also impossible to prove
whether the alga is coenocytic ornot (Fig. 3). The affinities offossil algae are only
cell walls fossil

calcareous wall
red green
Fig.3. Comparison between possible original organization of cortical filaments in Nanjinophycus

and Succodium and the resulting fossil morphology. If cross-partitions occur but are not calcified the
final appearance will be similar to specimens lacking original cross-partitions; yet this feature , if
preserved, could help to distinguish between a red and green algal affinity. (after Mu and Riding
1983)
160 X.Mu
based on the superficial similarity between them and living ones. It is well-
known that parallelism or convergence is common among different groups of
living algae, i.e. representatives of different groups may have similar habit and
internal organization. For example, the external and internal vegetative
structures of Halimeda, (Udoteaceae) are in many ways similar to those of
Codium (Codiaceae), they can also be comparable with those of some Chae-
tangiaceae (red algae) and other algae (Fig. 1). Thus, the assignment of many
fossil genera to the Udoteaceae in the literature is uncertain. The only generally
available criterion to distinguish between the Gymnocodiaceae (red algae) and
the Udoteaceae (green algae) is that of the reproductive structures: ovoid or
spherical in form, and located in the peripheral part of the thallus in the
Gymnocodiaceae, but usually absent in the Udoteaceae. In Recent Udoteaceae
the reproductive organs are external in Halimeda and possibly internal in
Penicillus (Hillis-Colinvaux 1984). Their shape, size and location are not
comparable with those found in some genera of fossil Udoteaceae, which
suggests that these latter genera may not belong to Udoteaceae. They possibly
belong to red algae like the Gymnocodiaceae. When reproductive organs are
not seen in fossil algae, it is very difficult, if not impossible, to distinguish
between the Gymnocodiaceae and Udoteaceae. However, it is also likely that
some ancestors ofUdoteaceae may have reproductive organs dissimilar to those
of Recent ones. Thus, the possibility of their green algae affinities cannot be
ruled out at present.
20 Suggestions for Future Work
From the foregoing it is evident that problems in determining the systematic

position and classification of Gymnocodiaceae and fossil Udoteaceae are the
result of the inadequate information available regarding fossil material. In
studying fossil calcareous algae we are, of course, not dealing with complete
organisms, but only with their calcareous skeletons which are generally in the
form of moulds. The most important information on the biological nature of the
algae, such as pigments, storage products, chemical characters of the cell wall,
and reproductive organs, which are critical to the classification ofliving algae,
have been lost during fossilization. Diagenesis of fossils makes the case worse
(Elliott, 1984). As a result it is very difficult, ifnot impossible, to determine the
systematic position offossil algae based only upon their skeletons which are also
often altered, just as we cannot recognize the health of a person by his clothes.
To advance our understanding of Gym nocodiaceae and fossil Udoteaceae,
and other groups of fossil algae as well, some suggestions are proposed as
follows:
1. Search for better preserved permineralized material. The fossil material of

calcareous algae preserved in limestones is not good enough to reveal the details
of their biological nature, due to the fact that algal-bearing limestones are
usually laid down under oxidizing conditions which permit the soft parts of the
algae to decay rapidly after death. The ideal materials are permineralized, e.g.
silicified. Silicified specimens formed in an early diagenetic stage may preserve
some details of algal structures, even of their soft parts, (e.g. cell walls, etc.).
Beautifully preserved material may be found in chert beds or concretions in
shallow marine carbonate sequences. These deposits should be attentively
examined and sampled in the field.
2. A thorough comprehension of the structure of the fossil is the first step in

taxonomic work. Although this sounds commonplace, in practice it too often
ignored. Many algal taxa in the literature have been created based on only one
section, or a few random sections from different individuals which are often
poorly preserved. This may lead to mistakes in reconstruction of the algae and
to confusion of one taxon with another. To avoid this, the adoption of orientated
thin -sectioning is recommended (see M u 1981). The acid dissolving a pproach is
useful for obtaining complete individuals which can reveal the gross mor-
phology and surface nature of the algae. This method is most suitable for
selectively silicified material in limestones (Mu and Riding 1983). Acid mace-
ration methods should be used together with sectioning.
3. Close co-operation between experts on fossil and living algae is urgently

needed. The present is the key to the past. Palaeo biological and palaeoecological
studies on fossil algae heavily rely on information about Recent algae. There is a
wide scope for joint work in comparative anatomy, ecology, ultrastructure, and
chemotaxonomy, for which multidisciplinary approaches are needed. The
results of such cooperation will be rewarding and can lead to a better under-
standing of the Gymnocodiaceae and fossil Udoteaceae.
21 Conclusions
Fossil Udoteaceae and Gymnocodiaceae are important groups of fossil calca-

reous algae. The former ranges from Palaeozoic to Cenozoic with abundant
living descendants; the latt,er is an extinct group ranging from Palaeozoic to
early Tertiary? It is generally accepted that the Gymnocodiaceae belong to the
red algae. Well-preserved silicified Gymnocodiaceae can reveal some details of
cell filaments, reproductive organs and parasitism. However, the systematic
position of this group still remains unresolved. The records offossil U doteaceae
are more abundant, but their biological features are less well known and much
discrepancy exists in their nomenclature, definition and classification.
The erect group of fossil Udoteaceae resembles the Gymnocodiaceae in
many ways: growth form, vegetative structure, palaeoecological and geo-
graphical distribution, etc. The major difference between them which can be
deciphered is that the gymnocodiaceans contain internal reproductive organs
which are usually absent in the Udoteaceans. In view of the fact that fossil
material can only provide limited data on the anatomy of the algae, and no
information is available at present about pigments, storage products, chemical
162 X.Mu
characters of the cell wall, or reproduction, which are critical to the classification
ofliving algae, and that parallelism is common among different groups ofliving
algae at different taxonomic levels, the Gymnocodiaceae and fossil U doteaceae
are considered heterogeneous. In addition, their biological assignments are not
conclusive, and in practice it is often difficult to distinguish between them. In
order to advance our understanding ofthese important groups search for better
preserved permineralized fossil material, and close cooperation between
students of fossil and living algae by using multidisciplinary approaches are
badly needed.
Acknowledgements. I am grateful to Dr. G .F. Elliott for critically reading the manuscript and giving
helpful advice.
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Chapter 10
Recent Calcified Halimedaceae
Abstract
Five Recent genera ofHalimedaceae, namely Halimeda, Penicillus, Rhipocephalus, Tydemania, and
Udotea calcify. Only Halimeda and Udotea, the genera with more than 10 species, are widely
distributed in the tropical Atlantic, Indian and Pacific oceans, but Halimeda alone produces
extensive populations worldwide making it globally significant. Because of the essentially tropical
distribution of all 5 taxa they can be considered strong indicators of tropical environments.
Halimeda is also widely distributed across the reef, with some of its 33 species indicating
different reefal conditions. Approximately three-quarters of its species are associated with con-
solidated or gravelly habitats as opposed to sand/mud substrata. Large populations may occur on
the deep fore-reef where Halimeda grows at depths of over 140 m, in the extreme shallows of back
reef moderate energy environments, and in lagoonal regions. Some lagoonal populations overlay
Halimeda bioherms which date from at least the early Pleistocene. Extensive present day
populations together with Halimeda bioherms provide dramatic evidence of the importance of
calcareous Halimedaceae in reef construction, and offer support for calling reefs algal rather than
coral. Although Halimeda is a principal component of many tropical reef systems, non-Halimeda
reefs are also part of the global reef picture.
1 Definition of the Group
The largest living green marine algae belong to the order Bryopsidales, formerly
Siphonales, and of the modern algae they are, paradoxically, among the least
known. Six genera out of a total of twenty-four calcify. Five of the taxa, namely
Halimeda Lamouroux, Penicillus Lamarck, Rhipocephalus Ktitzing, Tydemania
Weber-van Bosse, and Udotea Lamouroux (Figs. 1-5) have been assigned to the
family Halimedaceae: Pedobesia MacRaild and Womersley has been placed in
the family Bryopsidaceae (Hillis-Colinvaux 1984). Assignment to these families
is based principally on differences in morphology, reproduction and distribu-
tion, although the data available for some genera are still very limited. Selection
of the family name Halimedaceae (Hillis-Colinvaux 1984) is based on the
epithet having priority over both Caulerpaceae and Udoteaceae (Silva 1980).
The most taxonomically diverse genus of the Halimedaceae is Halimeda
with 33 living species. Five sections ofthis genus, Rhipsalis, Opuntia, Halimeda,
Micronesicae, and Crypticae, were delimited in 1980 by Hillis-Colinvaux using
behaviour of filaments at the nodes, or intersegmental regions, as the basis of
separation. Of the other genera, Udotea has approximately 26 species, Penicillus
five species, Rhipocephalus two species, and Tydemania two species. The em-
'Zoology Department, The Ohio State University, 1735 Neil Ave., Columbus, Ohio 43210, USA
168 L. Hillis
Fig. I. Grove of the sand-growing species H. incrassata at a depth of approximately I m on the north
shore of Jamaica. (magnification ca. O.2x; photograph by P.A. Colinvaux)
Recent Calcified Halimedaceae 169
Fig. 3. Rhipocephalus phoenix (Photograph

courtesy of W. Fenical; magnification ca.
nat. size)
I------l
lcm
Fig.4. Tydemania expedi/ionis. A small por-

tion of a sprawling thallus collected in the
lagoon of Enewetak Atoll
Fig. 2. Penicillus meadow, north shore of Jamaica. Thallus stages range from pre-capitulum to
senescent capitulum. Predominant species is capita/us (magnification approx. O.2x)
170 L. Hillis
Fig. 5. Udolea sp. from the Great Barrier Reef. (Photograph courtesy ofD. Meyer; magnification
approx.0.66x)
phasis of this paper is on the living representatives, and on features other than
purely morphological ones that are of present or potential interest in inter-
preting the geological record.
1.1 Architecture and Calcification of the Halimedaceae
The coenocytic filament is the basic structural unit of the order. Different
branching patterns and organization of the filaments produce distinctive
capitate (Penicillus) , globuliferous (Tydemania), phylloid (Rhipocepha/us and
Udotea) and catenulate (Halimeda) forms, some of which have associated stipes,
rhizomes and much enlarged hold fasts. Gepp and Gepp (1911), Hillis (1959),
and Hillis-Colinvaux (1980, 1984) provide details of organization and taxo-
nomic criteria as well as additional references.
All five genera deposit calcium carbonate in the conspicuous phase of their
life history, in the form of aragonite (McConnell and Colinvaux 1967).
Calcification is external to the filament, and for Halimeda begins in the newly
formed (apical) segment, in association with the filament wall, when the
segment is about 36 hours old (Wilbur et al. 1969; Hillis-Colinvaux 1980). The
crystals are first deposited among the fibrils ofthe pilose outer layers ofthe walls,
thereby forming a lining of the interfilamental spaces of the segment (Wilbur et
al. 1969). In species of Rhipocephalus, Penicillus and Udotea aragonite deposits
are formed within a sheath. A recent useful review of calcification in algae is
provided by Borowitzka (1982).
Extent of calcification varies with age, taxonomy and environment. In
general Penicillus, Rhipocephalus and Udotea are more lightly calcified, with
mature thalli containing ca. 29-59 percent CaCOa (Hillis-Colinvaux 1980,
Tables pp. 268, 269, 271) whereas the range for the mature Halimeda specimens
sampled is 47-90% if the lightly calcified, subtropical H. cuneata Hering is
excluded. Highest percentages were obtained for recently shed segments and the
oldest segments on the plants. When analyzing such data sets it is important to
realize that although apical segments of Halimeda are the youngest (Colinvaux
et al. 1965), they may, nonetheless, contain carbonate deposits.
There have been few studies of calcium carbonate content with depth.
Bohm (1973) examined seven Halimeda species growing on the north shore of
Jamaica over depths from < 1 to ca. 57 m. Percent calcium carbonate decreased
in simulans Howe and goreauii W. R. Taylor at depth, remained essentially the
same for tuna (Ellis and Solander) Lamouroux, and increased for discoidea
Decaise, gracilis Harvey ex J. Agardh, opuntia (Linnaeus) Lamouroux and
copiosa Goreau and Goreau.
1.2 Geographic Distribution: Genera as Indicators of Tropical

Environments
The calcareous Halimedaceae are almost entirely tropical in distribution, as

defined by water temperatures of 25 C and higher rather than by degrees of
0
latitude (Hillis 1959). This definition produces asymmetric bands across the
Pacific and Atlantic oceans (Fig. 6) that are distinctly broader in the west than
in the east, and makes the presence of some of these genera in Bermuda and
Japan, as well as the absence of all of them from Peruvian waters readily
understandable. Exceptions to the tropical pattern include two species of
Halimeda. H. cuneata appears to be restricted to the subtropics of both
hemispheres of the Indian and Pacific oceans (Fig. 6), whereas H. tuna grows in
the subtropical Mediterranean but also has an extensive tropical distribution,
being reported for the Atlantic, Indian and Pacific oceans. Presence in three
oceans has been attributed to only six other species of the genus, namely
incrassata (Ellis) Lamouroux, simulans, opuntia, copiosa, disco idea , and gracilis.
Some species of Penicillus and Udotea also grow in the Mediterranean as well
as in the tropics. The Mediterranean presence of these three taxa may have
resulted from a recolonization event following massive extinctions during the
Messinian crisis (Hillis-Colinvaux 1980).
The genus Udotea may be less stenothermic than the others, with, for
example, three species growing in subtropical waters off North Carolina in the
northwestern Atlantic (Schneider 1976). No representatives of Halimeda or
172 L. Hillis
4 H. c!'ulleau
J 4 5
Fig. 6. Generalized global distribution of the Halimedaceae. Genera are principally tropical, as
delimited by 25 °C i sotheres, one of the exceptions being the subtropical species H. cuneata, which
has an authenticated distribution ofS. Africa, southern Australia, and Gulf of Kutch , India. The six
biogeographic regions shown are: western and central Indian Ocean, eastern Indian Ocean and
western Pacific, eastern Pacific, western Atlantic, eastern Atlantic, and Mediterranean (Hillis-
Colinvaux 1980)
Penicillus have been found in these waters (Searles, Schneider, personal

comm unica tion) .
Only two genera, Halimeda and Udotea, occur in the Atlantic, Indian and
Pacific oceans. The known distribution of Penicillus outside the Mediterranean
is essentially Atlantic and restricted Indo-Pacific; Rhipocephalus is limited to the
Caribbean, and Tydemania to the Indo-Pacific. Because all five genera have a
fairly strict tropical range they can be considered strong indicators of tropical
environments. This characteristic has been used by Elliott (e.g. 1981) in analyses
of the distribution of fossil green algal floras .
1.3 Within-Reef Distribution : Genera and Species as Indicators of Specific

Reef Environments
There are very few data on the distribution of the different genera or of their
biomass within specific reef systems, even for the most studied genus Halimeda.
In this section I present an overview of reefal distribution for the five genera.
More detailed information, especially for Halimeda, is provided in the sub-
seq uen t discussion of sediment prod uction, as well as by H illis-Colinva ux (1977,

1980, 1985, 1986a,b,c).
Rhipocephalus, Penicillus and Udotea generally grow as discrete plants in
regions of unconsolidated substrata, 'rooted' with a distinctive, often very large
holdfast of free uncalcified filaments and adhering sand particles. All three
genera can be locally abundant (Stockman et al. 1967). Tydemania also develops
dense local populations (Hillis-Colinvaux 1977, 1980), although both its habit
and habitat, succinctly described as sprawling over sand and rock surfaces, are
different.
In comparison, when conditions are suitable Halimeda, a genus with a
variety of types of holdfast (Fig. 7) such as the large bulbous style similar to
Penicillus, Rhipocepha/us and Udotea (found in the members of section Rhip-
salis), but also relatively small matted ones with few adhering extraneous
particles, or else discrete patches of loose filaments, can develop very dense
populations at many sites across the reef on both sand and rock substrata.
Consequently, Halimeda is one of the most versatile of algal colonizers. For
those accustomed to thinking of the genus as predominantly a plant with a
conspicuous holdfast for growing in lagoonal sands and muds, it is noteworthy
that approximately three-quarters of its species grow principally on substrata
other than sand (Hillis-Colinvaux 1985, 1986a). Sprawler species (mostly
Section Opuntia) are associated with rock or sand-gravel substrata and have a
habit rather similar to that of Tydemania; other Halimeda species grow readily
~HO"fO"OfO ~ Hofdf~,~b~~"d
lew loo.e liloment.;
u.uolly .everol on
filoments which lorm
a mot which 1 ix e. onto
o plont rock>
SPRAW LER ROCK-GROWER
f
l eg . tl. 1IRJL!!!ig ) (eg olj. tuno )
segment
Node I uj nClure 01 Segments )
HoldloS! of bronched l ilomenl$ ond odher ing

port icle. 01 .ond
SAND-GROWER
(ego lj. incro"oto)
Fig.7. Basic hold fast systems of Halimeda are associated with the major reef habitats of this genus.
Species which sprawl over rock or sand usually have several inconspicuous points of attachment;
strictly rock or sand-growing thalli generally possess a single holdfast (Hillis-Colinvaux 1980)
174 L. Hillis
on rock surfaces. Where hard substrata predominate and the surface is essen-
tially vertical as on many fore-reefs , draperies of Halimeda may develop over
much of the reef wall.
None of the five genera achieves sustained space in high energy environ-
ments such as the algal ridge (Fig. 8), but Halimeda does develop close to 100%
cover in regions of moderate energy such as the H. opuntia zone of inter-island
channels at Enewetak Atoll (Hillis-Colinvaux 1980, 1985). Its populations are
also uncommon to rare on most spur and grooves (Fig. 8); for this region
Hillis-Colinvaux (1977, 1980) reported about 15% cover by Halimeda off a
leeward islet of Enewetak Atoll in 1975, but essentially none for sites examined
on the north shore of Jamaica in 1967-1969.
All five genera have broad bathymetric ranges (Taylor 1960), with
Halimeda extending from the intertidal to at least - 150 m (Hillis-Colinvaux
1982, 1985, 1986a; Blair and Norris 1985, 1988). Depth tolerances among the
species, where known, are generally broad (Hillis-Colinvaux 1986a), although
one Caribbean species, H. cryptica L.H . Colinvaux and Graham appears to be
restricted to deep water, that is, to 25 m and deeper (Colinvaux and Graham
1964).
Present data indicate, therefore, that neither the genera of Halimedaceae
nor species of Halimeda with the exception of H. cryptica and possibly one or two
others, are precise depth indicators within the euphotic zone (Hillis-Colinvaux
1980, Fig. 90 ; Johns and Moore 1988). However, within an entire reef system
genera and species can function as indicators, as the following examples show:
I. An abundance of rhizophyte ('rooted') taxa usually indicates back-reef

environments, since species with well-developed holdfasts (Penicillus,
Rhipocephalus, the larger Udotea, and the taxonomic section Rhipsalis of
Halimeda) generally are uncommon to rare on the deep fore-reef;
[ ""-------.
\
I
LAGOON REEF ISLET or SEAWAR D REEF
-....
INTER - REEF - CREST FORE - REEF
LAGOON BASIN LAGOON SLOPE LAGOON BACK
IPINNACLE (2000.) I
'SLAN~L ~
TERRACE CHANNE = REEF ~
SPUR &GROOvE TERRACE SLOPE (WALL)
-
POPuLATIONS MAY BE vERY OENSE CLOSE TO SHOAE
ANO ON TOP Of PNNACI.ES .....
AI"1"'ftOJI_ I ~
COV£IIII, &OWl 0
PAOlA",.
LOW,
.. CM.vUI
0
I'" COV{II ON
'8ITELHWNO -
1I'11ANt.!TlC)Ioj,uM:U
..... 10 C;:OIlAl "[AOI
".11'11 ' "ND J
~1' . DlY(AIj"""
ANO COVER
QIIIOWTH TO ~14(IA
IItOCt(. .. .
SffllAflOilI
ANO I ROCK - GROwERS
SOME SPRAWl.EAS
0«"', :,1 _. ....,
10.TIItA.TEo.. If''AflGEI oOlll1l&
ZONE
o a:~~R$ 0 fII~I(-GAOWE.'
4lCI1HIi.-u;q
ROCK & ?SANO
SlRATEOIES
ROCK - GROWERS
SOME SPRAWLER$
Fig. 8. Distribution of Halimedaceae within a reef. Transverse section of Enewetak Atoll shows
distribution of the different growth strategies of Halimedaceae across the reef (boltom line), and
general Halimeda biomass (second line [rom botton) (Modified from Hillis-Colinvaux 1986a)
2. An abundance of these same taxa generally indicates unconsolidated

substrata;
3. Dense cover (80% or more) by populations of a single Halimeda species, in
which the plants are tightly branched and segments are reduced in size,
often indicates an environment of moderate energy as exhibited by shallow
(-0.3 m) channels with strong currents. The H. opuntia zone at Enewetak
provides an example.
2 Sediments of the Halimedaceae
Halimedaceae have several roles in the reef system. By far the most impressive,
and also the best documented is the capability of contributing aragonitic skeletal
materials to the reef system. One of the consequences is an historic record better
than that known for most algae (Elliott 1981; Fillgel 1988; Mankiewicz 1988).
The extensive contribution made by these algae to the building and mainte-
nance of global tropical reef systems is the only role that will be discussed herein.
2.1 Some Mechanics of Sediment Production
Geologists working in tropical reefs have been much more aware of the
sediments of the Halimedaceae and their importance than most botanists and
zoologists. This is a natural result of the main source of evidence for geologists,
the years of impressive accumulation of the distinctive Halimeda segment or
'plate'; for many reef sites this is considerably more imposing than the
populations ofliving plants which are often cryptic because of their greenness.
Debris of the Halimedaceae ranges from coarse to fine-grained sediments;
that is, from the skeletal modules or segments of Halimeda to the fine particles
of lime muds (Folk and Robles 1964; Hillis-Colinvaux and Orme 1988).
Generally it remains loose, but it may also be consolidated into packstones and
wackestones (Orme 1985), that is, into skeletal or lime-mud buildups respec-
tively as defined by Heckel (1974).
Of the five genera in the group only Halimeda produces a distinctive 'tracer'
in the sediments, its characteristic segment or 'plate'. Sedimentary materials are
released from Halimeda as a result of 1) growth and the natural shedding of
segments and branches, as leaves are shed from deciduous trees; and 2) death
of the entire plant, as by burial or sexual reproduction (Hillis-Colinvaux 1980;
Figs. 9, 10).
In contrast, the other genera do not have an architectural unit that long
survives death under most circumstances. Debris produced by Penicillus and
Tydemania from causes other than burial may initially be loose filaments; that
from Rhipocephalus and Udotea pieces of blade of various sizes, or separate
filaments. However, the ultimate fate of both thalli and debris of these four
genera probably is complete disintegration, and transformation into fine lime
176 L. Hillis
FI .10
Fig. 9, 10. Two stages of sexual reproduction in Halimeda. Fig. 9. The segments of H. cuneata are
fringed by uncalcified stalked gametangia. (Photograph courtesy of L. B6hm; magnification
approx. 2.5x) Fig. 10. H. monile, showing the disintegration that follows sexual reproduction
(magnification approx. I.7x) (Hillis-Colinvaux 1980)
muds (Stockman et al. 1967). In reef sediments some of this fine fraction may
also be derived from Halimeda.
For all the genera the quality of preservation of internal detail in
sedimentary material can be affected by the cause of death. For example, old
segments of Halimeda as well as thalli belonging to any of the five genera that
have reproduced sexually are physiologically senescent (Hillis-Colinvaux
1980). In contrast, much of the buried material I have seen appears to have been
vigorous.
2.2 Transport of Sediments
The general fineness of the debris produced by the Halimedaceae enables it to

be transported by the currents within and surrounding a reef as well as by storm
surges. Hence, Halimeda sediments are observed in regions oflagoons where no
living thalli occur, and mud banks derived from species of the Halimedaceae
shift position. Yet more obvious are Halimeda segments which have been
transported down the fore-reefslope. My transects of the fore-reef ofEnewetak
Atoll by submersible showed Halimeda sands to be a common feature of the
terrain surveyed to a bout 150 m on both windward and leeward slopes (F ig. II).
Fig. 11. Calcareous sediments, a large percentage originating from Halimeda , at approximately 92
m depth on the windward reefslope ofEnewetak Atoll. Dark clumps contain in situ living Halimeda
thalli (Photo by R. Thresher)
178 L. Hillis
They were observed in crevices, floored tiny ledges, accumulated in the inner
reaches of caverns as far as could be observed, and in many regions, especially
on the leeward side, blanketed the sessile biota including Halimeda itself. Below
about 150 m, where there are relatively few organisms on the walls to impede the
downward flow of fine skeletal materials, Halimeda segments were trapped by
small surface irregularities or were dammed by boulder piles (Hillis-Colinvaux
1986a ). Halimeda sediments contin ued to the base of the Enewetak submersible
transects at ca. 365 m. Davies (1986) reports them as an important derived
constituent of slope sediments of the Great Barrier Reef to over 2000 m,
although steepness of the outer reef slope influences the full extent of transport
of Halimeda-derived debris (Johns and Moore 1988).
Significantly, some of the debris accumulates in situ. In lagoonal regions
and on some vertical slopes gravelly sediments from Halimeda are sometimes
restrained by enveloping algal and cyanobacterial mats (Hillis-Colinvaux 1980,
Fig. 72) or by actively growing plant communities, frequently including
Halimeda. Some downward shifting of loose materials can be attributed to
bioturbation. For example, at Enewetak, callianassid shrimp, living near
Halimeda beds, remove Halimeda surface debris to refuse galleries over 50 cm
below the surface (Suchanek et al. 1986).
Especially dramatic in situ accumulations are Halimeda banks which
apparently remain in place for thousands of years. Examples are the Halimeda
reefs or bioherms recently described for the northern Great Barrier Reef and the
eastern Java Sea (Davies and Marshall 1985; Orme 1985; Phipps et al. 1985;
Roberts et al. 1987a,b; Hine et al. 1988; Marshall and Davies 1988; Orme and
Salama 1988; Phipps and Roberts 1988). Some of these mounds, which occur at
depths of 20 m and more, are described as being 'as thick as the reefs (coral)
themselves, cover(ing) areas larger than individual reefs, and hav(ing) accreted
calcium carbonate at comparable rates' (Davies and Marshall1985).Ages so far
recorded range from about 5000 years before present to modern times.
3 Sediment Production, and Importance of Halimedaceae in Building Reefs
Sediment accumulations signal the relative biological successes of the con-

tributors and their significance in present-day and ancient reefs. A quantitative
measure of carbonate production for the Halimedaceae consequently becomes
important in the analysis of the role and importance of Recent Halimedaceae.
Carbonate production is a function of:
3-dimensional space occupied x yield of CaC0 3 unit space- 1 X time.
This statement when applied globally to the Halimedaceae shows that
Halimeda, the most widely distributed and abundant member is the greatest
sediment producer, Rhipocephalus the least so. Within separate reef systems,
however, Penicillus is sometimes the principal contributor. In the following
discussion, examples are provided of high density occupancy oftropical benthic
space by Halimeda and Penicillus, at widely scattered reef sites in the Atlantic
Recent Calcified Ha1imedaceae 179
and Pacific Oceans. For understanding entire reef systems additional analyses
of reef regions away from the ridge are needed; they are especially desirable for
the Indian Ocean. As studies of the unconsolidated regions of reefs increase we
may expect many additional sites to be found with dense present-day
populations of Halimedaceae.
3.1 Three-Dimensional Within-Reef Space Occupied by the Halimedaceae
The quantity of live Halimeda in reefs is generally deceptive, made so by its

cryptic. When photographed against rock surfaces resolution may be so poor
that identification even as a Halimeda is precluded. Reefs therefore need to be
examined with an emphasis on this genus if we are to obtain good data on the
space occupied. At present there are analyses of only three reef systems
(Enewetak Atoll, north shore of Jamaica, and the Great Barrier Reef at the
latitude of Townsville), which include this genus across most of a reef system,
that is from back-reef or lagoon to deep fore-reef, and which therefore can
provide a statement of distribution within entire reef systems (Gilmartin 1960,
1966; Colinvaux and Graham 1964; T.F. Goreau and Graham 1967; T.R.
Goreau and N.J. Goreau 1973; Moore et al. 1976; Hillis-Colinvaux 1977, 1980,
1985, 1986a,c,d, 1988; Drew 1983; Drew and Abel 1985, 1988a; Colin 1986;
Colin et al. 1986). For the other Halimedaceae there seem to be no similar
analyses of distribution, although extensive information on the distribution of
Tydemania at Enewetak is provided by Gilmartin (1960, 1966) and Hillis-
Colinvaux (1977). Scuba diving has been an important tool in these studies, but
it is only with the availability of submersibles and remote photographic systems
that there has been the opportunity to examine considerably wider expanses of
entire systems, and thus obtain for broad areas oflagoon or fore-reef a baseline
quantitative analysis of distribution.
The densest covers ofliving Halimeda found so far occur in three principal
places ofa reef: 1) relatively shallow to very shallow (ca. 0.3 m) rock platforms,
2) lagoonal or back reef areas with an unconsolidated substratum of Halimeda
gravels, sand (sometimes mixed with fine rubble to which some of the plants may
be attached), and muds, and 3) vertical walls as represented by the fore-reef
slope, or sides of bommies (pinnacles). The first two are the locations of
meadows, the third of draperies. Percent cover at any of these sites can be very
high.
Halimeda Meadows
The biomass of Halimeda over many horizontal reef surfaces is so dense as to
resemble terrestrial meadows. The H. opuntia zone found in moderate energy
interislet or back reef environments only a few centimetres deep (Hillis-
Colinvaux 1980, 1985, 1986a) is an example in the first of the three preceding
categories of habitat. In such an environment H. opuntia produces tightly
branched clumps, and cover is 80-100%. Other species, with habit similarly
180 L. Hillis
modified to small-segmented compact bushes, provide dense covers in similar

habitats (Hillis-Colinvaux 1980). Halimeda communities on the Hattish tops of
lagoon pinnacles provide a second example of a dense cover over lithified
substrata; where observed at Enewetak these communities were of mixed
species, although H. macrophysa Askenasy provided most of the cover
(Hillis-Colinvaux 1977, 1980).
Dense populations of Halimeda growing on unconsolidated habitats in-
clude Halimeda meadows of the northern part of the Great Barrier Reef (Drew
and Abel 1985, 1988a), meadows in the lagoons of several atolls of the Marshall
Islands (Colin 1986 and personal communication; Hillis-Colinvaux 1988) and
an atomic bomb crater at Enewetak (Hillis-Colinvaux 1980). Meadows of the
northern Great Barrier Reef, occurring principally at depths of20-40 m but with
smaller ones at 100 m or more, cover 2000 km2 or more of sea bed, at least some
of which is a post Pleistocene Halimeda bioherm (Davies and Marshall 1985;
Orme 1985; Phipps et at. 1985). Although the communities include Penicillus
and Udotea, the predominant genus for the sites examined is Halimeda (Drew
and Abel 1985, 1988a). The three commonest species belong to section Opuntia,
with copiosa the most prominent. Halimeda biomass of up to 4 kg dry weight m- 2
is reported.
In the Marshall Islands, Halimeda meadows on unconsolidated substrata
have been discovered in the lagoons of Rongelap (ca. 36 m), Kwajalein (ca. 32
m and deeper) and Enewetak (ca . -21 and -25 m) Atolls (Fig. 12). All these
Fig. 12. Halimeda meadow of mixed species, in lagoon of Enewetak Atoll at approximately 22 m
depth (Photo by P. Colin; magnification approx. O.lx)
Recent Calcified Halimedaceae lSI
meadows, like the Australian ones, contain a mixture of Halimeda species. The
most prominent taxa in the Enewetak meadows are incrassata, cylindracea
Decaisne, near gigas W.R. Taylorlxishaensis Dong and Tseng, disco idea,
opuntia f. opuntia, and a small segmented, fine form of opuntia. Where the
substratum was sand and small fragments of coral rubble a lax incrassata often
predominated. Where it was sand or gravel mixed with small rocks the taxon
presently delimited as gigas I xishaensis was the most abundant macroalga.
Other Halimedaceae were not obvious. Percent cover was high (Hillis-Colin-
vaux 1980, 1986c, colour Fig. p. 47), and the contribution by the hold fast alone
to total carbon investment was also often high: over 40% where the rhizophyte
H. cylindracea predominated (Hillis-Colinvaux 1988). Also at Enewetak, but at
an interisland-back reef site on the north end of Runit Islet, a single species
meadow on a mud-silt substratum was located at the bottom (ca. -11 m) of the
atomic bomb crater 'Cactus'. There the rhizophyte species H. incrassata
provided close to 100% cover (Hillis-Colinvaux 1980, Fig. p. 260).
Extensive Halimeda banks recently discovered elsewhere, such as east of the
central Sunda shelf margin in the Java Sea (Roberts etal. 1987a,b) and the south
west Caribbean (Hine et al. 1988) suggest that dense groves ofliving Halimeda
will be found for many other parts of the tropical world ocean.
Penicillus Meadows
I have observed dense growth of Tydemania in the western Indian Ocean and at
Enewetak, but the areas covered have not been sufficiently dense or extensive to
be called meadows. Hence, the only other genus of the Halimedaceae known to
produce extensive dense stands is Penicillus. Sizeable groves were found at some
of my field sites in Jamaica (.Fig. 2), and Stockman et al. (1967) record densities
of76-108 thalli m- 2 and higher in southern Florida. It seems likely that similar
stands occur elsewhere in the Caribbean.
Halimeda Draperies
Vertical, pendant colonies and communities of Halimeda may, in some respects,
be considered vertical meadows. They represent a very prominent kind of
Halimeda development on reef slopes and walls, regions that traditionally, and
especially in the deep fore-reef, have not been considered habitats for Halimeda.
The term 'drapery' is used because the extensive pendant colony forms of
members of section Opuntia, especially the species copiosa, which may well
prove to be the predominant wall species globally, suggest traditional wall
hangings. Their long and trailing fronds drape down and across rock surfaces,
caverns and crevices, to develop festoons that can be the major benthic car-
bonate source of a slope.
Other high biomass taxa on slopes and walls include the Halimeda species
distorta (Yamada) L.H. Colinvaux and cryptica. As in the meadows a single
Halimeda species may predominate, or there may be two to several prominent
Halimeda taxa. On the fore-reef of Jamaica, at -25 to -100 m H. cryptica, an
unusual uniaxial species predominates, but is often associated with copiosa (T.F.
182 L. Hillis
Goreau and N.1. Goreau 1973). At San Salvador, Bahamas, H. copiosa provides
up to 39% cover from -117 to -130 m but is associated with three other Halimeda
species (Littler et al. 1986). Halimeda copiosa also develops extensive hangings
on the sloping walls ofbommies (pinnacles) associated with Davies Reefin the
Great Barrier Reef(personal observation), and was one ofthe prominent species
collected from the deep fore-reef of Enewetak Atoll (Hillis-Colinvaux 1985,
1986a,d).
Other distinctive habits are exhibited by the wall-growing halimedas
(Hillis-Colinva ux 1985). Especially noteworthy in terms of biomass represented
were the large single species clumps or tussocks produced by H. distorta at about
-100 m (Hillis-Colinvaux 1986c, colour photograph p. 47). A single tussock
collected by submersible had a dry weight of 682 g.
Overall vertical cover of the fore-reef can be high. At Enewetak Halimeda
populations were found to cover 10-50% or more of the atoll slope down to
110 m, with at least a third of the species growing to more than 90 m (Hillis-
Colinvaux 1986a,d). At this depth cover by hermatypic corals did not exceed 1%
(Colin et al. 1986; Hillis-Colinvaux 1986a). The genus continued down to at least
-140 m. The full extent of its cover was, however, difficult to determine during
submersible observations and from photographs because sediments blanketed
extensive areas of the fore-reefslope. Sediment cover was conspicuously heavier
on the leeward side of the atoll.
There are essentially no data on distribution or biomass of the other
Halimedaceae on vertical surfaces. At Enewetak only Udotea and Tydemania
are known (Taylor 1950). Two species of Udotea grew to at least -138 m, and the
diminutive U. near javensis (Montagne) A. and E.S. Gepp was relatively
abundant on likely substrata. Udotea biomass, however, was insignificant
compared to the Halimeda contribution. No Tydemania thalli were located on
the fore-reef wall.
3.2 Yield of Carbonate Sediments
Most ofthe material shed as debris by these genera has a high as well as relatively
uniform carbonate content (Hillis-Colinvaux 1980, Tables pp. 268-269, 27 I) but
the rate at which a thallus yields sediments varies. Consequently, the major
question of concern to workers deriving their data from living material is the
number of generations, or of 'turnovers', produced annually. Turnover time
herein is considered to be the length of time taken to replace an individual. In
the context of carbonate production this definition, based on Lindeman's
analysis of energy flow in a Minnesotan lake (1942), does not have the problem
inherant in Lindeman's application of it to primary production since respiratory
losses are not coupled to carbonate production. Losses of parts of a plant from
grazing or from other causes must, however, be considered.
Data on generation times for the Halimedaceae are very limited. In labo-
ratory aquarium systems I found that Penicillus capitatus Lamarck replaced
itself about every 1.5-2 months (Colinvaux et al. 1965), a generation time in
accord with the field results of Stockman et al. (1967). Udotea flabellum (Ellis and
Solander) Lamouroux in the same laboratory system replaced itself in 6-9
months. Halimeda field census data from Jamaica and laboratory observations,
mostly on sand-growing species, indicate an annual turnover time of 112-3
generations, although some laboratory H. incrassata thalli replaced themselves
in 2 months. Field data based on number of segments gained rather than entire
thalli provide turnover times of39 days of populations of H. incrassata and H.
monile (Ellis and Solander) Lamouroux in Nonesuch Bay, Antigua W.I. (Multer
1986, 1988); about 1 year for H. opuntia in the Marquesas Keys, Florida, depth
< 2 m, (Hudson 1985); and 15 days for H. opuntia and H. copiosa combined at
Davies Reef, Great Barrier Reef, depth 5-8 m (Drew 1983). The different
approaches used to obtain generation-turnover times for Halimeda complicate
the evaluation of the resulting data; some of the techniques are mandated,
however, by the different habits of the plants. Species that are predominantly
sprawlers such as H. opuntia and H. copiosa do not have the discrete life form
usually exhibited by rhipsalian (rhizophyte) halimedas or Penicillus which
makes distinguishing different generations more straightforward. For turnover
figures to be used with confidence it is obvious that many more data are needed,
and that more species and a much wider range of habitats should be examined.
3.3 How Much Carbonate Does Halimeda Produce?
Data on carbonate Halimeda has become available in recent years from two
major sources: from biologists and geologists studying the growth ofliving thalli
and calculating total productivity based on biomass and turnover or genera-
tional data (Merten 1971; Hillis-Colinvaux 1974, 1980; Drew 1983; Abel and
Drew 1985; Drew and Abel 1985; Hudson 1985; Multer 1988), and from
geologists analyzing sedimentary deposits with seismic profiling and side-scan
sonar techniques, together with radiocarbon dating of reef cores to determine
vertical accretion (Davies and Marshall 1985; Orme 1985; Phipps et al. 1985;
Roberts et al. 1987a,b). Translated into rates of vertical accretion most of the
figures are in the range of 0.18-5.9 m 1000 years· I .
Different methodologies, environments, population sizes, and possibly
species differences account for this broad diversity, with the lowest of the figures
given (I have excluded figures one to two orders of magnitude below the above
range) resulting in part from a relatively low percent cover (= ca. 21%,
Hillis-Colinvaux 1974, 1980). Ofthe data available only those based on 'annual
contribution per single thallus' (Hillis-Colinvaux 1980; Multer 1988) provide a
ready basis for modelling and prediction. Thus, if a single rhipsalian thallus
produces 0.8 g carbonate yeac I (Hillis-Colinvaux 1980, considered conserva-
tive), a 60% cover (= 600 H. incrassata thalli, cf. Hillis-Colinvaux 1980, p. 274)
by the same species under similar conditions, a density which is still below that
of Halimeda meadows, would produce 0.4 m of carbonate vertical accretion in
1000 years. The porosity used for the calculations is 60%, following Stockman et
al. (1967). In the same way, a cover of 100% Halimeda, which approximates
184 L. Hillis
cover in the H. opuntia zone of Enewetak and the Halimeda meadows at -21m
in the lagoon west of Enewetak Islet, would yield a vertical accretion of 0.7 m in
1000 years. (The reader should be aware of two textual errors in earlier
publications which may be confusing. Hillis-Colinvaux 1980, p. 274, text line 12:
4066 mm year- 1 should read 0.066 mm year-I. Hillis-Colinvaux (l986c, p. 47, first
column last line) should read 7 cm 1000 years-I; and second column lines 1-2
should be 'years for a low cover, by sand growing species, to 3 meters'.}
The highest figures in the range provided in paragraph one are based on
cores taken from Halimeda banks east of the central Sunda Shelf margin in the
eastern Java Sea (Roberts et al. 1987a,b), and the northern Great Barrier Reef
(Davies and Marshall 1985). Both groups of workers obtained data indicating
rates of vertical accretion that increased throughout the Holocene, with average
rates for the time spans represented in the cores analyzed being 2.8 and 1.75 m
1000 years- 1 respectively. Both groups also suggest that the implied rapid growth
rates of Halimeda are related to nutrient enrichment from upwellings.
Although export or import of sediments may be involved in the develop-
ment of some Halimeda bioherms, those of the northern Great Barrier Reef are
considered to represent in situ accumulations of Halimeda, some of which are
presently topped by meadows ofliving Halimeda (Drew and Abel 1985). They
provide strong evidence for the reef-building activity and ecological success of
at least one genus of the Halimedaceae throughout the Holocene.
3.4 Major Accretion Events
Biotic and abiotic factors producing widespread or catastrophic death of the

plants lead to major episodes of accretion. Likely factors include temperature
and salinity stresses, disturbances resulting from tropical storms, sexual re-
production, and possibly epidemic infestation by cyanobacteria. Only a brief
elaboration of two biotic factors can be included in this paper, but it will serve
to underscore the potential of certain events. The first example, derived from
field studies in Bermuda, Jamaica and Enewetak, is the observation of extensive
populations of Halimeda shrouded by cyanobacteria (Hillis-Colinvaux 1980,
Fig. 72). Since similar events in culture have led to the death of the plants or else
a shedding of branches with subsequent regrowth, both entailing sediment
production, these same patterns are predicted for field populations.
The second example is based on field and laboratory observations of sexual
reproduction, an event which now has been reported for all five genera and
many Halimeda species (Hillis-Colinvaux 1959, 1973, 1980, 1984; Meinesz 1980
and personal communication; Drew and Abel 1988b), although its very exis-
tence for some of the genera was questioned until relatively recently (Dawson
1966). Death follows immediately after the formation of gametangia (Figs. 9,
10), the outpouring of the energy reserves of the entire plant into the gametes,
and their subsequent release (holocarpy). The process from the first readily
observable stages of gametangial formation to release of gametes occurs in
-36h. A widespread occurrence of sexual reproduction would therefore be
a major sediment-producing event. And, since the cytoplasmic contents of the

algae are channelled into gamete production, only the framework materials
of the filaments remain. Shed segments from sexually reproducing plants,
therefore, are essentially skeletal.
3.5 Impact of Halimedaceae on the Reef Mass of Modern

and Holocene Reefs
As one moves across a classical tropical reef composed of lagoon or back reef,
algal ridge-spur and groove, and fore-reef, the principal halimedacean (and
other) components change (Fig. 8). As noted earlier, percent cover by
Halimedaceae is essentially zero on the ridge and relatively low on most spur
and grooves. Cover by the Halimedaceae and principally by Halimeda can,
however, be very high in the other two major regions.
Reefmass ultimately depends on the rates of accretion in these three major
regions which, over geological time, are affected by sea level. During lowered sea
levels, for example, lagoons may become extinct, with a corresponding
decreased contribution by the halimedas living on the floor and on pinnacle
walls. Delimitation of different physiographic regions in a carbonate model of
a reef allows differences in rates of accretion to be considered; it also recognizes
the importance of the extent of their areas. Thus, in reefs with lagoons of
considerable size such as Enewetak and Bikini Atolls, lagoonal populations are
likely to contribute more than half the reef mass, even with accretion rates
significantly lower, as obtained by Smith and Kinsey (1976; 0.8 kg m- 2 year- t for
lagoon,4 kg m- 2 year- t for reef ridge).
In reef systems where accretions of lagoon and fore-reef are most likely to
be from Halimeda it is reasonable to suggest that the origin of reef mass depends
on the relative rates of accretion on lagoon floor, reef ridges (by their populations
of corals and coralline algae), and fore-reef as expressed by:
Rm = (HsL + RvRh + HsCdF)t,
where Rm = reef mass, Hs = deposition of Halimeda segments per unit area,
L = area of lagoon, Rv and Rh = vertical and horizontal components of reef
ridge growth, Cd = deposition of coral debris per unit area, F = area offore-reef
wall, and t = time. For Halimeda bioherms, since the ridge and fore-reef
elements are absent, the expression can appropriately be shortened to:
Rm = (HsL)t.
4 A Classification of Tropical Reefs by Principal Components
The predominant components of tropical reef systems are photoautotrophs, so

it seems especially quixotic to continue to use the term 'coral reefs' in scientific
publica tions. Recently I argued that 'algal reefs', a name which incl udes not only
186 L. Hillis
calcareous algae but also the symbiotic components and fleshy algae, more
accurately reflects both the ecological and physical basis of reef systems
(Hillis-Colinvaux 1986c). The evidence presented in this chapter, drawn from
both present and historical records, provides additional support for the use of
algal reef, or alternatively of coralgal reef in naming this tropical carbonate
system.
Halimeda, although one of the principal components of many Recent reefs,
is not, however, equally abundant in all tropical algal reefs. For reasons not
understood but which may be related to problems of successful dispersal, both
species diversity and biomass of Halimeda (and the other calcareous
Halimedaceae) are greatly reduced on certain reefs. There are, in other words,
Halimeda and non-Halimeda reefs. The absence of Halimeda from some reef
systems has important implications in the ecological and accretionary processes
of these reefs.
A cknowledgements. A significant portion of this paper was written while I was a Bunting Institute
Science Scholar of Radcliffe College at Harvard University. The scholarly and convivial support of
the Harvard and Bunting communities, and the financial support of the Office of Naval Research
for the Science Scholar Program are gratefully acknowledged. I also thank Robert Riding for
thoughtful discussion of the early history of the calcareous Bryopsidales, and the stimulus thereby
provided me for developing an analytical interface between living and fossil Halimedaceae.
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Chapter 11
The Genus Concept in Charophyta: Evidence from
Palaeozoic to Recent
M. FEIST l and N. GRAMBAST-FESSARD 1
Abstract
In fossil Charophyta, the generic criteria are based on characters of the calcified fructifications
(gyrogonite or utricle). However, the classification of living forms is based mainly on vegetative
parts. An alternative system employing gyrogonite characters might permit the classification of both
living and fossil members of the Characeae. Thus, the systematics of fossil Charophyta may be
considered as founded on natural taxa and not on organ-genera. Examples of Palaeozoic to Recent
representatives demonstrate the various criteria used to distinguish genera in the different families.
Phylogenetic lineages of species allow the recognition of phyletic genera, especially in the Clava-
toraceae. Relationships between genera outline the main phylogenetical trends in Charophyta. Two
new suborders are established as well as two new combinations concerning species figured in the
present paper: Palaeocharineae nov. subordo, Charineae nov. subordo; Nitellopsis (Tectochara)
morulosa n. comb.. Porochara sardiniae n. comb. In addition, the revision of all charophyte genera
has led us to put nine genera into synonymy: Minhechara Wei = Latochara Madler; Paracunea-
tochara Wang = Leonardosia Sommer; Caucasuella Kyansep-Romashkina = Aclistochara Mad-
ler; Gobichara Karczewska & Ziembinska-Twordzydlo = Microchara Grambast; Piri{ormachara
Liu & Wu = Mesochara Grambast; Qinghaichara Yang = Nitel!opsis Hy; Retusochara Grambast
= Peckisphaera Grambast; Sinochara Lin& Wang = Peckichara Grambast; Songliaochara Wang,
Lu & Zhao = Nodosochara Madler; Turbochara Wang = Nodosochara Madler; Tolypella (AI.
Braun) Leonhardi, pro parte = Sphaerochara Madler emend. Horn af Rantzien & Grambast.
1 Introduction
In Charophyta, the systematics offossil forms are based on the morphology of

the calcified female gametangium (oosporangium), called the gyrogonite,
whereas the classification of extant species is based on the vegetative parts.
Gyrogonites range from 200 to 2000 /Lm and can be investigated only by means
of optical and scanning electron microscopy. On the other hand, the size of the
thallus, from 1.5 to 200 cm (Wood and Imahori 1965), permits direct obser-
vations. The vegetative parts, always present in the living plant, even in species
that do not calcify or reproduce vegetatively, are commonly missing in the fossil
which predominantly comprises isolated gyrogonites. According to Horn af
Rantzien (1959), the gyrogonites do not represent the entire plant and should
therefore be classified as organ genera and organ species.
However, the fundamental structure of the oosporangium is similar in
extant and fossil representatives (Fig. 1). Oosporangia are spherical bodies with
1Laboratoire de Paleobotanique, URA n° 327 du CNRS, Universite des Sciences et Techniques du

Languedoc, 34095 Montpellier, France
190 M. Feist and N. Grambast-Fessard
,........ ...."
\
\
" \ I
/",'
I
I \I ,
: r --- c
8 p
Fig.IA,B. Chara hispida L. (Characeae). From Grambast 1958 (unpublished thesis). A Gyrogonite
(calcified oogonial ofa Recent specimen (published in Soulie-Marsche 1973); Bsection of the same:
m oospore membrane; a starch grains; Ca calcified part of the spiral cells; b basal plate (calcified
sister-cell of the oosphere); n nodal cell; p basal pore. In dOlled lines, uncalcified part of the spiral
cells and of the coronular cells (e), not preserved around the gyrogonite (unpublished)
a multicellular wall enclosing the oospore. The basal pore is obstructed by one
to three sister cells of the oosphere which constitute the basal plate. In species
which calcify, the multicellular wall, as well as the sister cells of the oosphere
become particularly resistant. In addition, the thickened membrane sur-
rounding the oospore may be preserved in the fossil state both in calcified
species, such as Rhabdochara praelangeri Castel (Fig. 4c), and in uncalcified
ones, such as Nitellites sahnii Horn afRantzien from the Jurassic ofIndia (Horn
afRantzien 1957). The vegetative parts are seldom preserved as fossils, but when
they are found, they do not differ significantly from those of extant species and
occur as long internodal cells alternating with short nodal cells. A new approach
to the genus concept is given by the use of electrophoresis in the living forms.
Applied by P. Gemayel (1988) to the genera Chara , Nitella and Tolypella, this
method totally confirms the validity of the generic criteria based on the gy-
rogonite morphology.
In this study, the first part is a survey of the successive morphological types,
from the Upper Silurian to Recent, which will give an outline of the main
distinctive criteria on which orders and families are based. Secondly, the various
generic criteria are defined by taking examples from different families and
periods, including examples ofliving Characean genera also represented in the
fossil state. This will help us to decide if the fossil charophyte genera should be
considered either as parataxonomic or as natural taxa.
The Genus Concept in Charophyta 191
2 Bases of Classification
The classification of charophytes is based on hierarchical characters which

reflect the phylogeny of the group. Evolution started with the originally vertical
gyrogonite cells spiralling either dextrally or sinistrally and reducing in number.
Subsequent modifications led to improvement in the enclosing and protection
of the egg. This took place in one of two different ways, either by the acquisition
of a supplementary cover around the gyrogonite or by the closing of the apical
pore of the gyrogonite.
In the current classification (see Appendix), the orders are based on the
orientation of the gyrogonite cells, i.e. vertical, dextral spiral or sinistral spiral.
The distinction of families is based on the number of the cells and, among the
Charales, either on the presence of an utricle (Clavatoraceae), an apical pore
always open (Porocharaceae), an operculum (Raskyellaceae) or a closed apex
(Characeae). The generic criteria are based on particular characters of the
gyrogonite apex, base, basal plate and the general outline of the gyrogonite.
Species are based on special characters of gyrogonite shape, ornamentation and
dimensions. In the Clavatoraceae, characters of the \ltricle are also taken into
account.
3 Successive Morphological Types
Praesycidium siluricum (Sycidiaceae), described by T. Ishchenko and A. Ish

chenko (1982) from the Upper Silurian (Pridoli) of Podolia (Ukraine, USSR) is
the oldest charophyte species presently known. The first and most important
stage of diversification occurred during the Devonian, when the three orders of
Charophyta were already represented (Fig. 2). During the Permian, Triassic and
Jurassic, gyrogonites were discrete forms, generally of small dimensions and
devoid of any ornamentation, but with high evolutionary potential. Thus, the
gyrogonite of the Triassic Porocharaceae Stellatocharoideae type, with its apical
pore at the end of a neck, or collar, is found again inside the utricle of the
Jurassic-Cretaceous Clavatoraceae. On the other hand, Porocharaceae gave rise
to the Characeae and Raskyellaceae by two different methods of closing the
a pical pore (F eist and Grambast-F essard 1984), namely by joined spiral cells or
by intercalation of supplementary opercular cells.
A second stage of diversification took place in the Lower Cretaceous, with
the rapid radiation of the Clavatoraceae which are characterized by the
evolution of the utricle. Grambast (1974) put forward three evolutionary
lineages and presented a phylogeny of the family. These lineages are amongst
the most striking of the plant kingdom.
A third diversification occurred in the Upper Cretaceous (Campanian-
Maastrichtian) with the expansion of the Characeae. Following the extinctions
at the Cretaceous-Tertiary boundary (Feist 1979) and a last diversification
during the Paleogene, the group regressed until the present state, with only six
extant genera, grouped in one family, the Characeae. In contrast to the fossil
MIOCENE
OLICZOCENE
1 . Pinnoputamen
EOCENE 2· Sycidium
~e 3 • Chovanclla
22
4. Trochiliscus
. I . 5 . Gcmmichara
~. 6· Eochara
7 • Palaeochara
8 • Stomochara
CRETACEOUS 9· Leonardos.ia
126 10· Stellatochara
11 • Latochara
12 • Echinochara
______ 13· Perimneste
14· Atopochara
~ 25
15 . Globator
JURASSIC '6 16 • Clava tor
'2
17· Flabellochara
1I~ 18 . Triclypella
19 • Embergerella
20 . Septorella
TRIASSIC 21 . Porochara
.t 22 . Raskyella
23 • Sphaerochara
PERMIAN 24 . Tolypel1a
.~ 9~ 25 . Aclistochara
26 • Amblyochara
27 • Platychara
CARBON IFEROUS ~8<® 28 . Dughiella
29 • Nilellopsis
30 . Harrisichara
7- 31 • Maedleriella
32 . Gyrogona
33· Rhabdochara
DEVONIAN ~6¢f!i; ",. .".R.e... 34· Chara
35 • Lamprolhamnium
Fig. 2. Structural evo lution ofthe fru ctifica tion in Charoph yta ( part ly f ro mGra mbast 1974)
record, the extant charophytes show structural and taxonomic impoverishment

(although the number of living species reaches 80 according to Wood and
Imahori 1965) and loss of calcified structures, such as ornamentation and
calcified multipartite basal plate.
4 The Generic Concept from the Fossil Record
The generic criteria vary in importance and significance in the different periods
and families as is shown by the following examples taken from the Palaeozoic
families as well as from the Clavatoraceae and Characeae.
4.l Palaeozoic Families
In two monotypic Palaeozoic families, the genera Eochara Choquette (Fig. 2,

No.6) and Palaeochara Bell (Fig. 2, No.7) differ in the number of gyrogonite
spiral cells (being more than six and just six respectively). In all post-Palaeozoic
charophytes, which are here placed in the new Suborder Charineae (see
Appendix), the number of spiral cells is fixed at five.
4.2 Clavatoraceae
The seventeen genera of the Clavatoraceae are well-defined by the characters of

the utricle. Grambast (1966, 1967, 1970, 1974) reconstructed inter- and in-
trageneric lineages linking successive stages of evolution. These lineages may be
illustrated by two examples:
1. Perimneste-A topochara lineage (from Grambast 1967, 1974; see Fig. 3a,b,c).
The two genera Perimneste Harris and Atopochara Peck seem at first sight quite
different, both in shape and size as well as in structure. In the Berriasian P.
horrida Harris (Fig. 3a), the antheridial casts are recognizable in hollows at the
utricle surface. Dissolving the calcite with weak acetic acid shows the position of
the utricle cells, which are grouped in three distinct clusters. From the Berriasian
to the Cenomanian, these clusters progressively condense and the number of
antheridial casts reduces. This proceeds until the typical structure of A topochara
is realized in A. trivolvis Peck of the Barremian, where the central basal cell of
each cluster has disappeared and the antheridia sites are reduced to one sterile
cell (Fig. 3b). From the Barremian to the Cenomanian, the cells of antheridial
origin disappear altogether and the upper cells become strongly spiralized. The
end member of this progressive evolution is seen inA. multivolvis Peck (Fig. 3c).
This progressive series of structural modifications shows how these forms
may be linked; the separation between the two genera being placed, perhaps
arbitrarily, at the appearance of A. trivolvis, when the cells are joined together
and the antheridia really disappear.
Fig.3a-f. Clavatoraceae. a Perimneste horrida Harris; Hils, NW Germany, Berriasian; basal view,
x33. b A topochara trivolvis Peck ssp. triquetra Grambast; Warlingham Borehole, S England, Lower
Barremian; lateral view, x40 (BGS. MPK 5875). c AlOpochara multivolvis Peck ; Le Revest, SE
France, Cenomanian, lateral view, x32 (Feist 1981). d Globator trochiliscoides Grambast: EI
Mangraner, ESpain. Barremian: lateral view. X32. e G/obator nurrensis (Pecorini) Grambast: Cala
d'Inferno, NW Sardinia. Berriasian: lateral view, X30. f Globator maillardi (Saporta) Grambast:
Broadoak Borehole, SEngland. Berriasian: lateral view, X50 (BGS. MPK5876)
2. Globator lineage (from Grambast 1966, 1974) (Fig. 3d,e,f). This series shows
the evolution from the Berriasian Globator maillardi (Sa porta) Grambast to the
Barremian G. trochiliscoides Grambast, by the reduction in the number of the
utricle cells and by the progressive acquisition of spiralling.
In this case, all the species are attributed to the same genus, because the
successive evolutionary stages represent modifications of the same basic
structure. However, if the end-members of the lineage had been found in-
dependently, they would probably have been attributed to two different genera
as was the case for Perimneste and A topochara. Globator may be considered as
a phylogenetic genus, in the sense that it is composed of a continuous lineage of
successive species.
5 Characeae
In the previous examples, generic distinction did not present particular

difficulties. This is not the case with the very homogeneous family Characeae
which includes about fifty genera, comprising nearly half of the presently
recognized genera of charophytes. The family is first undoubtedly known from
the Middle Jurassic (Feist and Grambast-Fessard 1982), and persists as the only
extant family.
In this family, the genera are distinguished not according to evolutionary
stages as in the Clavatoraceae, but on the basis of a. series of more or less
important morphological characters. The first most important character relates
to details ofthe structure ofthe apical part ofthe gyrogonite, such as the presence
or absence of a peria pical groove, the presence or a bsence of apical nodules, and
variations in the width ofthe cells around the apex. Another important character
is the structure of the basal plate, which represents the calcified sister-cells of the
oosphere. Observing these characters is not easy because of their minute size and
the problem is often made more difficult by the degree of calcification or
preservation of the gyrogonite. However, some of the genera of the Characeae
are well characterized and appear to constitute natural taxa, such as in the
following examples:
5.1 Genus Harrisichara Grambast
The gyrogonites in this genus are characterized by an undifferentiated apex, a

distinct basal column and, in most species, ornamented spiral cells. Harrisi-
chara, which includes more than twenty species, is reported from the Palaeocene
to the Lower Oligocene. In the Palaeogene sequences from the Isle of Wight
(southern England), an evolutionary sequence from H. vasijormis to H.
tuberculata can be clearly recognized (Feist-Castel 1977, Pl. 21, Figs. 1-5).
In this case, only fossil forms are concerned. Connections have, however,
also been established between some fossil gyrogonites and those of extant
genera, as is shown by the following examples.
5.2 NiteUopsis and Tectochara (Fig. 4a-b, d-f)
The former genus Tectochara Grambast, established for fossil species and
known from Palaeocene to Quaternary, is characterized by a decrease in both
the th ickness and width ofthe spiral cells in the peria pical zone and by a very thin
basal plate.
The extant, but rather rare, Nitellopsis Hywas poorly known for a long time.
The plant lives in relatively deep water (4-12 m), is dioecious and reproduces
mostly vegetatively by means of bulbiIs, so the female reproductive organs are
rarely encountered. Krassavina (1971), studying fragments of thallus and
calcified oogonia coming from the gut contents of ducks killed along the Azov
Fig.4a-f. Characeae. a Nitellopsis obtusa (Desvaux) Groves; Sarthe, NW France, Recent; lateral
view. x47. b Nitellopsis obtusa (Desvaux) Groves; Bordeaux. SW France. Recent ; bulbil. x92. c
Rhabdochara praelangeri Castel ; Marseille, SE France, Chattian; outer view of the oospore
membrane (ectosporostine), x600. d Nitellopsis (Campaniella) helicteres (Brongniart) Grambast &
Soulie-Marsche; Mont-Bernon, NE France, Sparnacian ; apical view, x45. e Nitellopsis obtusa
(Desvaux) Groves; same locality as a, Recent; apical view, x47. f Nite/lopsis (Tectochara) morulosa
nov. comb.; Saint-Gely-du-Fesc, SFrance, Cuisian-Lutetian; lateral view, x40; fossil representative
of the genus with spiral cells bearing tubercles (Feist-Castel 1972, PI. I, Fig. 5)
Sea (Ukraine, USSR), recognized the identity of these gyrogonites, belonging to

Nitel/opsis obtusa (Desvaux) Groves, and of Tectochara diluviana (Madler)
Grambast.
In this example, it has been possible to connect the living representative with
its fossil ancestors that are the Tertiary species formerly assigned to Tectochara;
this taxon is now considered as a subgenus of Nitellopsis.
5.3 TolypeUa and Sphaerochara (Fig. 5d-f)
Another example of the relationships between fossil and living charophytes can
be seen hy examining the characters of the basal plate of the gyrogonite.
Grambast (1956) showed that the morphological aspects of this plate could be
Fig. Sa-f. Porocharaceae a nd Characeae. a Aclistochara bransoni Peck; Elk Creek. USA . Upper
1urassic; apical view, x66. b Lamprothamnium papulosum Groves; Marseillan. S France. Recent;
apical view, x150. c Porochara sardiniae nov comb.; Cala d'lnferno. NW Sardinia. Berriasian; apical
view. x50 (Colin et al. 1984). d and e Tolypella pumila Grambast ; Hoogbutsel. Belgium. Lower
Oligocene; lateral view. x110 and isolated bipartite basal plate. x324. f Sphaerochara intricata
(Trent. ex Roth) Leonhardi n. comb. ; 1uignee. W France. basal view showing the undivided basal
plate. x90
used to recognize the genera within the Characeae. Subsequently Daily (1969)
and Sawa and Frame (1974) showed that the basal plate could be either simple
or multipartite in the living species of Tolypella, each of these types of basal plate
being characteristic of a section of the genus. Thus, this character is clearly of
taxonomic value; in the present classification ofthe living forms, however, itcan
define only subgeneric divisions. Applying this criterion, the fossil genus
Sphaerochara Madler emend. Horn af Rantzien and Grambast must be in-
cluded in the Nitelloideae (Feist and Grambast-Fessard 1982) since its gy-
rogonites are identical to those of Tolypella (AI. Braun), corresponding to the
section Rothia R.D. Wood; both have a simple calcified basal plate, and the two
taxa should thus be considered as synonymous, with Tolypel/a section Rothia
falling into the synonymy of Sphaerochara (see Appendix).
5.4 Lamprothamnium and Aclistochara (Fig. Sa-c)
The gyrogonites of both the Cretaceous-Recent Lamprothamnium Groves and

the Jurassic-Cretaceous Aclistochara Peck possess a deep peripheral groove at
the apex and are cylindrical and unornamented. Because of these similarities,
Soulie-Marsche (1989) considered the two genera to be identical. However, the
narrower apical zone of Aclistochara, resembling the small size of the apical
opening of the Porocharaceae (Fig. 5c), favours the maintenance of two distinct
genera (Feist and Grambast-Fessard 1982).
The examples taken from the Characeae (see also Fig. 6) demonstrate that
the study of the gyrogonites offossil forms may be useful for the recognition and
definition of present genera. Care must be taken, however, that rationalization
of the nomenclature of fossil and extant genera through the recognition of
apparent synonymies in temporally widely separated gyrogonite taxa does not
mask distinct stages in evolution.
6 Conclusions
As in the higher plants, the classification of the charophytes can be based on

characters of the female reproductive organs, which repre.sent the main part of
their fossil remains. The classification used here reflects the phylogeny of the
group and it expresses the main evolutionary tendencies as well as relationships
between genera and species. The gyrogonites may thus be considered as
characteristic of natural taxa rather than merely representing organ-genera. For
this reason, it is possible to try to harmonize the systematics offossil and living
forms and to look for a natural classification for the whole phylum.
Appendix: Systematic Section
1. Taxonomy
Since the last publication ofa complete classification (Tappan 1980, p. 955), two
.new families have been formed (Pinnoputamenaceae and Karpinskyaceae). In
the present paper, a new name is proposed for the Karpinskyaceae and two new
suborders are established: Moellerinaceae nom. nov. = Karpinskyaceae Wang
and Lu (1980, p. 196). Moellerinaceae is proposed to serve as a substitute for
Karpinskyaceae established by Wang and Lu (1980) as it is based on the earliest
described genus Moellerina Ulrich 1886 which also possesses the most typical
characters of the family; the presence of coronular cells in Karpinskya is a
particular character. Two new suborders, based on the number of cells com-
posing the gyrogonite, are introduced in order to present a more logical
classification: these are Palaeocharinae and Charinae (see Charophyta, below).
There are two new combinations relating to specimens figured in this
present paper:
Time (I: CHARACEAE NITEUOIDEAE ............

in / \ --.......
MY Series I Epoch To/ypel/a (SplJaeroclrara) To/ypel/a (Tolypella) Ni/ella
BP
Reoent calcified , uncalcified 9
o I
I
I
Pleistocene
I
Pliocene
I
,
el~
5,1 I
,I I
I
I
I I
Miocene I I
I I
I I
23,5 I I
I I
Oligocene I
I
.
36,5 I
I
I
I
Eocene
I
M,O
Pal90Cena
66,5
...
\
I
I
...
Upper I
..
Cretaceous .'
..- .
I
I
.'
I
97,0
~.
.
\
-100
lower
'.
,, ,
Crelaceous
~ ,
140.0
Upper
//
JuraSSic /
,
,,
I
160,0
l::
,,
Middle
.,,
Jurassic
184,0
lower *]9
210,0
Jurassic :~2 ,,
I
I
,/
Triassic .' I
POROCHAROIDEAE STEUA TOCHAROIDEAE
250,0 POROCI-IARACEAE
.0 Undivided basal plate 8 Multipartite basal plate

Fig. 6. Phylogeny of the post-palaeozoic charophytes. Apical structure and basal plate are con-
sidered the main criteria ; thus Tolypella is interpreted as polyphyletic. 1 Geological scale from Haq
and Van Eysinga (1987) 1-12 Lateral and apical views of selected genera. 1 Stellatoehara; 2
Latoehara; 3 Tolypella; 4 Atopoehara; 5 Septorella; 6 Feistiella; 7 Sphaeroehara; 8 Aclistochara; 9
Platyehara; 10 Dughiella ; 11 Chara; 12 Raskyella
Nitellopsis (Tectochara) moru/osa nov. comb. = Tectochara moru/osa

Feist-Castel1972, p. 2, PI. 1, Figs. 1-7.
Porochara sardiniae nov. comb. = Musacchiella sardiniae Feist and
Grambast-Fessard, in Colin et aI. 1984, p 348, PI. 1, Figs 6-13.
On the other hand, the reexamination of all charophyte genera for the
"Treatise on Invertebrate Paleontology" has led us to put several genera into
synonymy; they are indicated as follows, with new combinations based on the
type-species of each genus:
a. Porocharaceae
Minhechara Wei, based only on the wider apical pyramidal projection, is
not distinguishable from Latochara Madler 1955. Latochara columelaria
nov. comb. = Minhechara columelaria Wei in Hao et al. 1983, J. Wuhan
ColI. Geol., p. 174, PI. 43, Figs. 11-17.
Species from the Permian of China assigned to Paracuneatochara by Z.
Wang (1984) consist of isolated gyrogonites with the same characters as the
Brazilian Leonardosia Sommer 1954, known from casts only. Both have the
same apical morphology, with a long apical neck ending with the apical
pore. In addition they both are of Permian age.
Leonardosia jinxiensis nov. comb. = Paracuneatochara jinxiensis Z.
Wang 1984, Acta Micropalaeontol Sin., p. 56, PI. 2, Figs. 1-15.
b. Characeae
- Charoideae
Caucasuella Kyansep-Romashkina 1980 is not distinguishable from
Aclistochara Madler 1955 because both have the same apex, which is
sunken and convex in its centre. In addition, the genus Jurella, which is
considered by Kyansep-Romashkina as closely related to Caucasuella, has
been attributed by Liu (1982) to Aclistochara.
Aclistochara gulistanica nov. comb. = Caucasuella gulistanica Kyansep-
Romashkina 1980, Limnobios, Akad Nauk USSR, p. 81, PI. 1, Figs. 4-7.
Gobichara Karczewska and Ziembinska-Twordzydl0 1972 must be con-
sidered as a junior synonym of Microchara Grambast 1959, because they
are based on similar diagnostic characters: charoid apex, small size, com-
mon development of ornament.
Microchara deserta nov. comb. = Gobichara deserta Karczewska and
Ziembinska-Twordzydlo 1972, Palaeontol. Pol., p. 73, PI. 15-16.
Piri[ormachara Liu and Wu 1985 differs from Mesochara Grambast 1962
only by spiral convexity, a character which may depend on the degree of
calcification and which cannot be considered alone as a generic criterion.
Mesochara gumudiensis nov. comb. = Piri[ormachara gumudiensis Liu and
Wu 1985, Bull. Inst. Geol. Chin. Acad. Sci., p. 148, PI. 1, Figs. 4-7.
Qinghaichara Yang in Hao et al. (1983) presents the same diagnostic
characters as Nitellopsis: spiral cells "becoming narrower and remarkably
thinner at apical periphery, apical rosette produced by inflated ends of
spiral cells", basal plate thin ("plate-like"), "funnel-shaped outer depres-
sion around the basal orifice". .
Nitellopsis o valis nov. comb. = Qinghaichara o valis Yang in Haoet al. 1983,
J. Wuhan Coil. Geol., p. 156, PI. 36, Figs. 1-4.
Retusochara Grambast 1971 and Peckisphaera Grambast 1962 have similar
apical structures, with spiral endings which are slightly enlarged and a thick
basal plate. The larger size of Retusochara cannot be considered significant.
Peckisphaera macrocarpa nov. comb. = Retusochara macrocarpa Gramb.
1971, Paleobiol. Continent, p. 31, PI. 14-15.
All the salient features of Sinochara Lin and Z. Wang are found in
Peckichara Grambast 1957, namely an apex with nodules, surrounded by
a weak periapical furrow.
Peckichara rudongensis nov. comb. = Sinochara rudongensis Lin and Z.
Wang 1982, in: S. Wang, Huang Z. Wang et aI. GeoI. PubI. House, Peking,
p. 32, PI. 15, Figs. 5-9.
SongUaochara Z. Wang, Lu and Zhao 1983 and Turbochara Z. Wang 1978,
which differ only in the smaller size of the latter, are to be considered as
junior synonyms of Nodosochara Madler 1955, being based on the same
criteria: presence of an apical rosette in the center of the apex with
periapical narrowing of the cells, tapered base, thick basal plate.
Nodosochara heilongjiangensis nov. comb. = SongUaochara heilongjian-
gensis Z. Wang, Lu and Zhao 1985, Ed. Sci. TechnoI. Heilong-Jiang, p. 65,
PI. 28, Figs. 2-8.
Nodosochara speciaUs nov. comb. = Turbochara speciaUs Z. Wang 1978,
Mem. Nanjing Inst. GeoI. PaleontoI., p. 78, PI. 6, Figs. 1-9.
- Nitelloideae
Tolypella (A. Braun 1857) Leonhardi, pro parte = Sphaerochara Madler
1952 emend. Horn afRantzien and Grambast 1962.
Daily (1969) and Soulie-Marsche (1979), taking into account the characters
of the gyrogonites, revealed the identity of the living Tolypella section
Rothia Wood and the fossil Sphaerochara. This section includes a single
living species, T. intricata.
Sphaerochara intricata n. comb. = Tolypella intricata Trentepohl ex Roth
1797, Cat. Bot., p. 125; Leonhardi, emend. Wood 1965, p. 735.
2. Classification of the Charophyta

Order Sycidiales Madler 1952: gyrogonite composed of vertical cells.
Chovanellaceae Grambast 1962: vertical cells not subdivided.
Sycidiaceae Karpinsky 1906: numerous vertical cells, subdivided.
Pinnoputamenaceae Z. Wang and Lu 1980: cells forming two-branched
opposite systems.
Order Trochiliscales Madler 1952: gyrogonite composed of dextral spiral cells.
Trochiliscaceae Karpinsky 1906 emend. Z. Wang and Lu 1980: numerous
(often 18) dextral spiral cells.
Moellerinaceae nom. nov. not very numerous (5-12) dextral spiral cells.
Order Charales Lindley 1836: gyrogonite composed of sinistral spiral cells.
Palaeocharineae nov. subordo more than five sinistral spiral cells.
Eocharaceae Grambast 1959: numerous (more than six) spiral cells.
Palaeocharaceae Pia 1927: six spiral cells.
Charineae nov. subordo five sinistral spiral cells.
Porocharaceae Grambast 1962: apical pore always open.
Clavatoraceae Pia 1927: gyrogonite surrounded by a utricle.
Raskyellaceae Grambast 1957: apical pore closed by an operculum offive
202 M. Feist and N.Grambast-Fessard
cells leaving a rosette opening after being shed.

Characeae Agardh 1824: oogonium and gyrogonite with spiral cells joined
at the summit.
Acknowledgements. We are grateful to the Manager, Biostratigraphy Research Group, British

Geological Survey, Nottingham, for giving permission to figure specimens from Broadoak and
Warlingham Boreholes, and to C.l. Wood for improving the English text.
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fructifications. Stockholm Contrib GeoI4:45-197, 21 pi
Ishchenko TA, Ishchenko AA (1982) Novaia nakhodka kharophitov v verkhniem silure Podolii. In:
Sistematika i evolutsia drevnik rastenii Ukraini. Nauk Dumka, Kiev, pp 21-32, pi 5-6
Krassavina LK (1971) Sravnitelnoie izutchenie sovremenikh i iskopaemikh kharophitov:
plodonoshienia Nitellopsis obtusa i guirogoniti vidov Tectochara. Bot J 56, I: 106-117,2 pi
Liu J (1982) Jurassic Charophytes from Sichuan province. Bull Inst Geol Chinese Acad Geol Sci
(5):97-110 (in Chinese)
Sawa T, Frame PW (1974) Comparative anatomy of Charophyta. I. Oogonia and oospores of

Tolypella, with special reference to the sterile oogonial cell. BulI Torrey Bot Club 10 1,3: 136-144
Soulie-Marsche I (1973) Donnees generales sur les Charophytes du Plio-Pleistocene. BulI Assoc
Franc;. Etude Quaternaire (1973) 2:69-77
Soulie-Marsche I (1989) Etude comparee de gyrogonites de charophytes actuelles et fossiles et
phylogenie des genres actuels. Edit Revisee These Univ Montpellier 1979. Imprimerie des
Tilleuls, Millau, Fr, 237 p, 45 pI
Tappan H (1980) The paleobiology of plant protists, vol 1. Freeman, San Francisco, 1028 pp
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reference to classification and gyrogonite orientation of Trochiliscales and Sycidiales. Acta
Paleontol Sin 19,3:190-200,2 pI (in Chinese)
Wood RD, Imahori K (1965) A revision of the Characeae. Edit. Cramer. Weinheim. vol 1:904
pp, vol 2:395 pI
Chapter 12
Calcification of the Charophyte Oosporangium
A.R. LEITCH!
Abstract
The oosporangium is a unique and complex structure consisting of a central reproductive cell (the
oospore) surrounded by vegetative cells. In the subfamily Chareae, six vegetative ensheathing cells
(i.e. five spiral cells and one basal cell), completely surround the oospore and are in intimate contact
with it. After fertilization, a thick multilayered wall, the compound oosporangial wall (COW) forms
around the oospore. This wall is derived by the simultaneous deposition of three layers by the
ensheathing cells and three layers by the oospore. A calcified layer is deposited onto the COW by
the ensheathing cells. Calcification occurs outside the plasmalemma of each ensheathing cell but
within the confines of the cell wail; this is 'extracytoplasmic' calcification. The ensheathing cells
secrete an organic matrix that nucleates calcite development and is presumably involved in crystal
shaping. The calcite crystals of Chara are tabular and arranged in stacks, forming columns (like
gastropod shell). Columns of calcite are found in Lamprothamnium calcine, although no sub-
structure to these columns has been resolved. The differences seen in the calcified layer of Chara and
Lamprothamnium using light microscopy are largely a matter of perspective as to which is being seen,
the columns of calcite or the organic matrix.
1 Introduction
The family Characeae is a group of macrophytic green algae, anchored to

organic-rich sediments by colourless rhizoids. They occur in lakes and the
backwaters of streams and rivers, often forming high-density 'charophyte
meadows'. They favour oligotrophic calcareous water to a depth of 15 m (Stross
1979; Spence 1982) disappearing from lakes when they become eutrophic. A few
species are found in brackish water. One genus, Lamprothamnium Groves, is
exceptional in that it can survive highly saline conditions, although it requires
brackish water to complete its life cycle (DeDeckker and Geddles 1980).
The taxonomic affinities of the family Characeae remain controversial.
Their supposed intermediate position between green algae and bryophytes has
led workers to propose that they represent a subdivision of higher plants
(Bremer and Wantorp 1981). However, this is not widely accepted. Mattox and
Stewart (1984) considered that the Characeae belonged to a class of green algae
(Division Chlorophyta) while Round (1984) elevated the group to divisional
status (the Charophyta). Within the single extant family Characeae there are six
extant genera belonging to two subfamilies, the Nitelleae comprising Nitella
Ag., and Tolypella Braun, and the Chareae comprising Chara L., Nitellopsis Hy,
Lychnothamnus (Rupr.)Leonh .. and Lamprothamnium. The genus Tolypella
'J.I. Centre for Plant Science Research. Colney Lane. Norwich. NR4 7UJ. England
Calcification of the Charophyte Oosporangium 205
has two sections, acutifolia and obtusifolia, with such different oosporangial
morphologies (see Sawa and Frame 1974; and Daily 1975) that the validity of
the genus should be questioned.
The family Characeae is unique amongst the algae in that the reproductive
cell (termed the oosphere before fertilization and oospore after fertilization) is
completely ensheathed by vegetative cells. This forms an integral structure
called the oosporangium (or oogonium). The complex development and
morphology of the oosporangium is well documented (Sundaralingam 1954,
1962). Briefly, the oosphere/oospore is ensheathed by five elongate, sinistrally
spiralling cells, called spiral cells, and one (in Chareae and Tolypella section
acutifolia) or three (in Nitella and Tolypella section obtusifolia) small cells,
called sterile cells. In this work these cells are collectively called the ensheathing
cells. Subtending each spiral cell is either one cell (Chareae) or two cells
(Nitelleae) which together form a crown offive or ten cells, called coronula cells,
at the apex of the oosporangium. In addition, the oosporangium has two small
cells at its base that anchor the structure to the algal thallus. The ensheathing
cells and the oospore are of interest here because these cells are involved in
post-fertilization development.
Before fertilization the wall of the oosphere is in intimate contact with the
walls of the ensheathing cells. Both walls are microfibrillar and are considered
to be primary cell walls (Leitch 1986, 1989). After fertilization, a thick mul-
tilayered wall, called the compound oosporangial wall (formerly known as the
oospore membrane), is deposited around the oospore. The compound oospor-
angial wall (COW) is derived partly from the oospore and partly from the
ensheathing cells by the deposition of secondary wall layers. The oospore
deposits three secondary wall layers against the inside of its primary wall.
Simultaneously the ensheathing cells deposit three secondary wall layers against
the inside of their primary walls. The resulting COW is an integral unit that
surrounds the oospore and is built up of eight layers (Leitch 1986, 1989).
In the subfamily Chareae and Tolypella (section acutifolia), after the
deposition of the COW, a calcified 'shell-like' layer called calcine is deposited
(Horn af Rantzien 1956; Daily 1975). The calcine has a mineral component of
low magnesium calcite in freshwater species or high magnesium calcite in
brackish water species (Daily 1975; DeDeckker and Geddles 1980; Leitch
1986). The calcine is secreted by the ensheathing cells directly onto the COW.
Horn af Rantzien (1956) reported that the first calcine deposited (i.e. closest to
the COW) was distinctive from later formed calcine, calling the two types endo-
and ecto-calcine respectively. The calcine of Chara and Lamprothamnium are of
particular interest because Soulie-Marsche (1979) and Feist and Grambast-
Fessard (1984) reported that these genera typify two distinct forms of
calcification. In this paper techniques in light microscopy (LM), scanning
electron microscopy (SEM), and transmission electron microscopy (TEM) are
used to examine the differences between these calcine forms. Calcine from
Lamprothamnium papulosum Groves, Chara hispida L. and Chara delicatula
Agardh. is examined.
206 A.R. Leitch
2 Materials and Methods
Plant material of Chara delicatula and Chara hispida was collected from
drainage ditches in the Gordano Valley, Avon (ST 444 735) where they were
growing in clear, calcareous water to a depth of I m. Lamprothamnium
papulosum was collected from a small lake near a salt marsh at Lymington,
Hampshire (SZ 328 939) where the plants were growing in turbid, brackish
water to a depth of 1.5 m.
The oosporangia were fixed for TEM in 2.5% glutaraldehyde, 0.1 M
formaldehyde in 0.2 M sodium cacodylate buffer (pH 7.6) for 3 h at room
temperature. They were then post-fixed in 2% osmium tetroxide (in sodium
cacodylate buffer) for 3 h at room temperature and block stained overnight in
0.25% uranyl acetate. The oosporangia were embedded in LR white (medium)
resin and polymerized at 70°C overnight. Sections were cut with a diamond
knife, stained with lead citrate and uranyl acetate and examined at 80 kV in a
Philips 300 transmission electron microscope.
The oosporangia used for SEM were freeze-dried prior to examination. The
calcine, was isolated from the oosporangium using fine entomological needles
and air dried. All specimens were coated with gold using a Polaron Sputter
coater and examined at 15 kV in a Philips 501B scanning electron microscope.
The oosporangia for light microscopy were fixed and embedded as forTEM
and then ground to thin sections using carborundum powders. They were then
examined using an Olympus BHB compound microscope.
3 Development of the COW and the Calcine in Chara and Lamprothamnium
Following fertilization the COW develops around the oospore. The oospore
itself secretes three secondary wall layers in the following sequence: an amor-
phous layer; a helicoidal layer (i.e. a characteristically ordered microfibrillar
wall; see Neville (1986); and a random microfibrillar layer (Plate I.l). Simul-
Plate 1.1 A section through a COW. 1 the random microfibrillar layer; 2 the oospore helicoidal
layer; 3 the amorphous layer; 4 the oospore primary wall; 5 the spiral cell primary wall; 6 the
crystalline layer; 7 the pigmented layer. The ornamentation layer produces an irregular surface to
the COW (arrow). Chara hispida TEM x6700
Plate 1.2. Section through the lateral wall of a spiral rell of Chara hispida showing the calcine (c)
deposited on each side of the wall (w). Electron-der. ..: fragments in the calcine are calcite crystals.
TEM xiiOO
Plate 1.3. A weakly calcified oosporangium of Lamprothamnium papulosum showing concave spiral
profiles (arrows). Note that the apex is the most weakly calcified. SEM x50
Plate 1.4. A strongly calcified oosporangium of Lamprothamnium papulosum showing fiat spiral
profiles (arrows). Note that the apex is the most weakly calcified. SEM x50
Plate 1.5. Section through a calcifying spiral cell of Chara delicatula. An organic matrix (0) is seen
outside the plasmalemma (p). Note the electron-dense crystal nuclei (n). The holes (h) are sites where
crystals have been pulled out of the section. TEM x 7000
Plate 1.6. Section through the calcine of Chara hispida showing a complex organic matrix (0). Note
the organic layers in the endocalcine (arrowheads). TEM x3300
Plate 1.1-1.6
208 A.R. Leitch
taneously, the ensheathing cells deposit three secondary wall layers in the
following sequence: a crystalline layer, possibly of calcium sulphate (Leitch
1986), embedded in an electron-dense matrix; a highly pigmented helicoidal
layer; and an amorphous layer (Plate l.l; Leitch 1986).
After the development of the COW the ensheathing cells deposit the
calcine. Calcine is initially deposited against the COW and the inner half to
two-thirds of the lateral walls of the ensheathing cells (Fig. I). Further
calcification increases the thickness of the calcine (Fig. I). Because calcine is
deposited within the confines of the walls of the ensheathing cells, the form of
the calcine reflects the cell shape and so occurs as five spiralling units and a small
rounded/pentagonal unit of calcine. Each unit of calcine lies against the COW
and is separated by a cell wall (Plate I.2). The spiralling units are called spirals
and the basal unit is called the basal plate. With the death of the ensheathing
cells the calcification stops. The outer wall of the ensheathing cells then decays
:...... .
Fig. I. The development of calcine in Chara. a-c Progressive degrees of ca lcification. Calcine
(stippled) is laid against the COW (solid black) and the lateral walls of the ensheathing cells. The right
column of diagrams illustra tes the profile of calcine in longitudinal sections through varyingly
calcified oosporangia
away to reveal the underlying calcine. When this has happened a weakly
calcified oosporangium shows spirals with a concave profile (Plate 1.3) and a
strongly calcified oosporangium shows spirals with a flat or convex profile (Plate
1.4). The degree of calcification depends on the species, the environmental
conditions, and the physiological condition of the alga. The spiral apices are
often more weakly calcified than the rest of the spiral. This is particularly
noticeable in Lamprothamnium (Plates 1.3-1.4). On oospore germination the
weakly calcified apex cracks and falls away by the growth of the emerging
germling.
Fixing, embedding, and sectioning calcifying oosporangia for TEM is
difficult because the thick walls and the calcine reduce infiltration ofthe fixative
and resin. However, it is clear that calcification occurs outside the ensheathing
cells' plasmalemma (Plate 1.5) and within the confines of the cell wall. The
process reduces the volume of the cytoplasm. This site of calcification cannot be
considered as intracellular calcification, as reported by Daily (1975), because
calcification is outside the plasmalemma, it is 'extracytoplasmic' but within the
cell wall.
Calcification occurs by the secretion of an organic matrix, probably in the
form of precursors, that polymerizes into sheets, outside the plasmalemma of the
ensheathing cells (Plate 1.5). Under TEM the organic matrix that is closest to
the plasmalemma shows electron-dense patches (Plate 1.5). These patches are
interpreted as calcite nucleation sites. The organic matrix that is furthest from
the plasmalemma (hence older) shows electron-opaque cavities (Plate 1.5).
These cavities are sites once occupied by calcite crystals, the futher from the
plasmalemma the larger the cavities and hence the larger the crystals. Fully
developed calcine has calcity crystals ramified by an organic matrix (Plate 1.6).
Unfortunately, the staining requirement (uranyl acetate and lead citrate) to
show the organic matrix decalcifies the calcine. However, omitting uranyl
acetate reveals electron-dense calcite crystal fragments (Plate 1.7).
In a strongly calcified oosporangium of Chara the calcine that is closest to
the COW (hence older) is of a different form from younger, later formed, calcine
(Plate 1.6). First-formed calcine has a much higher organic matrix to calcite ratio
than the later-formed calcine and the matrix appears as thick, tightly packed
bands. First-formed calcine represents Horn af Rantzien's (1956) endocalcine
and accounts for less than 10% of the calcine in strongly calcified oosporangia.
However, very weakly calcified oosporangia show only calcine that is typical of
endocalcine (Plate 1.8). Bands of organic matrix are found in younger calcine
(ectocalcine) but they are more diffuse and are seen in transverse section as
concave bands arising from the lateral wall (Plate 1.9). Bands of organic matrix
are found in Lamprothamnium but they are much thinner and a distinct
endocalcine cannot be resolved.
A transverse fracture of a spiral from Chara shows in SEM a demarcation
line between the endocalcine and the ectocalcine in some, but not all, specimens
(Plate 1.10). A transverse fracture of the ectocalcine shows columns that are
long, thin, closely packed and radiate diagonally from the lateral walls of the
spirals and vertically from their inner wall (Plate 1.11). These columns may be
210 A.R. Leitch
Plate 1.7-1.12
simple or may form fans. Vertically aligned columns appear to blend into
diagonal columns. Lamprothamnium calcine (Plate 1.12) differs from Chara
calcine (Plate 1.11) in that the columns are more defined, they have a more
uniform direction, and vertical columns from the base form a distinct suture
with diagonal columns from the lateral wall (Plate 1.13).
In Chara at high resolution the columns are seen to be composed of stacked
layers of tabular crystals (Plate 1.14). This is similar to the crystal layers seen in
gastropod shell (Wilbur and Saleuddin 1983). The youngest calcine, which is
closest to the plasmalemma in the calcifying oosporangium, shows stacks of
crystals above each column (Plate 1.15). Greater resolution of each stack shows
layers of crystals that diminish in size towards the top (Plate 1.16). At the apex
of each stack is the smallest crystal and it represents a crystal growing after its
nucleation. Each crystal in the stack grows laterally forming tabular crystals.
The growth ofthe crystal is probably influenced by the organic matrix, as occurs
in molluscs (Degens 1976). With the lateral growth of the cyrstals the stacks fuse
together to form the columns. The layers of tabular crystals comprising the
columns were not resolvable in Lamprothamnium.
Soulie-Marsche (1979) and Feist and Grambast-Fessard (1984) noted that
the calcine from Chara and Lamprothamnium, when ground into thin-section
using carborundum powders, and examined under light microscopy appeared
different. They called the Lamprothamnium calcine the 'Y' form of calcification
because of an inverted 'Y' visible in transverse section (Plate 1.17). Likewise,
a concave substructure to Chara calcine gave rise to the term 'U' form of
calcification (Plate 1.18). An examination of the inverted 'Y' of Lamprotham-
nium calcine (Plate 1.17) shows a strong resemblance to the fractured face of
calcine examined under SEM (Plate 1.12). It is concluded therefore, that the 'Y'
form is generated by the suture between the change in column direction. Further,
the columns can be seen under light microscopy as radiating lines. An
examination of the calcine of Chara under light microscopy (Plate 1.18), and
comparing this with calcine examined under TEM (Plate 1.9), shows clearly that
the concave bands are bands of organic matrix found within the calcine. A close
...
Plate 1.7. Section through the calcine of Chara hispida showing the organic layers in the endocalcine
(0). Note the array of crystal fragments in the direction of the arrow. TEM x2700
Plate I.S. Section through a weakly calcified oosporangium of Chara hispida. Note the endocalcine
(e) on the COW. TEM x2000
Plate 1.9. Section through two spirals of Chara hispida showing concave bands of organic matrix
(arrowheads) in the calcine. The central area of calcine (c) sections least well. Note the lateral wall
(I). TEM x830
Plate 1.10. Fractured spiral of Chara hispida showing the endocalcine (e) and the calcine (c). Note
that in this specimen there is a distinct demarcation between calcine types (arrowhead). SEM x830
Plate 1.11. Fractured spiral of Chara hispida showing the columns (c) and the endocalcine (e) at the
fracture surface. SEM x450
Plate 1.12. Fractured spiral of Lamprothamnium papulosum showing columns (c) at the fracture
surface. SEM x620
212 A.R. Leitch
Plate 1.13-1.18
examination of Chara and Lamprothamnium calcine under light microscopy

also reveals, although faintly, columns in Chara and organic bands in Lam-
prothamnium. Therefore, the differences seen under light microscopy are largely
a matter of perspective concerning which of these is most readily visible, the
columns or the organic matrix (Fig. 2).
4 Discussion of Calcification
The study of calcification in lower plants and animals has led to a number of
generalizations. The generalizations concerning the mechanism ofbiomineral
precipitation involve a 'four compartment model' as proposed by Simkiss
(1986). The four categories of the model can be defined as follows: (1) the
modification of ion activity, (2) a diffusion limited site, (3) nucleating surfaces,
and (4) lattice modifiers and crystal inhibitors (Simkiss 1986). These categories
will be discussed with reference to the charophyte oosporangium.
As far as the author is aware no metabolic or ion flux experiments have ever
been carried out on charophyte oosporangia and so nothing is known about ion
activity (the first category ofSimkiss' 1986 model). However, it is known that in
other algae the active pumping of ions including calcium is directly involved in
biomineralization (see Borowitzka 1982 and references therein).
In the oosporangium calcification occurs 'extracytoplasmically', against the
COW and within the confines of the walls of the ensheathing cells. This site of
calcification can be considered as an example of Simkiss' (1986) diffusion
limited site. The isolation of a compartment bound by the cell walls and the
plasmalemma will influence the movement of ions. This is important because
slight modifications in the concentration of ions within the calcification site will
have an effect on the biomineral precipitated. There is a possibility offour major
calcium exchange pools in a system like this: the walls of the ensheathing cells,
the cytoplasm of the ensheathing cells, the calcite crystals, and the plant's
environment. Which pools have the most significant influence on the calcium
supply necessary for calcification remains unstudied .
..
Plate 1.13. Fractured spiral of Lamprothamnium papulosum showing the suture (arrowheads) at the
change in direction of the calcine columns. SEM x II 300
Plate 1.14. Fractured spiral of Chara hispida showing that the calcine columns are layers of tabular
crystals (arrows). SEM x6500
Plate 1.15. The spiral surface (last formed calcine) shows small crystal stacks (s). Note the fractured
face (j) of the spiral. Chara hispida, SEM x3loo
Plate 1.16. Detail of the last-formed calcine showing a single stack comprised of tabular crystals
(arrowheads). Chara hispida, SEM x22 000
Plate 1.17. Carborundum ground thin-section of Lamprothamnium papulosum spirals showing
radiating columns (large arrowheads) and concave bands (small arrowheads). LM x300
Plate I.IS. Carborundum-ground thin-section of Chara hispida spirals showing concave bands
(large arrowheads) and indistrict radiating columns (small arrowheads). LM xl90
214 A.R. Leitch
Fig. 2a,b. Diagram showing the relationship between the organic matrix (dashed lines) and the
calcite columns (complete lines) in the calcine (stippled) of Chara (a) and Lamprolhamnium (b)
In order for calcite nucleation to occur it is required that the solubility

product of calcium carbonate is exceeded. This is a rate limiting step unless the
crystal is nucleated by an organic surface (third category of Simkiss' 1986
model). It is likely that the sheets of organic molecules seen outside the
plasmalemma of calcifying enshea thing cells are providing the nuclea tion sites
necessary for crystal growth ; asimilar process occurs in molluscs (Degens 1976).
The calcite crystals in Chara calcine are tabular and do not appear like in
vitro-grown crystals. It has been suggested that organic molecules are involved
in crystal shaping by inhibiting and stimulating crystal growth in specific
directions (fourth category of Simkiss' 1986 model). Organic molecules are
known to be crystal inhibitors (Simkiss 1964). Wilbur and Bernhardt (1982)
suggest that carboxyl groups on proteins are responsible for this, the protein
interacting with the normal growth site of the crystal, preventing additions to the
crystal lattice. This inhibits normal crystal development or stops growth al-
together (Degens 1976). It is quite likely, therefore, that the complex three-
dimensional network of organic material in calcine is involved in controlling
crystal growth and shape. The crystal shape is similar to that found in the
gastropod shell where shape is modified by the organic matrix (Wilbur and
Saleuddin 1983).
The calcine of Chara and Lamprothamnium is similar in a number of
respects. The calcine of both genera has a calcite mineral component (although
in Lamprothamnium there is a higher magnesium content; Leitch 1986), a
complex ramifying organic matrix, and a columnar substructure which is
particularly visible under SEM. However, in Lamprothamnium the organic
matrix is less prominent and has a weaker banding pattern, the columnar
substructure is more uniform with a characteristic suture at the change in
column direction, and the columns reveal none ofthe substructure (i.e. layers of
tabular crystals) seen in Chara calcine. The question as to whether the
differences in the calcine of Chara and Lamprothamnium are of ecological or
taxonomic importance cannot be fully answered until calcine from a brackish
water species of Chara is examined. However, it is most likely that the organic
matrix influences the calcine form and that the matrix is under the control ofthe
alga. This implies a taxonomic importance to calcine differences.
It should be emphasized that, in interpreting fossil calcine substructure,
great care should be taken because selective removal of the organic component
or the mineral component of calcine during fossilization is likely to influence the
calcine form.
Acknowledgements. I thank NERC for supporting this work.
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Chapter 13
Calcareous Nannofossils
S.D. HOUGHTON}
Abstract
Calcareous nannofossils (mainly coccoliths) have been important constituents of pelagic carbonates
since the Early Jurassic, accounting for over 50% of some Tertiary sediments and around 30% of
Recent oozes. Coccolith biosynthesis by calcareous nannoplankton forms an important role in
marine biogeochemical cycles, contributing to the fluxes of carbon and calcium from the hydro-
sphere to the lithosphere. Most coccoliths are thought to descend through the water column 'sealed'
within the faecal pellets of zoo plankton grazers. This mode of sedimentation ensures that a constant
supply of nannofossil carbonate, largely free from dissolution effects, reaches the ocean floor.
Although post-depositional dissolution and diagenesis may modify coccolith biocoenoses, patterns
of accumulation of coccoliths in sediments do reflect the oceanographic regimes of the overlying
surface waters. Calcareous nannofossils are useful as tools for studying palaeoclimatic change and
also form very good biostratigraphic markers.
1 Introduction
Coccoliths are the micron-sized (ca. 1-25 /Lm) calcitic plates secreted by uni-
cellular haptophyte marine algae, the coccolithophorids. In the living cell,
coccoliths are extruded to .form a cover termed collectively the coccosphere,
which lies outside the cell membrane (Fig. 1). Between 10 and 100 coccoliths
comprise the coccosphere, which may range in size from 5 to 50 /Lm. The term
nannoplankton, as defined by Lohmann (1909) for the very small forms, 60/Lm
or less, that pass through phytoplankton nets, therefore includes both the
coccolithophorids and their skeletal elements. Individual dispersed coccoliths
are usually all that is observed in fossil assemblages because of post-mortem
disintegration of the coccosphere. However, intact or partially intact cocco-
spheres of the more robust forms may occasionally be found (Figs. 2, 3a). Fossil
coccoliths (Fig. 3), together with other calcite bodies of unknown origin which
occur in the same size fraction, constitute the heterogeneous group, known as
calcareous nannofossils. These other forms, termed nannoliths by some
workers, display a wide range of elaborate structures and often form a sub-
stantial component of the nannofossil assemblages. The most notable of these
forms are Discoaster species.
1Department of Earth Sciences, The Open University, Milton Keynes, Buckinghamshire,

MK76AA, UK
Present address: Adams Environmental The Business Centre 103-109, Lavender Hill, London
SWI15QF, UK
218 S.D. Houghton
- c
Fig. 1. Generalized diagram of a motile coccolithophorid

cell showing exterior at left, and section at right (after Tap-
pan 1980). c Coccolith ; ftwo flagella ; h haptonema; p plas-
n tids (chloroplasts); n nucleus; s storage body of leu cos in
Fig. 2. Cross-sectional view of a coccosphere of Reticulofenestra pseudoumbilica, showing inter-

locking coccolith construction (xlO 700) ; sample from Site 223 DSDP, Arabian Sea (Middle
Miocene age)
Calcareous Nannofossils 219
Fig. 3a-f. SEM micrographs of some common Late Cenozoic calcareous nannofossils. a Cocco-
sphere of Coccolith us pelagicus (x2600). Small coccolith at centre of coccosphere Gephyrocapsa
ericson ii, other coccolith at right side of coccosphere Calcidiscus leptoporus. b Gephyrocapsa
caribbeanica, distal view (xiS (00); C Braarudosphaera bigelowii, proximal view (xiO 500); d
Calcidiscus leptoporus, distal view showing slight etching of individual crystallines (x9100) ; e
Rhabdosphaera c/avigera , side view (x9300); f Syracosphaera pulchra, proximal view (x 14800)
220 S.D. Houghton
Species of calcareous nannofossils are defined on morphological charac-

teristics of fragments or components (coccoliths) of the original organism. Some
species are therefore probably no more than artificial categories. Studies on
Recent coccolithophorids have shown that some individuals bear two mor-
phologically distinct coccoliths on the same coccosphere (dimorphism). Ad-
ditionally, some coccolithophorids display dithecatism: in which the cocco-
sphere consists of a double layer of coccoliths, an inner layer (endotheca)
composed of caneoliths and an outer layer (exotheca) composed of cyrtoliths.
Further taxonomic problems are caused by coccolithophorid individuals
secreting morphologically distinct coccoliths during successive phases of their
life cycle. The dual plant and animal nutritional behaviour of some coc-
colithophorids (see ecology, below) has led to the group being claimed by both
botanists and zoologists. However, most micropalaeontogists have used the Code
of Biological Nomenclature for the systematic description of the taxa. There is
no general agreement on the classification of Mesozoic and Cenozoic calcareous
nannofossils. Many authors prefer to arrange nannofossil genera in alphabetical
order for their systematic descriptions. Tappan (1980) included coccolitho-
phorids in Class Coccolithophyceae and Division Haptophyta of the Plant
Kingdom, however, more recent studies (e.g. Westbroek et al. 1986) include
coccolithophorids in the phylum Prymnesiophyta (class Prymnesiophyceae).
This chapter reviews:
1. The main biological and ecological aspects of Recent coccolithophorids;
2. The main calcareous nannofossil evolutionary trends;
3. Calcareous nannofossil biostratigraphy;
4. Dissolution and diagenetic effects on nannofossil assemblages;
5. Recent advances in the use of nannofossils as tools in the elucidation of
palaeoclimatic; and palaeo-oceanographic change.
2 Biology of Recent Coccolithophorids
Most haptophytes (including coccolithophorids) are characterized by a hap-

tonema which lies between two smooth, whip-like flagella (Parke and Adams
1960). Internally, the coccolithophorid cell contains a single sub-spherical
nucleus and two golden-brown chloroplasts. The photosynthetic pigments in
the cell are chlorophyll-a and carotenoids (Sherwood 1974). The cell is enclosed
by a membrane, which in turn has an outer mucilaginous covering. External to
the cell membrane are the coccoliths. The life cycles of coccolithophorids are
complex, and for many species they are still poorly understood. Detailed
laboratory studies have been limited to a few species, mainly because of the
practical difficulties in obtaining and maintaining monospecific cultures.
Biomineralization aspects of coccolith formation have been most extensively
studied on the species Emiliania huxleyi (Lohmann) Hay and Mohler and
Pleurochrysis carteri (Westbroek et al. 1986; de-Vrind et al. 1986). These two
species may be cultured easily on synthetic seawater or semi-synthetic media in
large batches, and coccolith formation may to some extent be regulated by

chemical and other environmental parameters (Dorrigan and Wilbur 1973).
Coccoliths of E. huxleyi are formed in a specialized organelle, located near the
nucleus and consisting of a coccolith vesicle (which surrounds the growing
coccolith) and is joined to a system of anastomosing tubes, the reticular body.
Prior to calcification, the specialized organelle develops a high concentration of
Ca 2 + -binding, water-soluble polysaccharides relative to the other cell constit-
uents (van der Wal et al. 1983). Initial calcification occurs on the rim of a base
plate within the coccolith vesicle, and as calcification proceeds, complete
coccoliths are extruded to the periphery of the cell where they interlock to form
the coccosphere. Pure coccoliths can be isolated by ultrasonic disintegration of
the cell and centrifuging the resultant mixture in a silica or sucrose gradient (de
long 1975). Laboratory studies have shown that three different cell types of E.
huxleyi are commonly recorded:
1. Coccolith-bearing cells (C-cells);
2. Naked cells which lack conspicuous extracellular scales or coccoliths
(N-cells);
3. Flagellated cells that have organic scales on their cell surface but no calcified
coccoliths (S-cells).
However, it is not known what processes trigger the three cell types. Coc-
colithophorid division occurs inside the cell and the shell is either abandoned or
shared equally by the two daughter cells, with subsequent reconstruction of a
complete cell. When environmental conditions are favourable, cell division is
rapid, and in the logarithmic growth phase the cells may divide every 24 h.
Many haptophytes show an alternation of two distinct stages in their life
history (a dimorphic life cycle;· Tappan 1980). The motile cell alternates during
the life history with a non-motile stage which may be a pelagic cyst. Most
coccolithophores are thought to be free-swimmers during one stage of their life
cycle, using the pair of flagella for propulsion. Coccolithophorids, however, are
probably passive plankton, as laboratory studies (Leadbeater 1971) have shown
that they move on average 5 to 18 mm/min (at 18°C) which is certainly less than
the rate of most current movements. Studies by Black (1972) have shown that
coccolithophorids have a flexible skeleton during the free-swimming stage and
a rigid skeleton resulting from membrane calcification during the encysted
phase of the life cycle. Planktonic and benthic stages, both with and without
coccoliths, have been observed in several species, and although their
significance has yet to be fully understood, the benthic stages are thought to
occur in hostile environments. The ability of the coccolithophorids to secrete
coccoliths on their organic scales differentiates them from all other algae.
3 Coccolith Morphology and Crystal Structure
X-ray diffraction patterns and crystal forms have indicated that in nature
coccoliths are composed of low-magnesium calcite. Most coccoliths are con-
222 S.D. Houghton
structed of rhombohedral calcite crystals in a radial arrangement, a few have

hexagonal prisms, and some have a combination of these two forms in a single
crystal (Tappan 1980). On the basis of crystallization two main types of coccolith
are distinguished in coccolithophorids: (I) holococcoliths, (2) heterococcoliths.
The basis of this subdivision is the varying degree of crystal growth.
Holococcoliths have unmodified crystal forms with retention of their geometric
faces and angles and are precipitated outside the cell membrane (Parke and
Adams 1960). Crystallolith holococcoliths consist of simple rhombohedral
crystals scattered over the surface of the organic scale, although they do not
completely cover the scale (e.g. motile phase of Coccolith us pelagicus).
Calyptroliths include holococcoliths constructed of hexagonal crystals in the
form of proximally open caps, e.g. Sphaerocalyptra papillifera (Halldal and
Markali) Halldal. Zygoliths holococcoliths include forms which have an el-
liptical ring with a slightly or strongly arched crossbar (e.g. Zygosphaera
divergens Halldal and Markali). Holococcoliths, because of their small crystal
size and their loosely packed construction, are susceptible to rapid post-mortem
disintegration and are rarely found in the fossil record.
Heterococcoliths are internally formed and elaborately constructed. The
crystal faces and angles are modified, and the form ofthe coccolith is completely
controlled by the cell. Heterococcoliths are made up of numerous crystals,
commonly with radial construction, although rarely achieving exact radial
symmetry. These are termed heliolithae. The most commonly occurring
heliolith coccoliths are placoliths, which comprise discs built of two elliptical or
circular shields constructed of radially arranged crystallites. The central area of
placoliths may comprise an open pore [e.g. Umbilicosphaera sibogae (Weber-
van Bosse) Gaarder], which may be crossed by a bridge (e.g. Gephyrocapsa
caribbeanica Boudreaux and Hay, Fig. 3b), or may be filled with a lattice [e.g.
Reticulofenestra pseudoumbilica (Gartner) Gartner, Fig. 2]. Caneoliths are
basket-like coccoliths with a central area or bottom, of lamellae and petaloid
upper and lower rims (e.g. Syracosphaera pulchra Lohmann, Fig. 3f). Cyrtoliths
are basket coccoliths with a central process, which may be very elongated and
club-shaped, e.g. Rhabdosphaera clavigera Murray and Blackman (Fig. 3e), or
may be distally flared and trumpet-shaped as in Discosphaera tubifera (Murray
and Blackman) Ostenfeld. Helicoliths include coccoliths which have a marginal
flange, which increases in height around the periphery of the oval disc, over-
lapping the early part of the flange in a spiral construction, e.g. Helicosphaera
carteri (Wallich) Kamptner. Unusual, elongated, diamond-shaped coccoliths
with a central area of parallel laths and a raised rim are termed scapholiths and
are represented by living A noplosolenia and Calciosolenia.
Some calcareous nannofossils, constructed of a single or very few crystals,
may have radial symmetry. These forms are termed the Ortholithae and
include Discoaster species, ceratoliths and pentaliths. Discoaster species are
discussed under 'other nannofossil groups' (see below). Ceratoliths are large
horseshoe-shaped crystals. Each ceratolith acts as a single crystal of calcite, e.g.
Cera to lith us cristatus Kamptner (Fig. 4). Pentaliths are nannofossils formed of
five pieces, each consisting of a single crystal orientated 72 apart, e.g. Braa-
0
Fig.4. Optical photomicrograph

(using crossed polars) of Recent calca·
reous nannofossils from the Celtic Sea
(x 1600). Notice horseshoe-shaped Ce-
ratolithus cristatus (centre) which acts
as a single crystal as opposed to the
extinction patterns produced by the
surrounding coccoliths
rudosphaera bigelowii (Gran and Braarud) Deflandre (Fig. 3c). For a com-
prehensive review of coccolith morphology, see Tappan (1980).
Calcite crystals in coccoliths are arranged in such a way that a negative,
uniaxal, pseudofigure is seen under crossed polars. This results in each in-
dividual coccolith showing a well-defined extinction cross (Fig. 4). Kamptner
(1954) observed that each type of coccolith has a diagnostic interference pattern,
and that the extinction position of the calcite crystals is important for the
recognition of the species. The lines of the interference figure may curve either
to the left (laevogyre) or to the right (dextrogyre), as they are traced out from the
centre of the coccolith in distal view.
4 Other Calcareous Nannofossil Groups
Although all fossil coccoliths are generally thought to have been secreted by
coccolithophorids, other nannofossil groups have different and more obscure
affinities. These groups will now be discussed briefly.
Discoaster species (Fig. 4a) comprise nannofossils of star- or rosette-sha ped
construction. Most forms are flat-lying and remain dark when viewed with
crossed nicols (using optical microscopy) because the c-axis of the calcite is
vertical. The group is thought to contain numerous genera, most of which are
usually included in Discoaster. Tan Sin Hok (1927) attempted to separate the
Discoaster group into the genera H emidiscoaster, forms with fusion of alternate
rays, and Eudiscoaster, forms in which all the rays occur as independent crys-
tals. This subdivision is considered inapplicable because some otherwise
homogeneous species have variants belonging to both the suggested genera
(Bramlette and Reidel 1954). The secretion offused crystal units may also have
palaeoecological significance (see Sect. 6). Theodoris (1983) used Eu-discoaster
and Helio-discoaster as a replacement nomenclature and suggested that Dis-
coaster is an illegitimate name. However, most workers still retain the name
Discoaster for the whole group and this terminology is followed here.
The microrhabulids include rod-like cylindrical to pyriform calcareous
nannofossils of unknown origin which first appeared in the Late Jurassic
224 S.D. Houghton
although they were most abundant in the Late Cretaceous. The nannoconids
include forms with a thick wall of wedge-like calcite plates orientated per-
pendicular to and spirally surrounding an axial cavity or canal. Nannoconus
(Fig. 5c) appeared in the latestlurassicand is particularly common in the Lower
Cretaceous of the Tethyan realm, where they occur in fine-grained limestones
together with coccoliths, tintinnids, ammonites and planktonic foraminifera.
Nannoconus had a cosmopolitan distribution as it has been recorded from
Spitzbergen and also from the Falkland Plateau.
The schizosphaerellids include forms with subspherical tests (5-30 /Lm in
diameter) constructed of two overlapping or interlocking dissimilar valves (Fig.
8b). The valves are constructed of small tabular crystals regularly arranged in a
cubic lattice. The bipartite construction is unlike anything recorded in the
coccolithophorids, but many represent an encysted or non-motile phase, similar
to the braarudospheres (Tappan 1980). Schizosphaerella occur abundantly in
Jurassic marine sediments.
Fig. 5. a Discoaster nobilis (range Palaeocene) (x6000); b Thoracosphaera granifera (range

Pliocene-Recent) (x6500) ; c Nannoconus sp. (x8600); d Coccosphere of Ellipsagelospheara cf.
keftalrempti (x6400)
The Thoracosphaera (Fig. 5b) are spherical shells composed of a mosiac of

interlocking crystal units. The crystals are constructed of polygonal prisms that
may be solid or axially perforate. The genus Thoracosphaera, now classified as
a dinophyte and not a coccolithophorid (Tangen et al. 1982), has been tradi-
tionally included in nannofossil studies. Living Thoracosphaera are found in the
open ocean under normal environmental conditions, however, the Thoracos-
phaera blooms which followed the Cretaceous/Tertiary boundary may reflect
an adaption to special environmental conditions because of the absence or
scarcity of competing calcareous plankton.
5 Ecology of Recent Coccolithophorids
Recent coccolithophorids, being predominantly autotrophic plankton, are

largely restricted to the photic zone of the water column (0-200 m depth) where
sunlight can easily penetrate. Their maximum concentrations occur above a
depth of 100 m in the tropics and above 50 m in temperate regions; they
therefore live above the thermocline in the world's oceans. The vertical dis-
tribution of some of the more common coccolithophorid species in the oceans
is summarized in Table l. Upper photic zone species (0-50 m) are typically of
low density and include cyrtoliths which are characterized by elongated or
distally flared appendices (e.g. Discosphaera tubi[era and Rhabdosphaera
clavigera). A smooth, thin-walled form of Umbellosphaera (u. irregularis
Paasche) is also dominant in tropical surface waters. Mid-photic zone species are
typically placoliths with an open central pore (e.g. Umbilicosphaera spp.,
Table 1. Vertical zonation ofRecentcoccolithophorid species in the photic zone of

the Pacific and Atlantic oceans. Compiled mainly from Okada and Honjo (1973)
and Okada and Mcintyre (1977)
Species Vertical distribution in the photic zone
Emiliania huxleyi Throughout (0-200 m)

Gephyrocapsa oceanica
Discosphaera tubi[era
Rhabdosphaera clavigera Upper Photic Zone (0-50 m)
Umbellosphaera irregularis
Calcidiscus leptoporus
Umbilicosphaera hulburtiana
Umbilicosphaera sibogae Middle Photic Zone (ca. 50-150 m)
Oolithotus [ragilis cavum
Umbellosphaera tenuis
Anthosphaera quadricornu
Florisphaera profunda
Thorosphaera /fabellata Lower Photic Zone (150-200 m)
Hayaster perplex us
Oolithotus [ragilis
226 S.D. Houghton
Oolithotus fragilis cavum Okada and McIntyre) and U. tenuis (Kamptner)

Paasche, a more heavily constructed form of Umbellosphaera with ribs on the
distal shield. Lower photic zone forms include large coccolithophorid cells,
characterized by non-perforate coccoliths (e.g. Oolithotus fragilis (Lohmann)
Okada and McIntyre and Hayaster perplexus (Bramlette and Riedel) Bukry,
and forms with multilayered, unusual ellipsoidal to rectangular plate-shaped
coccoliths (Thorosphaera flabella Halldal and Markali, and Florisphaera
profunda Okada and Honjo). Two species, Emiliania huxleyi and Gephyrocapsa
oceanica, are commonly reported at all depths in the photic zone, possibly
indicating that these forms can live at various depths and temperatures.
However, this may not represent a true living distribution, but could represent
the disintegration of macro aggregates of the species at depth. Additionally, the
small cell size of E. huxleyiand G. oceanica ( < 10 p.m) may hinder their retention
by some filter-feeding zooplankton grazers (see Sect. 8). Some coccolithophorid
species have also been reported from depths as great as 4000 m in the Atlantic,
such forms are probably heterotrophic or saprophytic.
Coccolithophorids, together with dinoflagellates, predominate in the
plankton communities oftropical seas and they occur in lesser numbers in cooler
oceanic waters where diatoms are more abundant. Coccolithophorids are
particularly dominant in waters of low nutrient content, including regions of
'oceanic convergence', subtropical gyre centres (e.g. the Sargasso Sea) and in
marginal seas with anti-estuarine circulation such as the Mediterranean Sea
(Berger 1976). Over 300 living coccolithophorid species have now been de-
scribed, with the most diverse floras being those in the Atlantic. Emiliania
huxleyi is the most abundant living coccolithophorid and is "probably the most
productive lime-secreting organism on Earth" (Westbroek et al. 1986). The
species is a cosmopolitan form and has been reported from tropical to subarctic
waters.
Cold-water adapted coccolithophorids are also a consistent component of
subpolar and polar seas. Thirteen coccolithophorid species are known to survive
in the Weddell Sea, Antarctica (Thomsen et al. 1988) and 16 species occur in the
seas around West Greenland (Thomsen 1981). Few if any of these Greenland
species have a restricted distribution, and most seem indigenous throughout
boreal, subarctic and arctic areas. At least two of these species, Wigwamma
annulifer Manton, Sutherland and Oates and Papponamonas flabellifera
Manton and Oates occur in both hemispheres, which suggests that relatively
recently, in geological times, an exchange of biological material has occurred
between the polar latitudes (Thomsen et al. 1988). Species which occur at both
poles are perhaps able to withstand transport across the equator by surviving in
cold deeper wa ters, far below the photic zone (Thomsen 1981). Further studies
involving sampling of deep-water plankton below the photic zone in low
latitudes is needed to test this transport mechanism. W. annulifera also occurs in
the northern Pacific Ocean, off South Alaska (59 N, 151 W). This geographical
0 0
distribution suggests that the Northwest Passage could be a significant migration

route for cold-water plankton from the Atlantic to the Pacific Ocean (Manton
etal.1976).
Turrisphaera borealis Manton, Sutherland and Oates, Turrisphaera arctica

Manton Sutherland and Oates and Coccolith us pe/agicus (Wallich) Schiller are
species known to tolerate subzero temperatures. Turrisphaera is characterized
by unusual tower-shaped coccoliths composed of hexagonal crystallites rec-
ticulately arranged, each crystallite having a central perforation. Turrisphaera
occurs at Resolute Bay, Cornwallis Island in the Northwest Passage at a
temperature of ca. -1°C (Manton et al. 1976). C. pe/agicus has been reported
living under pack ice in the Eastern Greenland Polar Current (between Iceland
and Greenland) at temperatures as low as -1.7°C (Braarud 1979). The motile
phase of C. pe/agicus is reported to ingest bacteria, graphite and nannoplankton
cells « 5 ILm in diameter). The dual nutritional behaviour of the species may
explain its northern distribution in the Atlantic and may allow the species to
survive the low light intensities characteristic of the long dark season of the
Arctic.
The distribution of C. pe/agicus in the plankton of the North Atlantic is
limited by the maximum southerly position of the 14°C isotherm, which lies
about 40° N in the winter. In the South Atlantic, the species has been found in
postglacial cores south oflatitude 25 ° S, but was absent from all water samples.
This absence of C. pe/agicus in the Recent plankton has led to the suggestion that
until latest Pleistocene times, C. pe/agicus was still living in the South Atlantic
and that its disappearance was probably associated with postglacial warming in
the Southern Hemisphere (McIntyre and Be 1967). Although C. pe/agicus is
generally thought to be absent from the Recent plankton of the Southern
Hemisphere oceans, Hallegraeff (1984) has recorded coccospheres ofthe species
in waters off western Tasmania (latitude 40° S).
Coccolithophorids, although characteristically oceanic, also bloom sea-
sonally in coastal waters. Plankton studies in shelf waters from northwest
Europe have indicated that coccolithophorid blooms only occur in the
spring/summer period when the water column is thermally stratified with a
well-developed thermocline (Holligan et al. 1983; Murray et al. 1983). Surface
blooms of cells of E. hux/eyi may occur in numbers as high as 8.5 million cells/l
in the Celtic Sea (Holligan et al. 1983). There is, however, no record of significant
coccolithophorid growth in the tidally mixed shelfwaters of northwest Europe,
e.g. in the southern North Sea and in the central and eastern English Channel.
The abundance and species diversity of coccolith assemblages in the shelf
sediments of northwest Europe (Houghton 1988, 1989) mirror the distribution
of coccolithophorids in the overlying water. Figure 6 shows the species diversity
of coccolith assemblages in Recent sediments from the Celtic Sea and English
Channel. Highest species diversity assemblages (8 or more species, a maximum
of 18) occur in sediments from the Celtic Sea and western English Channel,
which are deposited below a seasonal thermocline (Houghton 1988). Low
diversity coccolith assemblages (six or fewer species) are deposited from the
tidally mixed waters in the central and eastern English Channel.
The restriction of coccolithophorid blooms to stratified shelf waters may be
explained by the nutrient and light preferences of the group. Coccolithophorid
growth above a thermocline is enhanced by the good light regime and is also
228 S.D . Houghton
9W 6 3 o
8 or more species
:::rrrm4 - 6 species
~ ~ ~~:~:~: 3 or fewer species
Fig.6. Species diversity of Recent coccolith assemblages in the surface sediments of the Celtic Sea
and the English Channel (after Houghton 1986)
favoured by the low nutrient levels. A diagnostic feature of many nanno-

plankton species is that the optim urn ra te of division is generally higher than tha t
of larger-net phytoplankton, following the broad generalization that rate of
uptake, growth and division is higher with reduced cell size (Raymont 1963).
Oceanic nannoplankton are thought to have a significant advantage over
larger-celled species in competition for nutrients, because they generally have
lower Ks values (Friebele et al. 1978). The Ks value of a species defines the
minimum concentration at which it can divide. Nannoplankton species such as
Emiliania huxleyi, which has a Ks value of ca. 0.1 pM nitratell (Eppley et al.
1969), are able to bloom in the nutrient deficient waters overlying a thermocline,
whereas large dinoflagellate cells which typically have large Ks values (e.g. ca.
5 flM nitrate/I) are generally excluded from the surface water plankton.
However, in tidally mixed waters, although there are sufficient nutrients for
algal growth , coccolithophorids are removed from the surface waters before any
significant growth can occur. Other low diversity continental shelf coccolith
floras associated with strong tidal mixing include examples from the Bay of
Fundy (Gran and Braarud 1935) and in the Torres Strait between Australia and
Papua New Guinea (Conley 1979).
Some coccolithophorid species prefer brackish waters and a few even prefer
freshwater occurring in rivers and lakes. The most commonly reported fresh-
water species is Hymenomonas rosa Stein (Tappan 1980). The euryhaline
characteristics of coccolithophorids as a group are evidenced by the occurrence
of 52 species in the Gulf ofElat (Aqaba), where the mean salinity is 41 parts per
thousand (ppt) (Winter et al. 1979). E. huxleyi has a wide salinity tolerance. It
has been found in the Gulf ofElat, thrives in the Black Sea (17-18 ppt), and also
occurs in the Sea of Azov (11 ppt) (Pitsky 1963). Coccolith us pe/agicus probably
has the most wide ranging salinity range, as it occurs in the Dead Sea at salinities
as high as 250 ppt. Extremely low salinities are tolerated by Cricosphaera which
occurs in supratidal rock pools influenced by wave splash (0.4-> 25 ppt).
Cricosphaera also occurs as a psammophyte, living in the interstitial waters of
intertidal sands at Roscoff (Brittany).
Although conventional plankton sampling studies have indicated that
coccolithophorids are important members of the plankton from many areas, it
is only since the development of satellite imagery that the accurate mapping of
the extent and quantitative significance of coccolithophorid blooms has been
achieved. Coccolithophorids, because oftheir external calcitic plates, have high
scattering signatures and may be recognized as high-reflectance patches at all
wavelengths in the visible band using satellite imagery (Aiken and Bellan 1988).
The recognition of coccolithophorid blooms by satellite colour scanners
(Holligan et al. 1983) indicates the potential of remote sensing techniques for the
identification and tracking of plankton blooms, and these images are essential
for the sampling requirements of international oceanographic surveys such as
the Global Ocean Flux Study (GOFS).
Coccolithophorids play an important role in marine biogeochemical cycles
and contribute to the ocean's effects on the world's climate. The secretion of
calcitic plates removes CO 2 (dissolved in the oceans as bicarbonate) from the
surface waters and transfers it to bottom sediments through the depositon of the
coccoliths. The occurrence of large blooms of coccolithophorids also helps
reduce the heat budget of the oceans by reflecting up to 30% of the incident
sunlight. Coccolithophorids are known to release volatile dimethyl sulphide
(OMS) which adds to atmospheric sulphur and probably to the acid rain of
European maritime countries. OMS also acts as a precursor for cloud con-
densation nuclei (CNN), which consequently causes a decrease in the Earth's
albedo (Charlson et al. 1987). The secretion of OMS may represent a Gaian
response to climatic warming (i.e. "the greenhouse effect") with the biota
blanketing the Earth with cloud and reducing the effect of solar heating
(Lovelock 1988; Aiken and Bellan 1988). Present studies (GOFS) on the
distribution of coccolithophorid blooms in the eastern North Atlantic will
investigate the oceanic uptake of CO 2 in the Northern Hemisphere. The North
Atlantic is well suited for such studies, since active processes in the area have
significant implications for the global carbon cycle. The Northern Hemisphere
is the major site of anthropogenic carbon input. Additionally, the North Atlantic
is thought to be a major sink for atmospheric carbon, both by the processes of
primary production and by the downwelling of CO 2 -rich water along its
northern margin. Such studies should help to determine whether coccolitho-
phorids and other marine algae are helping to modify the climatic warming
caused by the "greenhouse effect".
6 Evolution of Calcareous Nannofossils
Although there are a few reports of isolated coccolith-like bodies from

Palaeozoic sediments, calcareous nannofossils are generally thought to have
230 S.D. Houghton
..
~
........ . ...
.~
~
." '" .
=:!
t: ~
Q.
"5
"0
.D
"~
Q.
0
() 0
0 J:. ()
0
..,t:t: ~
~
~
u
~
0
u (:
:t II) ~
t
t
Fig. 7. Generalized stratigraphic range of some groups of calcareous nannofossils (after Bignot
1985)
originated in epicontinental seas during the Late Triassic (Fig. 7). The earliest
calcareous nannofossil to appear in the Triassic was Prinsiosphaera triassica
lafar (Jafar 1983). This genus comprises "spherical to hemispherical solid
nannofossils often containing a depression at one end and consisting of a
parallely stacked group of calcitic plates orientated in a random fashion" (Fig.
8a). Jafar recorded these forms from the Carnian to the Rhaetian of the Alps.
The assumed organic origin of Prinsiosphaera is based on its abundance,
stratigraphical occurrence, structural regularity and distribution in the Alpine
area. The biological affinity of Prinsiosphaera is unclear, as it has a completely
different morphology to coccoliths. However, it is probably the product of some
planktonic alga and may be related to a primitive calcareous dinoflagellate
(Bown 1987). Shizosphaerella puncticulata Deflandre and Dangeard (Fig. 8b)
appeared slightly later than Prinsiosphaera and is found in Upper Triassic/
Lower Jurassic boundary sediments. Schizosphaerella replaced Prinsiosphaera
as the dominant spherical nannofossil group in the earliest Jurassic and
they probably occupied the same niche in the epicontinental seas. Next to
evolve in the Late Triassic was Eoconusphaera zlambachensis (Moshko-
vitvitz) Bown and Cooper, a nannofossil with a truncated cone morphology
which is subcircular in plan view. The first Thoracosphaera (T. geometrica
Jafar) also appear at this time, although its relationship to younger members
of this genus is unclear.
Fig.8a-f. SEM micrographs of some Late Triassic-Early Jurassic calcareous nannofossils (a\l figures
from Bown 1987), a Prinsiosphaera triassica (x5700); b Schizosphaere/la puncliculala , side view
(x3300); c Archaeozygodiscus koessenensis. distal view (x19 800); d Crucirhabdulus minutus. distal
view (x17 400); e Mazagane/la pu/la. distal view (x7300); f Bisculum novum. distal view (x 10 500)
232 S.D. Houghton
The first 'true' coccoliths, A rchaeozygodiscus koessensis Hown (Fig. 8c) and
Crucirhabdus minutus Prins, ex. Rood, Hay and Barnard (Fig. 8d), are recorded
in the Norian Rhabdoceras suessi ammonite Zone (Jafar 1983; Bown 1987). The
comparatively sudden appearance of coccoliths and other calcareous nan-
nofossils in the Late Triassic may have been because a previously 'naked' group
of algae developed the capability to secrete micron-sized calcitic shields and
plates on their organic cell wall, with a consequent development of their fossil
record. The appearance of calcareous nannofossils followed a period of heavy
salt precipitation throughout the Permian and Triassic periods and perhaps the
salinity of the Late Triassic epicontinental seas was suitable for the secretion of
calcitic plates (Perch-Nielsen 1986). The initial location of coccolith-bearing
algae appears from previous records to have been in the Tethys Ocean. The
subtropical and tropical waters of much of the Tethys were probably the
optimum environment for coccolithophorid growth. It is also possible the
appearance of coccoliths in the Late Triassic represented the evolution ofa new
algal group which was filling a planktonic niche left vacant by the mass
mortalities of the Permian.
The next coccolith lineage to appear in the Early Jurassic was the tiered
placolith group represented by Mazaganella (Fig. 8e), although it has only been
reported from the Sinemurian and Pliensbachian of the southern Tethys area
(Bown 1987). Placoliths with radiating crystal sutures appeared in the Late
Sinemurian and are represented by Biscutum novum (Goy) Bown (Fig. 8f). The
final lineage to appear in the Early Jurassic (Pliensbachian) were placoliths with
imbricated crystal sutures (e.g. Lotharingius imprimus Bown).
Throughout the Early Jurassic new coccoliths suddenly appear with little
evidence as to their ancestry (Tappan 1980; Bown 1987). This is probably the
result of poor preservation, as occasionally sediments are found with a high-
diversity assemblage which contains some species which have no known an-
cestors. Goy et al. (1979) described a remarkably abundant and high-diversity
nannofossil assemblage from the Lower Toarcian 'Schistes Carton' of the Paris
Basin. This assemblage is the result of an exceptional preservation, a feature
rarely noted in Lower Jurassic sediments because of their diagenetic alteration.
Mitrolithus jansae (Wiegand) Brown and Young, a truncated conical nan-
nofossil, appeared in the early Sinemurian and was a dominant component of
Mediterranean-Tethys assemblages until the Early Toarcian (Bown and Cooper
1989). .
Species diversity and production initially remained low in the earliest
Liassic but then, in spite of minor fluctuation, diversity slowly increased
throughout the Early and Middle Jurassic. The first significant diversification
occurred during the early part of the Late Jurassic (Oxfordian). Oceanic
nannofossils show a peak in diversity in the Tithonian, when assemblages were
enriched with the first nannoconids and other nannofossils that flourished in the
Early Cretaceous (Roth 1983). Although nannofossil diversity steadily in-
creased throughout the Middle and Late Jurassic, nannofossil productivity
remained low, as evidenced by the rarity of pure pelagic carbonate sequences of
this age. Coccolithophores were abundant, however, in some Jurassic shelf seas
as evidenced by the Kimmeridgian coccolith limestones of the Dorset coast and

the German Solnhofen Limestone. In the Kimmeridgian and Tithonian, high
latitude nannofloras were dominated by Ellipsagelosphaera, whereas in low
latitudes Watznaueria was the dominant form (Cooper 1989). The migration of
calcareous nannoplankton from shelf and epicontinental seas into the oceanic
regime in the latest Kimmeridgian to Tithonian is reflected in an abrupt change
from impure limestone, marl and chert sequences to pure nannofossil car-
bonates. Nannofossil productivity was sufficiently high in the pelagic realm
during the latest Jurassic to depress the carbonate compensation depth (CCD)
in the Tethyan Ocean (Roth 1986). The end of the Jurassic is marked by a slight
decrease in nannofossil diversity, although only Schizosphaerella became ex-
tinct near the Jurassic/Cretacaeous boundary.
The general elevation in sea level accompanied by the development of
carbonate platforms throughout the Tethys during the Early Cretaceous
promoted the appearance of many new species and genera. Lower Cretaceous
sediments of northeast Europe yield high diversity (> 40 species/sample)
assemblages comprising a mixture of cosmopolitan, endemic, Tethyan and
Boreal-Arctic species (Crux 1989). A second marked increase in diversity
occurred in the Aptian and Albian, new forms include Broinsonia, Gatnerago
and Prediscosphaera. Most Early Cretaceous assemblages contain smaller in-
dividual coccoliths than those in the Late Cretaceous and are frequently
dominated by Ellipsagelosphaera. The widespread pelagic limestones of the
Early Cretaceous of southern Europe, and the chalks and marls of the Gulf
Coast and Mexico consist of abundant coccoliths together with nannoconids
and calpionellid tintinnids.
Maximum diversity, as well as abundance of calcareous nannofossils was
achieved during the Late Oretaceous when the worldwide chalks, which were
comprised mainly of coccoliths and their skeletal elements, were deposited.
Many of the Late Cretaceous nannofossil species were long-ranging forms
lasting for ca. 5-10 m.y. Diversity remained high during the Late Cretaceous
(> 130 species) until a decrease in the Middle Maastrichtian. At least 155 new
species originated during the 80 m.y. of the Cretaceous; a rate of ca. 2 new
species/m.y. over the whole of the period (Perch-Nielsen 1986). During the
same 80 m.y. around 126 species became extinct before the end of the Maas-
trichtian, which indicates a rate of ca. 1.5 extinctions/m.y.
At the end of the Maastrichtian came the well-documented extinctions of
marine fauna and flora and the calcareous nannoplankton suffered heavy losses
with only some 15 to 18 genera (about one-third of the Maastrichtian flora)
surviving into the Early Danian (Perch-Nielsen 1982). These species, pre-
sumably, must have comprised the total generic reservoir for the Cenozoic
evolution. Most of the surviving genera evolved during the Jurassic and have
relatively simple morphology, such as Biscutum, Braarudosphaeara and Mar-
kalius. Forms with a complex construction such as Prediscosphaera disap-
peared. Perhaps coccolithophorids which secreted simple coccoliths were able
to survive the stresses of the Cretaceous/Tertiary (K/T) boundary event,
whereas overspecialized species could not adapt to the environment and became
234 S.D. Houghton
extinct (Perch-Nielsen 1986). At low latitudes, Thoracosphaera blooms char-

acterized the plankton assemblages shortly after the boundary, although the
genus had only a sporadic distribution during the Cretaceous. None of the
nannofossil genera which survived the KIT boundary became successful during
the Cenozoic (Aubry 1989). Even the long-ranging genera Braarudosphaera and
Scapholithus, which continue through to the Recent, show no evolutionary
diversification during the Cenozoic. The other surviving genera (e.g. Biscutum
and Chiastozygus), with the exception of Markalius which continued to the
Oligocene, underwent progressive extinctions in the Palaeocene and Early
Eocene.
The mass extinctions of calcareous nannoplankton and planktonic fo-
raminifera at the boundary drastically reduced the flux of calcium carbonate
reaching the ocean floor. The boundary clay is virtually barren of pelagic
carbonate. Deposition of carbonate was reduced for at least 0.35 m.y., although
in some cases for as long as 1 m.y .. This decrease in carbonate is attributed to a
reduction in secretion of carbonate skeletons in the surface waters, rather than
a prolonged phase of dissolution on the seafloor. KIT boundary sequences also
contain a large negative anomaly in 813 C (up to 3%0) measured in the fo-
raminiferal and coccolith carbonate. This negative anomaly is considered to be
a worldwide event and probably indicates that the surface waters only supported
a very impoverished plankton. Minimum values of 813 C occurred ca. 40-50 kyr
after the boundary and did not return to pre-boundary values for at least ca.
300-400 K yr. Although the magnitude of the 813 C varies at different sites, it has
been detected only in the calcareous plankton. Primary productivity is thought
to have been suppressed for ca. 1 m.y. after the KIT boundary.
Various processes and events have been proposed to account for the mass
extinctions at the KIT boundary. These include a metorite impact event, a
severe regression at the end of the Maastrichtian, development of widespread
oceanic anoxia, temperature and salinity variations, changes in solar radiation,
nutrient limitation, and excessive volcanism. Perch-Nielsen (1986) has in-
dicated that although an iridium anomaly, microtectite concentrations andlor
shocked quartz (all features thought indicative of an extraterrestial impact)
occur at both the KIT and EocenelOligocene (E/O) boundaries, nannofossils
reacted quite differently to these events. Nannofossil abundance dramatically
decreased at the KIT boundary; however, only negligible changes occur at the
EIO boundary. This difference in the reaction of the nannofossil assemblages
may indicate a variation in the magnitude of the impact event, or may indicate
that most of the low-stress (,intolerant') forms were eliminated from the
plankton by the previous impact event. However, the nannofossil species which
became extinct at the KIT boundary represent those forms that were most
abundant and widespread, whereas those that survived were extremely rare and
had a restricted distribution. These extinction and survival patterns are not
thought to be indicative of an extraterrestial impact; such extinctions are
expected to occur at random, with the probability of survival being related to the
number of individuals in the population (Stanley 1979; Donovan 1987).
The mass extinction of calcareous nannoplankton at the K IT boundary has

recently been suggested to have caused a rapid global warming (Rampino and
Yolk 1988). The elimination of the marine calcareous phytoplankton may have
caused a severe decrease in DMS production leading to a dramatic reduction of
CCN and a concomittant decrease in marine cloud albedo. The decrease in
marine cloud albedo may have caused a rapid global warming calculated to be
> 6°C, with the highest temperature differences being recorded at higher
latitudes. Oxygen isotope analyses of marine sediments have indicated a general
warming at the KIT boundary and the resultant higher surface water temper-
atures may have been a significant factor in the suppression of the algal
productivity following the KIT boundary.
Following the major extinction of Cretaceous nannofossils at the KIT
boundary, new forms evolved in the Cenozoic, many being significantly larger
than the previous forms . Additional new and morphologically unusual forms
appeared, including Sphenolithus, Triquetrorhabdulus , Ceratolithus and Dis-
coaster. During the early Cenozoic (Fig. 9), new taxa evolved slowly at first,
rising to a total of ca . 20 nannofossil species by the end of the Danian, and up
to 60 species in the Late Palaeocene. Rapid evolution occurred in the Early
Eocene (120 species). This increase was partly due to the appearance of
Discoaster species in the Late Palaeocene (first occurrence datum at ca. 60 Ma).
The evolution of Discoaster species occurred during the heaviest values of S13 C
in CaC0 3 of any epoch in the Cenozoic. Peak values of S13 C (3.8 per mil in bulk
carbonate) occurred at ca. 60 Ma (Shackleton and Hall 1984) and indicates that
Discoaster species evolved during a period of very high surface productivity.
During the Late Palaeocene diversification of Discoaster species was rapid
(/) 40
w
U
W
Q.
(/)
III
::;)
(/)
o
~ 20
(/)
w
U
W
Q.
(/)
O~--~~---L----~----~--~~--~--~o
TIME (MILLION YEARS>
Fig. 9. Diversity of Discoasler (a) and coccolith species (b) in the Cenozoic, shown by subdivision
of the epochs (after Haq 1971)
236 S.D. Houghton
averaging 7.5 FOs/m.y. Overall nannofossil diversity then steadily declined

until the end of the Lower Miocene, when ca. 30 recorded species occurred. A
second, less dramatic, radiation of nannofossil taxa occurred during the Middle
Miocene, resulting in a total of over 50 species. Subsequently, the trend of
decreasing diversity continued until the Pleistocene. The Recent coccolitho-
phorid flora is still less diverse than that of the Cretaceous.
Within the Cenozoic, alternating diversification and decline of nannofossil
assemblages seems to be temperature related. Greatest diversity occurs in
warmer intervals, whereas less diverse assemblages consisting of cosmopolitan
taxa occur in cooler periods. Haq (1973) suggested that species with a limited
temperature tolerance may spread widely in equable seas, but when climates
became more extreme, these species had extremely reduced biogeographic
ranges which resulted in their extinction. However, the majority of the Recent
coccolithophorid species recorded in the Atlantic and Pacific oceans exhibit
eurythermal characteristics (Okada and Mcintyre 1979). The wide temperature
tolerance of coccolithophorids as a group suggests that effects other than simply
water temperature were controlling nannofossil diversity. Changes in nan-
nofossil diversity throughout the Cenozoic may also be explained by variations
in the depth of the photic zone in the world's oceans. In Recent oceans, highest
diversity coccolithophorid assemblages are found in the sub tropics and tropics,
where there is a deep (up to 200 m) photic zone, and based on coccolithophorid
assemblages, this zone has a tripartite subdivision into upper, middle and lower
photic zones (Okada and Honjo 1973). North of ca. 400N in the North Pacific,
the photic zone decreases to around 100 m and has a bipartite subdivision, with
loss of the lower photic niche and a contraction of the upper photic niche. The
consequent reduction in the availability of ecological niches lowers the diversity
of coccolithophore assemblages in these higher latitudes. During the warm
intervals of the Cenozoic, it is likely that the depth of the photic zone increased
and that the subtropical tripartite subdivision of the photic zone expanded into
more northern and southern latitudes, allowing the migration of higher diversity
coccoliths assemblages into these region. The patterns of evolution throughout
the Cenozoic are reflected in biostratigraphic zonation; nannofossil zones
during the cooler Oligocene have a duration of ca. 3 m.y., whereas warm epoch
zones average 1.7 m.y.
A major evolutionary trend shown by coccoliths throughout the Cenozoic
is a gradual reduction in the volume of calcite secreted per specimen. Palaeocene
and Eocene coccolith assemblages contain some very large (10-26 /Lm diameter)
forms, e.g. Chiasmolithus. Most Mio/Pliocene coccoliths range in size between
6 and 12 /Lm, although small recticulofenestrids < 5 /Lm may dominate Pliocene
high latitude floras. Pseudoemiliania lacunosa (Kamptner) Gartner, the first
coccolith species with slits in one shield, evolved in the latest Early Pliocene
(FOD = ca. 3.6 Ma). At about the same time small Gephyrocapsa species, forms
with a diagonal bar crossing the central pore, evolved. Small coccolith species (2
to 5 /Lm) occur throughout the Pleistocene, when the genus Gephyrocapsa
frequently dominated. Most Recent nannofloras are dominated by Emiliania
huxleyi (Fig. 13), another species with slits in its shields; the warm-water
ecophenotype of the species has slits in both shields, and the cold-water
ecophenotype has slits in one shield only.
The decrease in volume of calcite secreted per nannofossil specimen has
been suggested to be related to climatic cooling trends as carbonate is more
available in warmer waters. However, the assumption that large nannofossils
with a high density of calcite per specimen are associated with warmer waters is
not substantiated by observations of Recent plankton communities. Recent
warm-water morphovariants of coccoliths, e.g. E. huxleyi contain considerably
less calcite per specimen than the cold-water form. Other tropical forms are
fragile and of low density (e.g. U. irregularis Paasche). Cold-water coccolith
species are typically strong, imbricated, high density structures (e.g. Coccolith us
pelagicus). The same principle of increased calcite encrustment in cold-water
forms when compared with warm-water forms, is also strongly apparent in
planktonic foraminifera. Tropical forms have characteristically fragile, perfo-
rate tests.
The gradual reduction in volume of calcite secreted per coccolith specimen
is also reflected in Discoaster evolution (Bukry 1971). Most Palaeogene Dis-
coaster assemblages are characterized by compact, multirayed, heavily con-
structed forms with large central knobs. In plan view these early Discoaster have
a short 'free length' of rays, or the rays are very broad and the area encompassed
by calcite is very high (> 80% cover in plan view) (Fig.lOm-p). However, a few
more delicately constructed forms with a longer free length of ray also occur and
include D. okadai Bukry (ca. 28% calcite cover) and D. lodoensis Bramlette and
Riedel (ca. 40-50%). Mid-Cenozoic Discoaster species have a reduced number
of rays, although in plan view there is still a substantial area ofcalcite (Fig. lOi-k).
Discoaster assemblages in the Late Pliocene are restricted to very narrow-rayed
forms (Fig. lOa-e). The skeletons of these represent a mere framework « 25%
cover) of the massive Discoaster 'shields' typical of the Palaeogene. Only two
Discoaster species occur just before their extinction in the latest Pliocene at ca.
1.9 Ma: D. brouweri, Tan emend Bramlette and Riedel, a six-rayed form and, D.
triradiatus Tan, a three-rayed form.
The occurrence of a wide spectrum of Discoaster morphologies, even within
the same assemblage, was probably ecologically controlled and suggests that the
forms were depth-stratified. Test thickening and calcite crust formation is well
documented in deep-water planktonic foraminifera (e.g. Orr 1967) and high
density 'solid' coccoliths are typical of the lower photic zone in the Recent
Oceans (see Sect. 5). A similar depth-control on calcite secretion may have
occurred in the Discoaster group. It is likely that changes in the amount of
calcification of the Discoaster skeletons affected the passive bouyancy of the
Discoaster-bearing algae. Bouyancy in the water column may be accomplished
by numerous ways including a reduction in the specific gravity of the cell, by
increasing the surface-to-area ratio, or by added resistance. Coccolith calcite has
a density of between 2.7 and 2.95, whereas seawater varies between 1.021 and
1.028 (Tappan and Loeblich 1973). A reduction in specific gravity may be
achieved by economic use of skeletal secretion. (Parsons and Brasier 1987). Be
(1968) has shown that in Recent planktonic foraminifera populations, surface
238 S.D. Houghton
Fig. lOa-po Discoaster shape variation (mainly after drawings by Prins 1971). a Discoaster chal-
lengeri Bramlette and Riedel (M. Miocene- L. Pliocene); b D. pentaradiatusTan emend. Bramlette
and Riedel (M. Miocene- L. Pliocene); cD. tamalis Bramlette and Riedel (L. Pliocene) ; dD. brouweri
Tan emend. Bramlette and Riedel (M. Miocene- L. Pliocene); e D. surculus Martini and Bramlette
(L. Miocene- L. Pliocene); f D. kugleri Martini and Bramlette (M . Miocene); g D. quinqueramus
Gartner (L. Miocene) ; h D. neohamateus Bukry and Bramlette (L. Miocene); i D. bollii Martini and
Bram lette (M.- L. Miocene); j D. adamant us Bramlette and Wilcoxon (Oligocene- L. Miocene); k D.
deflandrei Bramlette and Riedel (Eocene- M. Miocene); I D. broennimannii Stradner (L. Eocene);
m D. multiradiatus Bramlette and Riedel(U. Palaeocene- L. Eocene) ; nD. gemmeusStradner(L.-M.
Eocene): 0 D. munitus Stradner (L. Eocene) ; p D. mirus Deflandre (L. Eocene)
dwellers have tests with larger mural pores and pore densities than deeper
cold-water forms.
The heavily calcified Late Palaeocene Discoaster forms such as D. bram-
letfei (Bukry and Percival) Romein and D. multiradiatus Bramlette and Riedel
(Fig. 10m), which had greater than 90% calcite cover in plan view, probably
evolved and lived deep in the photic zone of the water column. More delicately
constructed forms such as D. okadai (28% cover) and D. lodoensis (40-50%) lived
near the surface. The trend towards an increased dominance of narrow-rayed,
weakily encrusted forms throughout the Cenozoic may reflect an increased
preference of the algal group for a shallow-depth habitat. As the group migrated
upwards through the water column, forms preferring deeper water became
extinct but were not replaced by new forms. During the cool Oligocene Epoch,
Discoasterdiversity was severely restricted , dropping to less than five species per
nannofossil zone. Oligocene Discoaster species are dominated by the D. tanii
group, simple six- and five-rayed forms with untapered arms (ca. 40% calcite
cover). Two other common species D. adamanteus and D. deflandre , are heavily
calcified (ca. 70- 75% cover) forms in which three of the six-rays are fused to form
a single crystal and are separated from the other fused crystal unit by well-
defined interradial sutures (Black 1972). The Oligocene represents the only
Epoch when weakily encrusted « 33% cover) forms do not occur in Discoaster
assemblages. This may be explained by an absence of Discoaster species in an
upper photic habitat this time, or perhaps because of the cool temperature of the
Oligocene oceans. the Discoaster-bearing algae favoured the secretion offused
and heavily calcified forms, even in a shallow-water niche. Alternatively, their
absence may be due to a period of enhanced calcite dissolution during the
Oligocene.
Most of the broad-ray, heavily encrusted, Discoaster species (e.g. D.
deflandrei Bramlette and Riedel and D. adamanteus Bramlette and Wilicoxon)
became extinct in the early Miocene, and this suggests that a niche became
available for forms that were adapted for a lower photic zone habit. The
narrow-rayed Discoaster forms dominant in the Late Pliocene were probably
restricted to surface-dwelling phytoplankton populations. Many of the Dis-
coaster species which appeared in the Miocene and Pliocene deviate from a
discoid form and develop arms which are slightly bent (D. brouweri, D. pen-
taradiatus, D. berggrenii and D. triradiatus) and which tend to raise the central
structure of the Discoaster above the tips of the arms in an umbrella-like
construction. Assuming that Discoaster specimens represented a skeletal
covering for some near spherical-shaped algal cell (Deflandre 1952), the in-
creased curvature displayed by these younger species suggests that the group
evolved towards forms with a smaller cell volume.
Detailed studies on the evolutionary processes of calcareous nannofossils
are still relatively few. Deep ocean drilling undertaken by the Deep Sea Drilling
Project and subsequently by the Ocean Drilling Program should provide
sufficient opportunities to evaluate the process of nannofossil evolution and
help test the theories of gradualism versus punctuated equilibrium. Oceanic
sediments facilitate such studies as sample retrieval is usually good (100/s of
specimens of the same species occur in one assemblage), morphological char-
acteristics are well-preserved, and a good latitudinal coverage helps ensure that
evolutionary patterns are separated from ecologically controlled migration.
Ideal oceanic sequences for evolutionary studies should be characterized by
high sediment accumulation rates, little or no dissolution, low amounts of
bioturbation and should be integrated with established magnetostratigraphies
(Backman 1986).
7 Calcareous Nannofossil Biostratigraphy
The object of biostratigraphy is to group strata into units on the basis of the
variety and abundance of their contained fossils. Biostratigraphers use various
biozones (usually referred to simply as 'zones') as a means for the calibration of
stratigraphic sequences. Four types of biozone are employed in stratigraphical
classification:
I. Assemblage Zone: strata characterized by a distinctive fossil assemblage;

2. Range Zone: strata representing the stratigraphic range ofa particular fossil;
240 S.D. Houghton
3. Acme Zone: strata characterized by the acme or maximum abundance of a

certain form;
4. Interval Zone: the stratigraphical interval between two biohorizons.
Most nannofossil zonations utilize 'interval zones' whose boundaries are

considered synchronous biohorizons, represented by unique events such as the
evolutionary first appearance and extinction of particular species.
At present there is no calcareous nannofossil zonation for the Late Triassic
(Carnian-Rhaetian). Biostratigraphical zonation for the Jurassic is based
mainly on the first occurrence of one nannofossil species relative to the first
occurrence of another. Bernard and Hay (1974) produced the first tentative
zonation ofthe Jurassic of southern England and France. Perch-Nielsen (1985a)
reviewed the zonation schemes for the Jurassic and concluded that the zonations
are far from satisfactory. However, Bown (1987) has recently updated the
nannofossil zonation for the Lower Jurassic of northwest Europe. Additional
work still needs to be done to improve the nannofossil biostratigraphy of the
Middle and Upper Jurassic. Important biostratigraphical markers for the
Jurassic include species of Parhabdolithus, Biscutum, Discorhabdulus, Ste-
phanolithion and Stradnerlithus. The first occurrence of Nannoconus is almost
synchronous with the Jurassic/Cretaceous boundary, occurring in the latest
Tithonian. Species which have their first occurrence just above the boundary, in
the earliest Berriasian, include Rotelapillus laffittei (Noel) Noel and Lithra-
phidites carniolensis Deflandre.
The nannofossil zonation of the Cretaceous (particularly the Late Cre-
taceous) is more advanced than that of the Jurassic and has been reviewed by
Perch-Nielsen (1985a). Crux (1989) has recently refined the biostratigraphy of
the Lower Cretaceous (Ryazanian- Barremian) of northwest Europe into 16
nannofossil zones, and has integrated these zones with the ammonite and
belemite zonantions. Important biostratigraphical markers for the Cretaceous
include forms of Reindarditites, Eiffellithus, Quadrum, Micula, Tranolithus and
Prediscosphaera. Future biostratigraphical studies of the Upper Cretaceous
should concentrate on the integration of the calcareous nannofossil and
macrofossil (belemnite and ammonite) zones and their correlation with the
magnetostratigraphic and chronostratigraphic scales.
The Cretaceous/Tertiary boundary may be placed at the first occurrence of
Bianolithus sparsus Bramlette and Martini and/or Cyclagelosphaera alta
Perch-Nielsen (Romein 1979; Varol 1989). The standard zonation schemes of
Martini (1971), Bukry (1973), and Okada and Bukry (1980) are usually used for
the subdivision of the Cenozoic. Finer subdivisions have been developed for
local environments; however, these schemes still use the Martini or Bukry
zonations as a framework. Martini (1971) used the letters NP and NN and
numbers to designate the Nannoplankton Palaeogene zones and the Nanno-
plankton Neogene Zones respectively. Okada and Bukry (1980) proposed CP
and CN (Coccoliths Palaeogene and Coccolith Neogene) to codify the original
zones of Bukry (1973).
The standard Palaeogene nannofossil zonation of Martini (1971) is shown
in Fig. 11 b. Early Palaeocene subdivision is accomplished by the first occurrence
Martini.1971 MY Martini.1971
MY
CD
c: NN21 Emilianla hux/eyi CD 25.0
CD f-0.27 (;j Sphenolithus
NP25
....:c
0
0 NN20 Gephyrocapsa CD
...J clperoensis
;; oceanica c: 2S.2
.-
CD
-0.47 CD
0 NP24 Sphenolithus distentus
Q. Pseudoemiliania 0 "0 30.2
NN19 lacunosa ~fhenolithus
-:E
1.66- OJ NP23 redistentus
34.6
-1.89 0 NP22 Helicosphaera
>- reticulata
~ 35.1
NN18 Olscoaster brouweri w
CD NP21 Ericsonia subdisticha
(;j -2.33
...J
f-- - 36.7
Spheno/ithus
CD
c:
NN17 O.pentaradiatus NP20 pseudoradians
CD -2.43
0
NN16 O.surculus NP19 Isthmolithus recurvus
.- r-- -3.56
-Q.
0
37.S
Reticulofenestra
NN15 pseudoumbi/lca CD
~ - 3.7 (;j Chiasmolithus
:;; NN14 Oiscoaster ...J NP1S oamaruensis
w - asymmetrtcus
4.1
NN13 Ceratolithus rugosus
f- 4.5
r-- -
NN12 Amaurolithus - 39.S
I
tricorniculatus
r- 5.6 CD NP17 Oiscoaster saipanensis
c:
CD
0 42.3
Oiscoaster 0
Oiscoaster tani
NN11 quinqueramus w NP16
CD nodifer
(;j CD
...J :c 46.0
"0 NP15 Nannotetrina fulgens
:E 49.S
r- 8.2 .- NP14 Oiscoaster
sublodoensis
NN10 Oiscoaster ca/caris 52.6
NP13 Oiscoaster lodoensis
f-S.S5 .- 53.7
Tribrachiatus
- NP12 orthos.tylus
NN9 Oiscoaster hamatus ~
:;; 55.3
CD f-10.0 w NP11 Oiscoaster binodosus
c: NNS Catinaster coalitus 56.3
CD
.!! f-10.S NP10 Tribrachiatus contortus
-
0
.-
0 "0
"0
NN7 Oiscoaster kug/eri - :;7.S
::;: :E f-13.1 Oiscoaster
NN6 O/scoaster exilis NP9 mu/tiradiatus
r-14.4 59.2
- NN5 Sphenol/thus hetero-
morphus NP8 Heliolithus riedelil
CD
f-16.2 (;j 60.0
Helicosphaera NP7 Oiscoaster mohleri
NN4 ampiliaoerta CD ...J
f-17.4 c: 60.4
~ NN3 Sphenolithus CD NP6 Heliolithus kleinpel/ii
be/emnos 0
~. r- 21.5 0 61.6
NN2 Oiscoaster druggii CD F asciculithus
NP5 tympanlformls
r-23.2
-asas f-- 62.0
23.7- NN1 T.carinatus Q. NP4 Ellipsolithus macel/us
24.0 63.8
>- NP3 Ch/asmolithus danicus
64.8
a ~ NP2 Cruciplacollthus tenull
w
65.9
NP1 Markal/us Inversus
b 66.4
Fig.n. a Neogene nannofossil zonation of Martini (1971) integrated into the chronostratigraphic
framework of Berggren et al. (l985b); b Palaeogene nannofossil zonation of Martini (1971)
integrated into the chronostratigraphic framework of Berggren et al. (l985a)
242 S.D. Houghton
of coccoliths such as Cruciplacolithus, Chiasmolithus and Ellipsolithus. Within

the Heliolithus kleinpelli Zone (NP6) the first occurrence ofthe Discoaster group
(D. bramletii) occurred. Subsequent subdivision of the Late Palaeogene is
accomplished by the first occurrence of Discoasterspecies (D. mohleri Bukry and
Percival, D. nobilis Martini and D. multiradiatus Bramlette and Reidel). An
improved biostratigraphical zonation scheme for the Palaeocene has recently
been proposed by Varol (1989). The zonation, applicable in low to mid-lati-
tudes, comprises 20 zones and 24 subzones (prefixed NTp, Nannofossil Ter-
tiary-Palaeocene) and is defined upon last occurrences of species and their
acmes. The Palaeocene/Eocene boundary is usually placed at the top of
NP9/CP8, which approximates to the last occurrence of Fasciculithus tym-
paniformis Hay and Mohler (Varol 1989) and the first occurrence of the
planktonic foraminifera Pseudohastigerina Perch-Nielson 1985b). The Eocene
(zones NP 10-20) is subdivided by forms of Tribranchiatus, Discoaster,
Chiasmolithus, Isthmolithus and Sphenolithus. Designation of zones NPI8-
NP20 may be difficult in low latitudes because of the scarcity of the zonal fossils
in the assemblages.
The Eocene/Oligocene boundary, defined in terms of nannofossils, is
usually placed at the top of NP20 which corresponds to the last occurrence of
disc-sha ped D iscoaster species (D. barbadiensis Tan and D. saipanesis Bramlette
and Riedel). This event occurs slightly younger than the disappearance of the
planktonic foraminifer Hantkenina (Perch-Nielsen 1985b). Throughout the
Oligocene species, diversity is low and zonation is accomplished by the first
occurrence and last occurrence of forms of Ericsonia, Helicosphaera and
Sphenolithus. The Oligocene/Miocene boundary is usually placed at the top of
NP 25 or within NN 1.
The standard Neogene nannofossil zonation of Martini (1971) is shown in
Fig. Ila. The subdivision of the Miocene is based predominantly on species of
Discoaster and Sphenolithus. Biostratigraphic zonation is hindered in high
latitudes and in marginal sea assemblages, where Discoaster species may be
scarce or absent. New, medium to large-sized, placolith species which appeared
in the Miocene include Calcidiscus macintyrei (Bukry and Bramlette) Loeblich
and Tappan (NNl) and Reticulofenestra pseudoumbilica (NN2) (VaroI1985).
Within the NN9 Discoaster hamatus Zone is the first occurrence of new five-
rayed Discoaster species which include D. hamatus Martini and Bramlette, D.
bellus Bukry and Percival and D. prepentaradiatus Bukry and Percival. The first
occurrence of Amaurolithus primus (Bukry and Percival) Gartner and Bukry, the
oldest horseshoe-shaped calcareous nannofossil in the Neogene, occurs in D.
quinqueramus Zone (NN 11). Other species of Amaurolithus appear soon after A.
primus. The upper part of Zone NN 12 is characterized by the first occurrence of
birefringent ceratoliths (Ceratolithus). In some high latitude Pliocene North
Atlantic sediments only the last occurrence of R. pseudoumbilica (3.56 Ma)
and/or the first occurrence of P. lacunosa (ca. 3.6 Ma) may be used for
subdivision. Zonation of the Late Pliocene is based on a series of extinction
datums of Discoaster species; in chronological order these include D. tamilis
Kamptner, D. asymmetricus Gartner, D. surculus Martini and Bramlette, D.
pentaradiatus Tan emend. Bramlette and Riedel and D. brouweri. The last 0.18
m.y. of the range of the Discoaster group (2.07-1.89 Ma) is characterized by a
higher (> 20%) proportion of D. triradiatus and this forms a useful indication for
the extinction of the group and an important biostratigraphical event, even in
the presence of considerable reworking (Backman and Pestiaux 1986).
The Pliocene/Pleistocene boundary, now placed at 1.66 Ma, occurs between
the extinction datums of D. brouweri at 1.89 Ma and C. macintyrei at 1.7-1.45
Ma. The boundary, in terms of nannofossils, is therefore placed towards the base
of the P. lacunosa (NNI9) Zone. The Pleistocene is primarily subdivided by
small placolith species and also by C. macintyeri and H. selli. The NN 19 Zone
of Martini (1971) can be subdivided by nannofossil events which occurred in the
following order; first occurrence of G. oceanica s.l., last occurrence of C.
macintyrei and last occurrence of H. sellii Bukry and Bramlette (Fig. 12). The last
occurrence of H. sellii, however, is diachronous across latitudes, ranging
between, 1.04 and 1.42 Ma (Backman and Shackleton 1983; Pujos 1988).
Thierstein et al. (1977) studied the global synchroneity of Middle and Late
Pleistocene nannofossil datums. The most important nannofossil event in the
NANNOFOSSIL ZONES
Pseudoemil/ania lacunosa Zo
small Gephyrocapsa Zone

1 - - - - - - - - - - - - - 1 NN19
Helicosphaera Sellii Zone
Calcidiscus maclnlyrei Zone
Discoasler brouweri Zone NN18
Fig. 12. Calcareous nannofossil zonation schemes for the Pleistocene

244 S.D. Houghton
Middle Pleistocene is the last occurrence of P. lacunosa" which marks the top of
the NN 19 Zone. This occurs as a globally synchronous datum within the middle
of oxygen isotope stage 12, dated at0.458 Ma. The first occurrence of E. huxleyi,
which designates the NN20/NN2l boundary, occurs consistently late in oxygen
isotope stage 8, dated at 0.268 Ma.
The dominance reversal of Gephyrocapsa spp.1 E. huxleyi has been used to
subdivide NN21 Zone. During the first two-thirds of its range E. huxleyi occurs
in relatively low abundances and is subordinate to Gephyrocapsa; however, it
subsequently becomes the most dominant form. The Gephyrocapsal E. huxleyi
reversal in dominance is time-transgressive. In tropical and subtropical
sediments it occurs between oxygen isotope stages 5a and 5b (0.085 Ma),
whereas in transitional sediments it correlates with oxygen isotope stage 4c (ca.
0.073 Ma). The dominance of G. oceanica and the rarity, or even absence of E.
huxleyi in sediments underlying monsoon-influenced regions of the Indo-
Pacific (see Sect. 9.2) may not allow the NN20 and NN21 Zones of Martini
(1971) and the E. huxleyi Acme Zone of Gartner (1977) to be separated. In such
assemblages where E. huxleyi is rare because of an ecological control, an
overestimation of the age of the sediment may occur. Heckel (1973) also
indicated that the absence of E. huxleyi from nearshore sediments from the
Capricorn Basin was probably a facies-control, and should not necessarily be
used as evidence for exculsion of a sediment from the E. huxleyi Zone. A
potentially useful biostratigraphical species which may be used as an alternative
marker for the E. huxleyi Zone in the Indo-Pacific region is Geminilithella
subtilis (Muller) Yarol. This species was used by Hughes et al. (1986) to date
uplifted, latest Pleistocene sediments in the Solomon Islands.
Calcareous nannofossils have several characteristics which facilitate their
use as biostratigraphical markers. Many nannofossil species have been shown
to have synchronous extinctions and first appearances (Backman and Shack-
leton 1983; Backman and Pestiaux 1986; Pujos 1988); therefore, their datums
may be used with some degree of confidence, regardless of water mass char-
acteristics. As a group, nannofossils are far more eurythermal than planktonic
foraminifera. This enhances their use in oceanic correlation because zonal
species frequently have a cosmopolitan distribution. Nannofossils may be
recovered from a surprisingly wide spectrum of sediment types, ranging from
fine-grained silts and clays to sands and muddy gravels. Their occurrence in
coarse-grained sediments may be attributed to their mode of settling, which is
primarily within large faecal pellets of zooplankton grazers (see Sect. 8).
8 Sedimentation, Dissolution and Diagenesis of Nannofossil Assemblages
The extent to which a calcareous nannofossil assemblage preserved in bottom

sediments reflects the living floral assemblages in the surface water depends
upon the processes of sedimentation, dissolution and diagenesis. Dissolution
probably is the most important ofthese, and begins soon after the organism dies
and sinks into waters undersaturated with respect to calcium carbonate. Dis-
solution is inhibited by the fact that most nannofossils are thought to descend to
the seafloor within the faecal pellets of zooplankton grazers. Copepods are
major grazers of coccolithophorids (Honjo 1976), and daily may consume up to
1200 individuals, ejecting the calcite plates in some 200 faecal pellets. A single
pellet may contain up to 100000 coccoliths, which represents ca. I /Lg ofCaC0 3
(Ta ppan 1980). Although the gut of cope pods is though t to be acidic, labora tory
studies (Honjo and Roman 1978) have shown that coccoliths contained in
copepod faecal pellets (produced by Calanus and Acartia species) showed no
signs of mechanical destruction or chemical dissolution. Even fragile coccolith
specimens of the macrococcoliths of Umbellosphaera irregularis and many
species of holococcoliths remained intact. Large cocco1ithophore cells such as
Syracosphaera (ca. 30 /Lm) and Pontosphaera (ca. 50 /Lm) were found as isolated
coccoliths within faecal pellets, but Emiliania and Gephyrocapsa « 10 /Lm) were
often found to occur as intact coccospheres. This suggests that small cells of ca.
10 /Lm are inefficiently ingested and may be the lower size limit of cope pod
retention for filter feeding.
An organic coating envelops faecal pellets and seals the coccoliths from
corrosive waters throughout the major part of their descent (Honjo 1976). The
descent of the coccoliths within the "faecal pellet express" accelerates deposi-
tion dramatically, estimated to be ca. 100 years in the oceans for an isolated
coccolith to a few weeks for a mean-sized faecal pellet. This mode of deposition
ensures that a constant flux of coccolith carbonate, relatively free from dis-
solution, reaches the bottom sediment, and also promotes a better correspon-
dence between plankton assemblages and fossil assemblages. In the equatorial
Pacific, 92% ofthe coccolith carbonate produced in the surface waters may reach
the bottom within faecal pellets (Honjo 1976).
Faecal pellets, although protecting coccoliths during descent through the
water column, rapidly disintegrate on the seafloor because of bacterial
action. Once coccoliths spill out of the pellets they become susceptible to
dissolution. In deeper waters a coccolith lysocline (the depth below which all
coccolith skeletons show signs of dissolution) has been identified between 3- and
4-km depth. Selective dissolution ofless resistant species occurs between these
depths; dissolution increases dramatically below 4 km resulting in preservation
of only solution-resistant nannofossil assemblages dominated by placolith-type
coccoliths. Below the carbonate compensation depth (CCO) which separates
predominantly calcareous sediments from carbonate poor pelagic sediments,
few nannofossils survive. In the North Pacific, the coccolith CCO varies con-
siderably (Roth and Coulbourn 1982), with deepest values (5500 m) being found
in the Central Equatorial Pacific, and shallower values occurring towards the
margins of the basin, particularly in the Transition Region of the California
Current, where it may shoal to a depth of 3300 m.
Sedimentation of coccoliths in macroaggregates other than faecal pellets
has been described from several localities. Macroaggregates (ca. 300 /Lm)
consisting of ca. 15 000 intact cells of E. huxleyi (Fig. 13) embedded in mucoid
material were recorded from a sediment trap in the Fladen Ground, North Sea
(Cadee 1985). Other macroaggregates have been described of Oolithotus fragilis
246 S.D. Houghton
Fig. 13a-c. SEM micrographs of

Emiliania huxleyi. a Distal view of
the warm-water ecophenotype
showing well-preserved T-shaped
elements (x33 000). b Distal view,
showing solid proximal shield and
central ring structure but with dis-
tal T-shaped elements absent
(x17 000); c Both proximal and
distal shields absent, remnant cen-
tral ring preserved alone (x 13 000)
('palmellid' stage) from the Mediterranean (Bernard 1963) of Umbilicosphaera

sibogae (Weber-van Bosse) Gaarder in traps from the Panama Basin after a
bloom of the species in the surface water (Honjo 1982), and clusters of Ge-
phyrocapsa oceanica cells from Australian coastal waters (Hallegraeff 1984).
Such aggregates of algal cells are thought to facilitate rapid removal from
inhospitable environments that may occur in the surface waters (Smetacek
1985). Sedimentation of coccoliths in macroaggregates is probably an important
mode of transport of coccoliths to the seafloor and may be widespread among
coccolithophorids, particularly in shelf seas where environmental conditions
are liable to fluctuate rapidly.
Berger (1970) has estimated tha t a bout four-fifths of the calcite supplied to
the ocean bottom is being redissolved; however, nannofossil skeletons are more
resistant to this effect than other microfossil tests. Part of this increased resis-
tance to dissolution is attributed to the incorporation of acid-resistant, cel-
lulose-like polysaccharide compounds within the skeletal calcite (Franke and
Brown 1971). Organic coatings on the surface of coccoliths are also thought to
inhibit dissolution of coccolith carbonate. The dissolution characteristics of
nannofossils are utilized by species of tintinnid and planktonic foraminifera
which incorporate nannofossils into their wall structure (Fig. 14a).
Initial dissolution of coccoliths is reflected in etching of individual crys-
tallites resulting in rounded corners and serrate margins of shields (Fig. 3d).
Fig. 14. a Detailed view of planktonic foraminifera test of Globigerina (Zeaglobigerina) aperta
Cushman showing incorporation of placolith coccoliths into the wall structure (x800). b Well-
preserved distal view of Coccolith us pelagicus showing no overgrowth (x4200). c Distal view of
Coccolith us pelagicus showing overgrowth on radial elements, although most of the crystallites can
still be distinguished (x5200). dDistal view of severely overgrown coccolith of Coccolith us pelagic us,
some of the radial crystallites have overgrown at the expense of others (x6000)
Usually coccoliths constructed of smaller crystallites are more susceptible to

dissolution. More advanced stage dissolution isolates the individual elements of
coccoliths and removes loosely attached plates. Finally, disintegration of the
coccolith may result in the production ofa mass of micron-sized coccolith debris.
Figure 13a-c shows coccoliths of Emiliania huxleyi in various stages of dis-
solution. Initial dissolution of coccoliths of E. huxleyi occurs at the rim of the
structure within the T-shaped elements. High resolution electron microscopy of
the T -sha ped elements has shown that they consist of a mosaic of microdomains
of 300-500 A in diameter, whereas the crystal lattice of the more resistant
radially arranged elements is continuous (Parker et al. 1983). The central ring of
these coccolith is the most resistant structure in this species (Fig. 13c).
Post-depositional diagenetic overgrowth on nannofossil specimens com-
monly occurs in Cenozoic and Cretaceous sediments (Bukry et al. 1971 ; Roth
1973). Such processes, although they may be used as a tool for identifying early
diagenetic processes within sediments, often cause taxonomic problems, and
may alter the apparent diversity of assemblages. In the deep ocean, overgrowth
248 S.D. Houghton
on nannofossil specimens seems to be related to the early lithification of

calcareous sediments. Diagenesis of nannofossil oozes, which consist almost
exclusively of low Mg calcite, occurs by deposition of syntaxial overgrowth
cement on Discoaster species, coccoliths, foraminifera and other fragments
(Matter 1974). Precipitation of overgrowth onto Discoaster rays occurs soon
after burial, usually preceding calcite overgrowth on coccoliths which is thought
to be initiated at ca. 100 m depth below the seafloor. Wise and Kelts (1972)
suggested that these differences in the timing of overgrowth are caused by
differences in the construction of nannofossils: Discoasterrays are usually single
crystals and act as large seed crystals similar to echinoderm fragments, whereas
coccoliths consist of many sub-micron sized elements which have varying
optical orientation. Laboratory evidence (Adelseck et al. 1973) has indicated
tha t Discoaster ra ys and large placoliths grow at the expense of small coccoliths
under simulated diagenetic conditions. In the south and central Pacific,
secondary calcite is reported to occur regularly only on the large coccolith
species Calcidiscus leptoporus and Umbilicosphaera sibogae, and is usually
restricted to the distal shield (Roth and Berger 1975). Overgrowth on placoliths
is usually apparent by the abnormal thickening of the distal shield crystallites
(cf. Fig. 14b-d).
Results from the Deep Sea Drilling Project taken from many ocean depths
have shown that the most solution-resistant Cenozoic genera, Coccolith us,
Calcidiscus, Reticulofenestra and Dityococcites, occur in most deep-water and
shallow-water calcareous sediments. Solution prone genera absent from deeper
water sediments include Micrantholithus, Tranversopontis, and Scyphosphaera.
Overgrowth on nannofossils can also occur in surface sediment from
near-shore environments. Greater than I % of nannofossil assemblages in late
Quaternary sediments from the Gulf of Aqaba (Elat) and the Red Sea were
reported to have overgrowth (Winter 1982a). Although overgrowth on nan-
nofossils in the deep oceans is usually associated with differential solution,
similar processes are not thought to occur in the 1800-m-deep gulf, where the
entire water column is supersaturated with CaC0 3 • Calcite overgrowths on the
nannofossil assemblages from the Red Sea were therefore concluded to occur
within the sediment because of the supersaturation of the water column and
interstitial waters.
9 Calcareous Nannofossils as Palaeoclimatic and Palaeo-Oceanographic

Indicators
9.1 Calcareous Nannofossil Assemblages as Indicators of Palaeodepth
Coccolithophorids, because of their predominantly planktonic habit, are not

thought to be particularly useful as indicators of the depth of the sediments in
which they are found. However, holococcoliths are usually restricted to as-
semblages deposited at shallow depths up to a few 100 m. Some caution must be
applied to this general conclusion, as holococcoliths may survive transport into
greater depths by turbidity currents. Okada (1984) has used the abundance of
Florisphaera profunda Okada and Honjo in marginal seas along the western
Pacific Ocean as a tool to determine palaeodepth. F. profunda was not recorded
in shallow seas but was found to increase to over 50% of the assemblages at ca.
2000 m depth. It is generally accepted that Recent Braarudosphaera bigelowii has
highest abundances in near-shore areas and despite a few exceptions is very
scarce or absent from pelagic sediments. In bottom sediments from Sendai Bay,
Japan, the B. bigelowii component of the nannofossil assemblages increased
from < 10% in deeper water sediments (> 100 m depth) to nearly 50% in
near-shore sediments shallower than 24 m (Takayama 1972).
Quantitatively, coccoliths are generally less abundant in shallow, near-
shore sediments than in open-oceanic sediments. Heckel (1973) reported that in
fine fractions of sediments from the Capricorn Basin deposited above 40-m
water depth coccoliths were rare « 2%); the coccolith component increased to
over 10% in water depths greater than 100 m. In the seas around Britain, Recent
coccolith abundances decrease from 109 coccolith per gram (> 10% of the fine
fractions) in deep water sediments (> 100 m depth) from the Celtic Sea to < 10 7
coccolith per gram « 0.1%) in sediments shallower than 30 m in the English
Channel (Houghton 1986, 1988).
9.2 Calcareous Nannofossil Assemblages as Indicators of Marginal

Sea/Open Oceanic Sediments
Thierstein (1976) recorded the differences between Cretaceous nannofossil

assemblages in the Tethys and those found in Pacific oceanic sites. Species
present in the epicontinental and marginal seas of the Tethys but absent from
the oceanic Pacific sites include Nannoconus spp., Conusphaera mexicana Trejo,
Micrantholithus obtusus Stradner and Lithastrinus floralis Stradner. Atlantic
Cretaceous marginal sea and shelf assemblages contain abundant Broinsina and
Nannoconus, whereas oceanic Atlantic assemblages are dominated by Watz-
naueria barnesae (Black) Perch-Nielsen (Roth and Bowdler 1981). In the
Cenozoic, genera which seem to be more common in shelf areas rather than in
oceanic conditions, include Pontosphaera, Scyphosphaera, Braarudosphaera,
Micrantholithus and Pemma. Discoaster is a typical component of oceanic
Cenozoic assemblages particularly at low mid latitudes. The genus has not been
recorded in the Upper Pliocene shelf assemblages from the Capricorn Basin
(Hecked 1973) and is also absent in the near-shore Upper Pliocene St. Erth Beds
and Coralline Crag of England (Jenkins and Houghton 1987; Houghton and
Jenkins 1988).
Okada and Honjo (1975) investigated the Recent coccolithophorid dis-
tribution in plankton samples from the Pacific marginal seas of Asia. They
concluded that the dominance of Gephyrocapsa oceanica and the very sparse
occurrence of Em iliania huxleyiwas diagnostic of coccolith assemblages in these
marginal seas. With minor exceptions, G. oceanica was noted to predominate in
the plankton of the East China Sea, South China Sea, Java Sea, Timor Sea and
250 S.D. Houghton
1 1
K
•
G.oceanlca Asia
Africa
Indian Ocean
~ G. oceanica dominant
~ E. huxleyl dominant
~ Neither specie's dominant
40 E0 'SOGE
Fig. 15. Map showing areas of dominance of G. oceanica and E. hux/eyi in Recent coccolith
assemblages from the marginal seas of the Indian Ocean and western Pacific Ocean, including both
sediment and plankton samples, compiled from many sources. Marginal seas with no shading
represent areas with no suitable coccolithophorid data
off the Northern coast of Australia (Fig, 15). The species was most dominant in
coccolithophorid samples from the South China Sea (95-100% of nannoflora).
A trend of increasing dominance of the species in the plankton towards land was
also recorded, Similar reversals in dominance of E. hux/eyi over G. oceanica
towards the coast have also been noted in sediments from the Great Barrier Reef
(Heckel 1973; Conley 1979), the west Tasman Sea (Burns 1975) and in the East
China Sea (Wang and Samtleban 1983). G. oceanica is also the most abundant
species in assemblages from the Bay of Bengal, the Arabian Sea and in the
Adaman Sea in the northern Indian Ocean (Guptha 1976, 1981). Coccolitho-
phorid assemblages in Australian coastal waters have a latitudinal dominance
shift from G, oceanica-dominated in the north (North West Shelf, Coral Sea and
northern Great Barrier Reef), to E. huxleyi-dominated in the south (Tasman
Sea, Great Australian Bight; Hallegraeff 1984).
The abundance of G. oceanica in the marginal seas of the Indian and Pacific
oceans is certainly not characteristic of shelf waters from the North Atlantic,
where from the tropics to the subarctic, E. huxleyi is by far the most abundant
species (sometimes forming a monospecific flora) and G. oceanica is usually rare.
G. oceanica, a warm-water species with a maximum temperature range of
between 12 and 30°C (Okada and Mcintyre 1979), is not expected to be a
common component of high latitude shelf assemblages. The species is replaced
in temperate waters by G. caribbeanica, a cold-water preferring Gephyrocapsa
species. However, in shelf regions of the North Atlantic, G. caribbeanica tends
to be rare and is certainly not so adaptive as G. oceanica to the near-shore
environment. The rarity and absence of E. huxleyi in the Asian marginal seas
occurs in waters which are well within the temperature and salinity tolerance of
the species, and perhaps the distribution pattern of the species is better ex-
plained by reference to changes in water mass fertility. Winter (l982b) cor-
related the percentage abundance of E. huxleyi and G. oceanica in bottom
sediments along a traverse from the Gulf of Eden-Red Sea to the Gulf of Aqaba,
with changes in the nutrient characteristics in the overlying watermass. G.
oceanica was suggested to favour high fertility waters and E. huxleyi low fertility
waters. Studies on cores from the Gulf of Aqaba indicated that during inter-
glacial periods, particularly near rnajor Wadi systems, when the Gulfhad a higher
fertility caused by nutrient supply from the land, G. oceanica was found in
highest percentages. A preference for higher nutrient levels may also explain the
E. huxleyilG. oceanica dominance reversal recorded towards land from several
marginal seas.
The dominance of E. huxleyi and G. oceanica in Atlantic and Indo-Pacific
marginal seas generally reflects their abundance patterns found in oceanic
sediments. E. huxleyi is more abundant in the Atlantic than in the Pacific, and
this difference is especially discerniable when comparing equatorial assem-
blages. Schneiderman (1977) noted E. huxleyi to be ubiquitous in the Atlantic
Ocean, the species forming over 50%ofthe flora in tropical regions and increasing
to near 100% in some subpolar sediments. Okada and Mcintyre (1979) recorded
the average annual species composition of coccolithophorid populations from
five weather stations in the North Atlantic, ranging from tropical to subarctic
environments. E. huxleyi had a mean abundance (data compiled from com-
bined surface water and 1OO-m levels) which ranged from 69.7 to 79.5%, whereas
G. oceanica was rarely recorded in the Atlantic survey, and had a maximum
value of6.1% at the tropical locality. In the Pacific Ocean, low-latitude, high
abundances of E. huxleyi (> 40%) are generally restricted to assemblages
underlying waters of low fertility and include the East Pacific Rise, near the
Galapagos and below the Central Pacific gyre at 20 N to 30 N latitude (Roth
0 0
and Coulbourn 1982). G. oceanica is most abundant in assemblages deposited

below the high-fertility equatorial water mass.
The variations in abundance of G. oceanica and E. huxleyi in the Atlantic
and Indo-Pacific regions may be related to major sedimentary facies patterns
and oceanic circulation fractionation. There is a distinct tendency for carbonates
to accumulate in the Atlantic and for siliceous oozes to accumulate in the Indian
252 S.D. Houghton
and Pacific oceans (Sverdrup et al. 1942). These broad differences in

biogenous output are related to the fractionation of both silica and carbonate
between the ocean basins as controlled by deep-circulation patterns. The North
Atlantic is dominated by an anti-estuarine, nutrient-poor circulation, the North
Pacific by a nutrient-rich, estuarine circulation (Redfield et al. 1963). These
difference in fertility are related to the nutrient elements being concentrated in
the direction of major circulation of deep-water from the North Atlantic
through the South Atlantic, Indian and South Pacific to the North Pacific. The
dominance of E. huxleyi at all latitudes in the North Atlantic suggests that the
nutrient-deficient anti-estuarine circulation is the optimum oceanic environ-
ment for the species.
The dominance of G. oceanica in the low-latitude marginal seas of the
Indo-Pacific occurs in a region of strong seasonal variation in phytoplankton
crop, related to monsoonal influences. These changes are most pronounced in
the northwest Indian Ocean during the (winter) southwest monsoon (roughly
May-August) when there is strong upwelling and nutrient-enrichment in the
surface waters (McGill 1973). The richness in nutrient elements during the
monsoon upwelling regions, contrasts markedly with the general paucity of
surface nutrients over wide areas of the open tropical and subtropical oceans.
Monsoonal-influenced upwelling occurs off Somalia, in the northern Arabian
Sea, Bay of Bengal, off the Adaman Islands, off northern Australia and in the
marginal seas of southeast Asia, all regions where the nannofossil assemblages
contain dominant G. oceanica. The high nutrient-preference of G. oceanica
explains its dominance pattern in monsoonal-influenced regions and also
explains the change to E. huxleyi-dominated assemblages in the subtropical
regions of the southern Great Barrier Reef, Tasman Sea and southern coast of
Australia where there is no monsoonal influence. Changes in the ratio of E.
huxleyilG. oceanica in low to mid-latitude assemblages may therefore be used
as an index of water-mass fertility in both marginal and oceanic environments.
These ecological controls on the abundance of E. huxleyi and G. oceanica,
however, may prove to be a hinderance in the resolution of Late Pleistocene
nannofossil biozonation (see Sect. 7).
Okada and Honjo (1975) also reported the common occurrence of mal-
formed coccoliths in the plankton ofthe Asian marginal seas. Such coccoliths are
susceptible to dissolution and have rarely been reported in bottom sediments
from the same area; malformed coccoliths are therefore unlikely to be of
palaeoecological significance. Additional studies on other shelf and marginal
sea areas, particularly from the western Indian Ocean and South Atlantic, may
further help to identify those features which are characteristic of such
environments.
Coccoliths are also found in Recent estuarine and tidal flat sediments.
Coccolith assemblages are transported into such environments as suspended silt
on flood-tides and are comparable in species-composition with those assem-
blages found in offshore sediments (Houghton 1986, 1988). Such processes were
pro ba bly common place during interglacials when sea -level standings were high
and will allow interglacial sediments to be integrated into the biostratigraphic
framework established for oceanic sediments. The Pleistocene marker species E.

huxleyi, Gephyrocapsa oceanica, and Pseudoemiliania lacunosa have all been
recorded in various interglacial deposits of northwest Europe (Houghton,
unpubl. work). Coccoliths therefore have good potential to form a biostrati-
graphic link between the oceanic regime and the continental record.
9.3 Calcareous Nannofossils as Palaeotemperature Indicators
McIntyre and Be (1967) rna pped the occurrence of the more common coccolith
species in both plankton and bottom sediment samples from the Atlantic Ocean.
From this study the coccolith species were divided into five latitudinal climatic
assemblages: tropical, subtropical, transitional, subarctic and subantarctic (Fig.
16). The diagnostic species of the climatic assemblages are listed in Table 2,
although all samples containing coccoliths were dominated by the species
Emiliania huxleyi and Gephyrocapsa species. Tropical and subtropical as-
semblages usually contain three times more species than subpolar assemblages,
confirming the strong latitude control on species diversity.
Biogeogra phic ranges of coccoliths determined from bottom sediment and
plankton samples show that living species have a slightly wider distribution than
those preserved in the sediment. This is attributed to the rapid warming of the
Atlantic during post glacial times. However, post mortem dispersion ofcoccolith
species beyond their living biogeographic range by ocean currents is negligible.
The present-day latitudinal control on coccolith distribution indicates that fossil
Fig. 16. The coccolithophorid climatic zones of

Atlantic Ocean sediments. 1Tropical; 11 sub·
tropical; 1I1 transitional ; IV subarctic, subantarc·
tic (after Mcintyre and Be 1967)
254 S.D. Houghton
Table 2. Diagnostic Recent coccolith species in Atlantic climatic sediment assemblages
Tropical Subtropical
Umbellosphaera irregularis Umbilicosphaera sibogae
Neosphaera coccolithomorpha Rhabdosphaera clavigera
Oolithotus fragilis Calsiosolenia murrayi
Umbellosphaera tenuis Syracosphaera pulchra
Discosphaera tubifera Umbellosphaera irregularis
Rhabdosphaera stylifera Umbellosphaera tenuis
Transitional Subpolar
Calcidiscus leptoporus. type C Coccolith us pelagicuS'
Polar
Barren of coccoliths
aSpecies found today in North Atlantic but absent from South Atlantic.
After McIntyre and Be (1967) and Ruddiman and McIntyre (1976).
coccoliths should be excellent tools for the elucidation of past climatic and
water-mass fluctuations.
Laboratory studies on cultures of E. huxleyi have shown that the species
synthesizes long (C 37 -C 39 ) straight-chain. unsaturated ketones (Volkman et al.
1980). The degree of unsaturation of the ketones is controlled by variations in
the environmental temperature of the algal culture (Dias and Bradshaw 1988).
Identical compounds have also been identified from marine sediments which
range from Recent to Miocene in age. The distinctive chemical structure of the
ketones and their resistance to biological degradation suggests that the occur-
rence of these compounds in marine sediments may be used as a tool to
reconstruct palaeoenvironments. Additional studies are needed to determine
whether such ketones are synthesized by other algal groups and to identify the
extent of the ketones' stratigraphical distribution. If these ketones are found to
be uniquely synthesized by coccolithophorids, then it may also be possible to
recognize coccolithophorid input into sediments even in areas of strong dis-
solution, such as below the carbonate compensation depth (Volkman et al.
1980).
McIntyre et al. (1972) studied Pleistocene nannofossil and planktonic
foraminiferal assemblages in sediments from the North Atlantic to delineate the
intensity and position of polar-front movements. Pleistocene polar, subpolar,
transitional and subtropical water masses were identified and latitudinal shifts
of these assemblages southwards were correlated with polar front movements.
During the last 225000 years six southward polar front migrations were
delineated with up to 10° latitude climatic shift being recorded from the open
ocean. Latitudinally related coccolith assemblages have also been described
from the Cretaceous (Thierstein 1976), and Palaeogene (Haq and Lohmann
1976).
D iscoaster species are thought to have great potential as palaeotemperature
indicators. From their first appearance in the Late Palaeocene until their
extinction in the Late Pliocene, Discoaster species have showed an ecological
preference for tropical and subtropical environments, i.e. for warm water
masses. Bukry (1973) has used the DiscoasterlChiasamolithus ratio as an
indicator of palaeotemperature. Early Eocene sediments at latitude 14 S 0
yielded 76% warm-water Discoaster and 24% Chiasmolithus, whereas higher

latitude assemblages (37 S) contain 46% Discoaster and 54% Chiasmolithus.
0
Oxygen isotope variations, microfossil data, and cyclic variations in the

carbonate content of sediments have been used to demonstrate that the Ceno-
zoic palaeoclimate varied at periodicities correia table to the orbital parameters
(e.g. Hays et al. 1976; Pisias and Leinen 1984). These data suggest that the
Earth's surface temperature varies in response to periodic variations in the
Earth's orbit and axis. Over ca. 100 Kyr the shape of the Earth's orbit is known
to oscillate from a circular path to an elliptical one (eccentricity period). A
second variable results from tilting of the Earth's axis from ca. 21.5 to 24.5 and
0 0
back over a period of ca. 42 Kyr (obliquity period). A third variable is the
precession of the equinoxes which results from the 'wobble' of the Earth's axis,
and has a period of ca. 21 Kyr. All these variables affect the amount of solar
radiation reaching the Earth's surface, and patterns of change can be calculated
from astronomical data.
Backman and his co-workers have initiated detailed quantitative studies to
determine the usefulness of the genus Discoaster as an indicator of sea-surface
palaeotemperature change. These studies are based on a quantitative analysis
of Discoaster abundance from closely spaced samples in deep sea cores. The
abundance values obtained have been shown to be proportional to the ac-
cumulation rate of the species in question (Backman and Shackleton 1983).
Backman and Pestiaux (1986) have determined the variations in the total
abundance of Discoaster species for the Late Pliocene time interval between 1.9
and 3.6 Ma (Ocean Drilling Programme Site 606, latitude 37 N, longitude
0
36 W).
0
Two patterns of accumulation are revealed in the time-dependent variation

in Discoaster accumulation (Fig. 17):
1. A long-term trend of decreasing total Discoaster abundance with
decreasing age;
2. An overprint of short-term oscillations.
Spectral analysis of the Discoaster abundance plots was carried out to test
if palaeoclimatic forcing of all the main orbital elements (eccentricity, obliquity,
precession) is imprinted on the Discoaster record. This study showed that the
413-Kyr eccentricity period, the 41-Kyr obliquity period and the precessional
periods could be identified, although the eccentricity period was the dominant
cycle. The long-term trend of decreasing accumulation of Discoaster specimens
was interpreted by Backman and Pestiaux (1986) to indicate a progressive
decrease in sea-surface temperature in the Late Pliocene. A similar investigation
was carried out by Backman et al. (1986) from Deep Sea Drilling Programme
Hole 522A (56 N, 23 W) cored on the Rockall Plateau. From this higher
0 0
latitude site a similar long-term trend of decreasing D iscoaster abundance with

decreasing age overprinted by short-term abundance oscillations was identified.
256 S.D. Houghton
2329 I
200 0
'i
>.
.>t!
E
o
N )( 1500
Ie
E
c
.2
iii
~ 1000
::I
o
'.."
o
G>
Ui
'oo" 500
o'"
0L-~~~~~~2~.~
5------~3~.0~------~3~
. 5~~
Age(Ma) at DSDP Sit e 606
Fig. 17. Plots of total Discoaster accumulation versus time in Upper Pliocene sediments from Deep
Sea Drilling Project Site 606. Notice the trend of decreasing accumulation with decreasing age and
the overprint of short-term oscillations (after Backman a nd Pestiaux 1986)
Spectral analysis ofthe abundance oscillations for the preglacial period revealed
a dominant quasi periodicity associated with obliquity-induced temperature
variations in the surface waters. Spectral peaks corresponding to the 100 Kyr
eccentricity and 21 Kyr precession periodicites were also detected.
The interpretation that the long-term trend of declining Discoaster abun-
dance throughout the Late Pliocene is a reflection of a prolonged cooling of
northern subtropical waters in the North Atlantic may be treated with some
caution . Since the Middle Miocene, the Discoaster group had been undergoing
a long-term decline in importance which continued until they eventually
became extinct in the latest Pliocene (Fig. 18). The Late Pliocene decline of
Discoaster species recorded from sites 522A and 606 could merely be the 'tail' in
the final decline of the group. With more Discoaster species becoming extinct in
the Pliocene (up to 5.5 extinctions / m.y.) than new species evolving (max. 2.5 first
occurrences / m.y.), the extinction of the group seemed inevitable, regardless of
temperature change. The decline in Discoaster importance to almost negligible
values in the plankton communities of the North Atlantic just prior to their
extinction is reflected in Discoaster/coccolith ratios of the order of 1/10 000 to
1/ 5000 (Backman 1984).
..
...
.....
,... il• ;;
il•
CI
•
"ti:; •e :::• •e
U) 0) .!! ~ ".
<Ii c'" l:
;
~
::;; ".
() 0'-
~
..
0 .::.S e -ti
Co lilt: ". :! ..co CI
I •
& • ••'"' :! g
CD
W ~
<
CI
~
CIII
~~ Q
l:
Q Qci Q ci ci
Early
1.5 Pleistocene Discoaster Species Absent
(--------119
~.o-
Late
18
:::rz::
Ii
~.5 Pliocene I
I
Fig. 18. Distribution of Discoaster I
I
species in the Late Pliocene (compiled 3.0
16 I I
from Backman and Shackleton 1983; I
I
Perch·Nielsen 1985b; Backman and
Pestiaux 1986)
I
I j I
I
Backman and Pestiaux (1986) have argued that Early Pliocene surface
water temperatures were considerably higher than than in Late Pliocene. The
basis of this hypothesis was the greater (x4) abundance of Discoaster species
and total specimens in the Early Pliocene than in the Late Pliocene interval.
However, this difference may merely reflect the general decline in dominance of
Discoaster species in the plankton of tropical and subtropical waters.
Confirmation of higher surface water temperatures in the Early Pliocene may be
provided by a similar decline in importance of other warm-water-preferring
nannofossil groups (e.g. ceratoliths) throughout the Pliocene. Alternatively,
cold-water forms should show an increase in dominance over the time interval.
Shoaling of the Central American seaway between 4.5 and 3 Ma is thought to
have intensified the surface water circulation of the North Atlantic subtropical
gyre system (Backman and Pestiaux 1986). However, the introduction of warm
water into the North Atlantic did not appreciably alter the overall decline in
Discoaster abundance.
If temperature variation is assumed to be the main control on Discoaster
decline and extinctions, it might be expected that Discoaster extinction datums
would not be synchronous, but would vary with latitude with species dying out
in cooler northern waters as their niche contracted, the remaining forms
surviving in the south. Most of the available evidence in the North Atlantic
suggests this is not the case. The final extinction datum of the Discoaster group
has been dated at l.89 Ma for both high- and low-latitude sites. Bukry (1978)
reported that within the Discoaster group, different Discoaster species had
different temperature optima, with D. surculus Martini and Bramlette
dominating in cooler areas. If a decline in temperature was the main cause of
Discoaster extinctions and decline, then D. surculus (Fig. 7e), best adapted to
cooler waters, might have been expected to be the last survivor of the group as
258 S.D. Houghton
palaeotemperature deteriorated in the Late Pliocene. However, D. surculus

became extinct at ca. 2.42-2.46 Ma (Backman and Pestiaux 1986) and therefore
appears not to have survived the first major phase of Northern Hemisphere
Glaciation (based on isotopic values and on the first evidence of ice-rafted
debris) dated at 2.55-2.44 Ma (Shackleton et al. 1984). This extinction datum
suggests that effects other than temperature were responsible for the removal of
the species from the plankton. Ecological factors, such as competition for
nutrients from other warm-water plankton groups or perhaps increased
predation of the Discoaster-bearing algae, probably influenced the decline of
the group.
Backman's pioneer orbital forcing studies on Discoaster abundance have so
far been restricted to the North Atlantic. Similar studies should also be carried
out in the Pacific and Indian Oceans to see how orbital forcing varies with
different palaeoceanographic regimes and latitudes. Orbital forcing studies
from Pleistocene examples will have to be determined from other temper-
ature-sensitive nannofossil groups.
9.4 Oxygen Isotopes in Nannofossils
The estimation of palaeotemperature by use of oxygen-isotope analysis of

calcitic microfossil skeletons has provided a detailed insight into how the Earth's
climate has varied. Such studies document ratios of oxygen isotopes as a relative
deviation (818 0 per mil) from the mean ratio of a standard:
8180 = 1000 [ (180/160) sample ] -1 .
(180/ 16 0) standard
The usual reference standard is PD B-1, which represents the rostrum of a
Cretaceous belemnite from the Pee Dee Formation, South Carolina. The
variation in isotopic composition of the ocean waters over time is documented
by the changing ratio 0[180 ;t60 in fossil carbonates found in oceanic sediments.
Foraminifera, both planktonic and benthic forms, have been commonly used
for such studies and have yielded information on the oceanographic regimes on
the seafloor and in the upper water column. Isotopic analysis of coccolith
carbonate is also a potential tool in the elucidation of palaeo-oceanographic
change in the world's oceans. Coccoliths are used for isotopic analysis because
they have several advantages over foraminifera. Recent coccolithophorids grow
and secrete calcite only in the photic zone and therefore their coccoliths should
incorporate an isotopic signal characteristic of the upper part of the water
column (ca. 0-200 m depth). Foraminifera, however, not only secrete calcite
from waters of specific density and at a depth corresponding to the chlorophyll
maximum in the water column but also may continue secreting calcite during
gametogenesis far below the euphotic zone. Secondly, coccolith species have a
wide biogeographic distribution, being found preserved in all but polar
sediments. Thirdly, coccoliths have a lower magnesium content and larger
individual crystallites in their skeletons than foraminifera and are consequently
less susceptible to dissolution and recrystallization. Coccoliths are also more

abundant in many Tertiary and Cretaceous pelagic carbonates. Despite these
advantageous properties, coccoliths, until relatively recently, have not been
widely used in isotope studies. Unfortunately, beca use of the small size of
nannofossils it is thought to be impossible to isolate individual species of
coccolith from sediments. Isotope analysis of coccolith carbonate is therefore
usually carried out on fine fractions of sediments (ca. < 25 ftM) which contain
polyspecific assemblages.
The oxygen isotope composition of coccolith carbonate from a Quaternary
core in the Caribbean (Anderson and Steinmetz 1981) follows the same pattern
of glacial! interglacial oscillations characteristic of those recorded in planktonic
foraminifera (Fig. 19), including the asymmetrical sawtoothed pattern in-
dicative of rapid deglaciation events. However, although the isotopic variations
in coccolith and foraminifera carbonate are generally synchronous the nan-
nofossil I) 18 0 values are typically offset from planktonic foraminifera values by
up to + 2 per mil during glacial cycles and ca. + 1 per mil during interglacials.
Other studies have produced contrasting results, with the oxygen isotope signal
of coccoliths isotopically lighter relative to equilibrium values (Anderson and
Cole 1975; Dudley 1976).
Dudley et al. (1986) investigated oxygen isotope fractionation in Recent
coccolithophorids grown in culture. This study showed two distinct groups of
non-equilibrium but temperature-controlled fractionation in coccolith car-
bonate precipitation: l80-rich coccoliths including the species Emiliania huxleyi
and Gephyrocapsa oceanica (+ 1 per mil relative to equilibrium calcite) and
l80-depleted coccoliths (-2 to -3 per mil relative to equilibrium calcite) in-
- 2 .0 - 2 .0
.~ G.sacculi fer (Emili an i, 197 2)
\ A\'· . ./J\ :/. . . .\\ f\/V'/\(\.'~

"1'" ~\"- '\:
- 1.0 - 1.0
:. /,,\.\.Jv'/\
m
o
Il.
(5 1.0
.' 'N
,,<,.,,1 \..' .....

• • •"
"~M
\' / ,
;
\
\
.....
0
+ 1.0
co , ..
c.o 2 .0
. t t m + 2.0
7 9 " 11 13 "'"
3-25}<
frac ti on
3 .0 + 3 .0
B A
I I I
o 400 600 800 10'00 1200
Depth in core (em)
Fig. 19. Oxygen isotopic results on the 3-25 /Lm fraction of core P6304- 4. Emiliani's oxygen isotope
data on the planktonic foraminifera species G. sacculifera is also shown. The numbers beneath the
3- 25 /Lm '" 0 curve identify the interglacial isotopic stages. A , B, C indicate three globally
synchronous Late Quaternary coccolith datum levels: A Exti nction of Pseudoemiliania lacunosa
(0.458 Ma); B first appearance of Emiliania huxleyi (0.268 Ma); C reversal in dominance of G.
oceanica and E. huxleyi (0.085 Mal from Anderson and Steinmetz (1981)
260 S.D. Houghton
cluding the species Calcidiscus leptoporus, Umbilicosphaera sibogae, U. hul-

burtiana Gaarder and Syracosphaera pulchra. These data imply that the di-
rection and magnitude of deviation of the isotopic record of coccoliths relative
to foraminifera may be due to variation in species composition, rather than
preservational characteristics of the coccolith assemblages. Biological frac-
tionation ("vital effect") is therefore the most likely cause of resulting non-
equilibrium values.
Although there are limitations to the isotope analysis of coccoliths, it
remains an excellent tool for the study of Late Cenozoic climatic change. As well
as providing an indirect indicator of palaeotemperature oscillations, the tech-
niq ue also provides a record ofthe glacial! interglacial cycles, of changing global
ice volumes, and of glacio-eustatic variations of sea level. The identification of
the isotopic stages in deep-sea cores serves as an additional stratigraphic
framework which can be integrated with the palaeomagnetic and biostrati-
graphic records. Much work still needs to be done in this comparatively new
field; the mechanisms that control isotopic fractionation in coccolithophorids
are still poorly understood. Future studies should also concentrate on the extent
to which the l) l3C in nannofossils reflect the l) 13C in the surface ocean waters.
Such studies may throw light on enhanced phases oflocal upwelling 0[12C and
nutrient-enriched waters. Fractionation of polyspecific assemblages may also
improve the quality of isotopic nannofossil results. Separation of individual
species or genera could be facilitated if detailed studies were carried out on the
settling properties of nannofossils specimens. Discoaster species, because of
their stellate shape and generally large size (ca. > 10 /Lm) should have different
settling properties from coccoliths thus allowing fractionation as a monogeneric
assemblage.
10 Conclusions
Calcareous nannofossils, since their emergence in the latest Triassic, have been
important members of the phytoplankton and significant contributors to the
carbonate budget of pelagic sediments. Their rapid mode of sedimentation in
faecal pellets and other macroaggregates and their solution resistant skeletons,
ensure that there is usually a good correspondence between fossil and living
assemblages. Calcareous nannofossils are good biostratigraphical markers and
excellent tools in the elucidation of palaeoclimatic and palaeoceanographic
change.
Acknowledgements. K. Perch-Nielsen, D.G. Jenkins, J. Jenkins, L. Harper, and N. Swinburne are

acknowledged for their critical comments on an early draft of the manuscript. O. Varol is thanked
for valuable discussions on Cenozoic nannofossil biostratigraphy. The figures were drafted by A.
Bach-Lloyd and J. Taylor. P. Bown is thanked for supplying the SEM micrographs for Fig. 8.
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Chapter 14
Fossil Calcareous Dinoflagellate Cysts
H. KEUPpl
Abstract
Present knowledge of fossil calcareous dinoflagellate cysts is summarized briefly. All genera of the
cyst family Calciodinellaceae Deflandre 1947, including most of the Mesozoic "Calcisphaerulidae"
Bonet 1956, known from the fossil record since the late Triassic, are listed here and subdivided into
three subfamilies.
Several specimens show a more or less complete para tabulation which is expressed by different
modes. A monophyletic character of the traditional peridinian family can be proposed owing to a
corresponding tabulation pattern (ortho-hexa-tabulation of the general formula 4',3a,7",
4-6c,5"',2""). A more or less strong control over cyst morphology by environmental factors is pos-
tulated due to the quasi-extracellular mineralization. This is obviously problematic for sys-
tematics. But on the other hand, these mineralization features can be used for environmental and
stratigraphic interpretations.
Two new names (Pentadinellum, Wallidinellum) are introduced here for the calcareous cysts
of Recent Ensiculi[era cf. mexicana and Scrippsiella sweenyae.
1 Calcareous Cysts vs Thecae
The study of fossil dinoflagellates is the domain of palynology due to their

acid-insoluble, sporopollenin cysts. In general, the motile stages of the Pe-
ridiniales with cellulose thecae are not preserved. Since the work of Deflandre
(1947, 1948), calcareous cysts (which he had described as thecae) have been
known from Mesozoic and Cenozoic sedimentary rocks. Their sizes range from
10 to 100 pm (mostly about 40 /Lm). Despite their global abundance, especially
of the Pithonella-like calcispheres during the Cretaceous and Jurassic
(Beschoren 1926; Wanner 1940; Locker 1967; Keupp 1981,a.o.},ourknowledge
of this fossil group is still very poor. Two main reasons are responsible for this:
l. Most micropaleontologists isolate their material using sieves (63 /Lm),
washing out most of the calcareous cysts. And, in addition, the investigators
of calcareous nannoplankton deal with the fraction smaller than 20 /Lm;
therefore, the middle size range is missed.
2. The sample preparation techniques ofpalynologists involve the use of acids
which destroy calcareous dinoflagellates.
Two groups of Recent dinophyceans containing calcareous walls can be
distinguished:
1 Institute of Palaeontology, Freie Universitiit, Schwendener Str. 8, D-IOOO Berlin 33, Germany
268 H. Keupp
1. Order Peridiniales Haeckel 1894

Family Calciodinellaceae Deflandre 1947 emend. Bujak and Davies
1983: calcareous cysts only
2. Order Thoracosphaerales Tangen 1982: coccoid vegetative stages
The first group, with many genera and species, is reported from sedimentary
rocks from Late Triassic onwards, especially from the Cretaceous. Only one
species representing the order Thoracosphaerales, Thoracosphaera heimii
(Lohmann 1920), is known from the fossil record since the Palaeocene. The
calcareous shell of Thoracosphaera does not show the tabulation patterns
characteristic for most motile stages of dinophyceans. Therefore, the separation
of thoracospheres (including some fossil and Recent species of, as yet,
unconfirmed affinity) from resting cysts ofthe Calciodinellaceae, which often do
not have signs of the thecate tabulation patterns (= para tabulations), is still a
problem in fossil and Recent material. Both calcareous shells sometimes show
more or less complete morphological conformity (Th. heimii with some Ob-
liquipithonelloideans). According to Tangen et al. 1982, the coccoid vegetative
stages of Thoracosphaera have evolved from resting cysts.
This may have happened in connection with the migration of calcareous
dinoflagellates from more neritic to pelagic paleoenvironments (see below).
Before the culturing of isolated clones of Thoracosphaera heimii was achieved,
the differentiation of both Thoracosphaera and Pithonella-like calcispheres was
determined according to size: about 40-100 /Lm for the pithonelloids and 7-24
/Lm for the thoracospheres (Bolli 1974; Keupp 1979a). However, later findings
involving small calcareous cysts show that this criterion is useless in the
differentiation offossil calcareous bodies (Keupp 1981). Many calcareous cysts
are probably included among the numerous Recent and fossil species described
as Thoracosphaera (e.g. Stradner 1961; Kamptner 1963,1967; Bramlette and
Martini 1964; Fiitterer 1976,1977; Keupp 1978, 1979a; Jafar 1979; Gilbert and
Clark 1983). All species of these "thoracospheres", which represent calcareous
cysts, should be placed in the cyst genera Obliquipithonella Keupp 1984 and
Orthopithonella Keupp 1984.
2 Taxonomic Aspects
The taxonomy of dinoflagellates used currently is ambivalent. The classificatory

system of Recent dinoflagellates is based on motile stage morphology, while that
of fossil specimens is based on cyst morphology. In peridinians the mor-
phological development of both thecae and cysts (with organic as well as
calcareous walls) seems in many cases to be independent (Sarjeant 1974,
pp. 86-87; Tappan 1980). For example, the modern motile stages of Scrippsiella
sweenyae (Balech 1959) and Scrippsiella trochoidea (Stein 1983) show a very
close affinity owing to the corresponding tabulation pattern of their thecae. But,
using cyst taxonomy. their calcareous cysts (Wallidinellum n.g. resp. Rhab-
dothorax Kamptner 1958) belong to different subfamilies. Due to practical
Fossil Calcareous Dinoflagellate Cysts 269
reasons, palynologists use separate classifications for thecae (often with a

conservative morphology) and cysts (using more advanced development as a
criterion).
All calcareous genera of certain and probable dinoflagellate cysts from Late
Triassic to Recent are listed in Table I showing their stratigraphic ranges. Three
subfamilies of the Calciodinellaceae can be distinguished owing to their
different wall structures (Keupp 1987). Their definition is based on the crys-
tallographic orientation of calcite crystals forming the calcareous wall (or at least
its outer part; Figs. 9-12):
l. Perpendicular to cyst surface: Orthopi thonelloideae
2. Irregularly oblique to tangential: Obliquipithonelloideae
3. Uniformly oblique: Pithonelloideae (only fossil)
Some cyst genera show more or less clear paratabulations which reflect, from the
motile theca, a complete or a reduced "orthohexa-tabulation" of the general
formula 4', 3a, 7", 4-6c, 5111 ,2"". In this way the theca is built by different cycles
of single plates: three ofthe four apical (') and three intercalary plates (3a) form
the apical area round the centrally positioned plate 3'. The ventral sulcus
(position of the longitudinal flagellum) interrupts the circular cycles as a small
area between plate 3' and the antapex. It includes the hexagonal plate I' which
looks symmetrical due to the onlap of six other plates (= "ortho-hexa"). After
the cycle of seven precingular plates ("), the girdle (circular furrow of the
transversal flagellum), built by 4 to 6 small plates, follows. The hypocyst below
the girdle is built by a cycle of 5 postcingular plates (III). The antapex shows two
large pentagonal plates intersected ventrally by the end of the sulcus. Mech-
anisms which are responsible for the replication of this thecate plate pattern
on the cysts are still unknown. This described plate pattern represents a quite
conservative tabulation of the Peridiniineae (Bujak and Davies 1983). Often a
reduced number of paraplates can be observed which results from fusion of
single plate equivalents. This decrease of plate number characterizes only the
cysts and does not occur in the motile stages. The realization of para tabulations
varies (for terminology see Sarjeant 1982):
l. Holotabulate: sutures, crests (e.g. Calciodinellum Deflandre 1947; Fig. 2),
or edges of the cyst (e.g. Heptasphaera Keupp 1979a, Fig. 3) correspond to
the thecate tabulation;
2. Intratabulate: processes arise from centre of thecate plates (e.g. A lasphaera
Keupp 1979c; Fig. 6);
3. Cingotabulate: expressed only by indications of the cingulum and ventral
sulcus (e.g. Carinellum Keupp 1981; Figs. 4-5);
4. Cryptotabulate: only by shape and position of the archeopyle (e.g. partim
Orthopithonella; Fig. 7).
A high variability in cyst shape results from both specific paraplate reductions
and from the different modes of para tabulations. In addition, the localization of
plate-equivalent structures also shows a wide range in the cysts:
Table 1. Genera of Calciodinellaceae and their stratigraphic ranges' N
Cl
Cyst genus Stratigraphic distribution Theda
Suhfamily Orthopithonelloideae Keupp 1987:

CalclOdinellum Dellandre 1947 Miocene-Recent &
Calcigonellum Dellandre 1948 Miocene-Quaternary
Praeealcigonellum Keupp & Versteegh 1989 Ha uterivian-Quaternary
Calcipterellum Dellandre 1948 Miocene
Eehinodinella Keupp 1980b Hauterivian-Albian
Keuppisphaera (Keupp 1987) Lentin & Williams '89 Late Albian
Orthocarinellum Keupp 1987 Late Albian
Orthopithonella Keupp 1984 ? Late Triassic, Jurassic-Recent
Sphaerodinella Keupp & Versteegh 1989 Miocene-Recent
Rhabdothorax Kamptner 1958 Albian-Recent Scrippsiella trochoidea (Stein)
"tetrahedral cyst type" Wall & Dale '68 Recent ?
Suhfilmily incertas sedis:
Dimorphosphaera Kcupp 1979a Barremian-Albian
Subfamily Obliquipithonelloideae Keupp 1987:
Ohliquipithonella Keupp 1984 o Late Triassic, Jurassic-Recent Scrippsiella patagonica Akselman & Keupp
incl. "spherical cyst" Wall & Dale 1968 Ensiculifera "rex" Dale & Tangen (partim)
Alasphaera Keupp 1979c Valanginian/Hauterivian
Biearinellum (Dell. 1948) Keupp 1984 Oxfordian-Eocene
o inel. "bicarinate cyst" Wall & Dale Recent
Calciearpinum Dellandrc 1948 Eocene
Carinellum Keupp 1981 Late Hauterivian-Eocene
Cubodinellum Keupp 1987 Albian-early Cenomanian
Gonellum Keupp 1987 Albian
Heptasphaera Keupp 1977a Late Jurassic-early Barremian
Nephrodinella Keupp 1981 Late Hauterivian-Barremian
Pcntadinellum n.g. Recent Ensiculifera cf. mexicana Balech ;z:
Sliteria Krasheninnikov & Basov 1983 Turonian-early Campanian
Tetramerosphaera Willems 1985 Maastrichtian ...~
C
"0
"0
Pleistocene-Recent Scrippsiella sweenyae Balech ."
Wallidinellum n.g.
~
"clavate cyst type" Wall & Dale '68 Recent ? ~
"keeled cyst type" Wall & Dale '68 Quaternary (fossil) n
"pyriform cyst type" Wall et a!. '73 Holocene ? n
'"
"wedged cyst type" Wall & Dale '68 Quaternary (fossil) ~
'"o
Subfamily Pithonelloideae Keupp 1987 ~
Pithonella (Lorenz 1901), Villain '77 Late Barremian-Maastrichtian tJ
5'
inc!. "Palinosphaera Borza 1972" o
? Bonetocardiella Dufour 1968 Albian-Danian (JO
""'"
Wallia Keupp (1990) Campanian ~
0;-
? Centosphaera Wise & Wind 1976 Maastrichtian ro
n
aThe taxa of motile stages of Recent calcareous cysts are given where possible.
~
tv
-.]
272 H. Keupp
-....
..........
""-
\
\
\
\vI
I
J
I
/
/
/
Ektocoel //
----
Au'*-nd _
Fig. 1. Obliquipithonella showing the three to four possible organic phragms (I endophragm ; II
mesophragm ; III periphragm; IV ectophragm). Instead of the phragms I (inc!. II) and Ill, an
autophragm can exist. The mineralization of one, two, or in some cases three calcareous wall layers
can occur independently in each interspace of two phragms. (Keupp 1981)
Figs. 2-7. Some different modes of para tabulations developed in calcareous dinoflagellate cysts
(terminology according to Sarjeant 1982)
Fig. 2. Holotabulate paratabulation expressed by surface·crests in Calciodinellum operosum
Deflandre 1947 from late Miocene, EI Medhi, Algeria (original sample material collected by G.
Deflandre). Diameter of cyst: 30 Jtm
Fig. 3. Holotabulate reflection of an ortho-hexa-tabulation expressed by edges of the cyst in
Heptasphaera michaeli Keupp 1979a. Early Barremian of Sarstedt (Moorberg), NW-Germany.
Diameter of cyst : 50 Jtm
Figs. 4-5. Stereopair of Carinellum parasolis Keupp 1984 (view on the epicyst) showing a
cingotabulate paratabulation. Eocene from Caussy/France (sample collected by G. Deflandre).
Dia meter of cysts: 45 Jtm
Fig. 6. The processes built by both inner and outer calcareous wall-layers of Alasphaera caudata
Keupp 1979c reflect intratabularly the orlhoperidinian tabulation . Lower Hauterivian from Sar-
stedt (Moorberg)/NW-Germany. Diameter of cyst: 65 Jtm
Fig. 7. Orthopithonella gusta{sonii (Bolli 1974) with cryptotabulate paratabulation expressed by
polygonale archeopyle (= Orthopithonella ''paratabulala'' Keupp 1980b). Late Aptian from
Mittellandkanal nea r SchwiecheldtlNW-Germany. Diameter of cyst: 42 Jtm
274 H. Keupp
I. Outside the outer calcareous wall layer by crests, tubercles, and sutures: e.g.
Calciodinellum (Fig. 2), Bicarinellum Deflandre 1948 emend. Keupp 1984,
Orthopithonella "paratabulata" Keupp 1980c (Fig. 7);
2. Outside the inner calcareous wall layer only by crests: e.g. Tetrameros-
phaera Willems 1985;
3. Outside both the inner and outer calcareous wall layer: e.g. Alasphaera
(Fig. 6);
4. Inside the organic membranes only (Compare Fig. 1):
periphragm: e.g. Obliquipithonellapatriciagreeleyae(Bolli 1974); Keupp
1980a; (Fig. 8);
endophragm: Obliquipithonella loeblichi (Bolli 1974); Keupp 1981,
PI. 4817;
autophragm: Obliquipithonella organica (Hultberg 1975)
Except for a few specimens representing single species of the genera Ortho-
pithonella (0. "para tabulata "), ObliquipithoneUa (0. patriciagreeleyae, cf.
thayeri, organica), and Echinodinella Keupp 1980b (E. erinacea, Keupp 1980b),
the most spherical to ovoid cysts, the prominent specimens of the so-called
"Mesozoic and Cenozoic calcispheres", do not show any paratabulations. But
their wall structures and apical archeopyles (= openings for excystment) prove
close affinity to true Calciodinellaceae (Figs. 9-13). Corresponding to this,
calcareous cysts of the Recent motile dinoflagellate Ensiculifera sp., as sampled
by Tangen and Dale from plankton of the Oslofjorden in August of 1982 (Figs.
14-17), are congruent morphologically with common Mesozoic cysts (Jurassic,
Cretaceous) of the Obliquipithonella quittyi (Bolli 1974)-type. In contrast, there
is no evidence for a dinoflagellate nature for the "Palaeozoic calcispheres",
Fig. 8. Broken specimen of Obliquipithonella patriciagreeleyae (Bolli 1974) from early Barremian
of Speeton/England showing the ventral area of a complete ortho-hexa-paratabulation inside the
organic periphragm only. Size of fragment (cyst 3878) 25 I'm
Figs. 9-13. The three calciodinellacean subfamilies are characterized by the structure of their (at
least outer) calcareous walls
Fig. 9. Fragment of the single-layered calcareous wall (autophragm inside) of Echinodinella
erinacea Keupp 1980b showing the orthopithonelloid structure with perpendicularly orientated
calcite crystals. Thickness of wall: 6 fJ.m. Early Albian from Altwarmbiichen near Hannover /NW
Germany
Fig.l0. Wall of Obliquipithonella multistrata (Pflaumann and Krasheninnikov 1978). The calcified
pericoel (= inner wall layer) show an irregular microgranular structure, while the calcified ectocoel
(= outer wall layer) has been constructed by irregularly oblique crystal-orientation, the so-called
crossing-bars-structure (Keupp 1979b). Hauterivian/Barremian-boundary from Sarstedt (Gott)/
NW Germany. x2000
Fig. 11. Double-layered calcareous wall of obliquipithonelloid Bicarinellum calvum Keupp 1979a.
The inner wall layer shows a loosely arranged "crossing-bars structure" due to poor calcification.
Early Barremian from Sarstedt (Gott)/NW Germany. X 2500
Figs.12-13. The Wallia (Keupp I 990)from thelate Cretaceous, South Dakota/USA, belongs to the
Pithonelloideae: its wall-crystals show a uniquely oblique orientation. Diameter of cyst: 90 fJ.m,
thickness of wall: 7 fJ.m
276 H. Keupp
Figs. 14-17. The living motile stage Ensiculifera sp. hitherto undescribed , from Oslofjordenl
Norway produces calcareous cysts very similar to Obliquipilhonella quilfyi (Bolli 1974): see Keupp
1981. PI. 34. The photographs were made by Tangen, Trondheim and Dale, Oslo, from plankton
samples collected in August 1982
Fig. 14. Empty hypotheca of the motile stage including sulcus and girdle which show the genus
character. namely a long spine protruding from the girdle (= ensicule)
Fig. 15. Partly collapsed calcareous cyst with rigid surface which is probably caused by immaturity,
since these morphotypes have been found together with motile thecae in the plankton. Diameter of
cysts: 48 iJ.m
Fig. 16-17. Apparently smooth cyst, 43 iJ.m diameter. representing the characteristic Obliquipith.
onella qui/fyi-type. and detail of its surface (X 10000)
which represent a heterogeneous group of algal spores and microproblematica

from a more or less restricted environment.
The archeopyle in general lies apically and includes plate equivalences to
apical or intercalary series or both. Sometimes the number of plates included is
quite variable within one species [see for example Orthopithonella
''paratabulata'' (Keupp 1980c, pI. 31 )]. In Mesozoic cysts, the archeopyles are
mostly small using the apical plate equivalences, often 3' only. In some ortho-
pithonelloideans, a large archeopyle including both apical and intercalary
plates seems to become dominant in the Tertiary (e.g. Calciodinellum, Ortho-
pithonella albatrosiana, Kamptner 1963; Calcipterellum, Deflandre 1948;
Calcigonellum in[ula, Deflandre 1948; see Keupp 1984; Keupp and Versteegh
1989), perhaps in connection with the change of some cysts to oceanic envi-
ronments (see below).
3 The Wall of Calcareous Cysts
The various reaons why dinoflagellates make cysts seasonally are still under
discussion (Sarjeant et ai. 1987).
According to Evitt 1985, the formation offossilizable cysts occur in modern
marine peridinians during sexual reproduction (hypnozygotes) rather than in
reaction to unfavourable environmental conditions (resting cysts). Observa-
tions ofa few non-calcareous species have shown (Pfiester 1975, 1984; Spector
et ai. 1981; Dale 1983) that the process of encystment varies specifically. In
Peridinium cinctum (MUller) Ehrenberg 1932, for example, three separate
organic membranes are constructed step by step, beginning with a progressively
enlarged planozygote covered with a theca of single cellulose plates (this is very
similar to the vegetative motile stage), and ending within the nonmotile hyp-
nozygote. In contrast to most other species, which shed the theca after forming
the cyst, the theca of the planozygote in Peridinium cinctum has been made
resistant by impregnation with chitin. In other species four more or less resistant
wall membranes arise during cyst formation (von Stosch 1973).
In calciodinelloids, th e na ture of the cysts (h ypnozygotes or resting cysts or
both) is still unknown. Cenozoic pelagic species with thin calcareous walls
probably represent hypnozygotes (Dale 1986), while thick walled, more or less
neritic cysts may be chiefly resting cysts. It appears that before the formation of
calcareous cysts two to four organic membranes are in existence (Fig. 1). While
the outermost "ectophragm" is normally not fossilizable, the inner membranes
("autophragm" or instead "periphragm" + "endophragm" - sometimes an
additional "mesophragm" too) are often preserved consisting of acid insoluble
material, probably sporopollenin.
Taxonomic confusion originates from isolated phragms of calciodinelloids
in palynological preparation procedures (H ultberg 1985). For example, the new
organic-walled dinoflagellate cyst Neuffenia, created by Bremer and DUff
(1986) from Oxfordian limestones rich in calcispheres, seems to be such an
isolated phragm. By analogy with organic-walled cysts, it can be assumed that
278 H. Keupp
the interspaces between the poreless membranes are filled with a polysaccharide
mucus. Due to a strong buffer effect from the polysaccharides, a considerable
supersaturation of dissolved calcium carbonate is possible. Similar buffer
solutions prevent the calcification of the mucus covering photosynthetically
active cyanobacterians. But the mineralization of this mucus takes place very
rapidly when the maximum effect of buffering is weakened (S. Golubic, pers.
comm.). In a similar way, calcite crystals may grow to form the wall in many
calciodinelloids. Independent of the primary number of organic membranes,
one to three separate calcareous layers can be found. Different crystalline
structures in each layer suggest their independent mineralization. Despite
crystal growth inside the organic membranes, the mode of mineralization
appears to be extracellular. Therefore, golgiderived vesicles characteristic of
genetically governed mineralizations could not be observed during wall for-
mations of thoracospheres (Tangen et al. 1982). The common formation of
dendritic, sometimes hollow, crystals in calciodinelloids support the interpre-
tation of rapid mineralization from supersaturated solutions. The primary
orientation of crystals seems to be determined by an epitactic overgrowth of
distinct membrane structures or by an organic matrix (specific structure of the
polysaccharides) (Bandel and Keupp 1985). Further crystal growth appears as
an inorganic pattern (Keupp 1981).
The phenotypic development of wall-forming crystals seems to be con-
trolled by environmental factors, such as temperature and salinity. Forexample,
the analysis of sedimentary rocks and fossils from the Late Aptian/Early Albian
ofthe Lower Saxonian Basin (northwestern Germany) allows reconstruction of
a coastal upwelling system (Kemper 1982). Comparison of synchroneous cal-
careous dinoflagellate associations from the central basin and the marginal
upwelling area show clear trends of morphological change within cosmopolitan
cyst species (Keupp 1981, 1982). Particularly the following features changed
with reduced temperature conditions:
1. Progressive reduction of inner calcareous wall layers;
2. Increase in the size of crystals, producing a rough cyst surface;
3. Disappearance of ovoid to elongate cysts.
On the other hand, similar trends have been shown in the vertical calcisphere
succession of the D-Beds (Berriasian to Early Hauterivian) ofSpeeton, England,
which are characterized by contemporaneous freshwater influence during the
Early Valanginian (Keupp and Mutterlose 1984).
Taxonomic problems at genus and species level arise more in the Ob-
liquipithonelloideae than in both ofthe other subfamilies Orthopithonelloideae
and Pithonelloideae. They contain a wide range of primary individual vari-
ability, and, additionally, morphological control by environmental factors.
Often all intermediate morphotypes can be shown in a single sample of cysts.
Environmental influences on cyst morphology are recognizable through
reversible morphological trends in vertical successions of cyst associations
(Keupp 1980b, 1981). The following variable features seem to be influenced by
temperature and salinity:
Fossi.l Calcareous Dinoflagellate Cysts 279
A Obliquipithonelloideae (for explanations see e.g. Keupp 1981, PI 19 20, 0
24-25,37-39,46-50):
Number of mineralized wall layers: normally two separate calcareous walls
exist, but in well-preserved material from the Lower Cretaceous, cysts with only
poorly or noncalcified pericoels occur (Fig. I; Keupp 1981, PI. 20/10-14;
Structure of the inner wall layer, independent of the primary rate of crystal
nucleation: microgranular to "crossing bars structure" (Figs. 10-11; Keupp
1979b);
Size and shape (more or less dendritic) of calcite crystals modifying the cyst
surface: for example, the size of simple rhombohedric crystals of the Ob-
liquipithonella loeblichi group, is not correlated with the diameter of cysts
(Keupp 1981, Fig. 19; PI. 46, 50). Their sizes range from 3 to 14 /Lm (diameter of
cysts about 45 /Lm in the middle) in the Lower Cretaceous, with one sample
containing nearly the same sizes;
Intensity of surface sculptures in connection with para tabulations, which are
independent of the content of calcium carbonate, e.g. the vertical succession of
Bicarinellum calvum Keupp 1979a in the Hauterivian/Barremian of Speeton
(Keupp 1981, pp. 47-50) shifts from elongated strong nodular morphotypes to
smooth, nearly spherical cysts. This may be due to decreasing Tethyan influence
of the Boreal Realm;
Cyst shape of some species (elongated versus spherical: see example above:
Bicarinellum calvum).
B. Orthopithonelloideae:
Size and shape of wall crystals in some species (e.g. groups of Orthopithonella
gustafsonii and O. porata (Keupp 1982): Keupp 1980a, 1981);
Intensity of surface sculptures in connection with paratabulations (e.g.: mor-
photype succession of Praecalcigonellum polymorphum from ssp. den tatum,
with high sutural crests, to ssp. tenue, virtually without sculpture, in the
Aptian/ Albian (Keupp 1980b). This corresponds to a contemporaneous re-
striction of benthonic organisms, dominated by textulariid foraminifera, sug-
gested to be caused by cold water conditions (Michael 1979, pp. 314-316);
Cyst shape of some species analogous to the Obliquipithonelloideae (e.g.
the elongate derivation from Praecalcigonellum mutterlosei (Keupp 1979a),
the morphospecies P. sulcatum (Keupp 1979b), marks, during the Hauterivian/
Barremian, the warmer water influences of the Tethyan: Keupp 1987; Fig. 3).
Number of calcareous walls and structure of inner calcareous wall (only in two
observed cases):
1. Orthopithonella cf. tuberosa (Kamptner 1963) characterized mostly by a
single layered wall, occurs sometimes with an additional small inner
calcareous layer (Keupp 1981, pI. 12);
280 H. Keupp
2. In Oligocene Orthopithone/la sp. (Weiler 1982) two double-walled mor-

photypes occur: (a) with the innerla yer constructed by radial fibrous, partly
hollow crystals (obviously dominant in the Rupelton facies) and (b) with
the inner layer of irregular crystal orientation (dominant in the carbonate
bearing sandy facies).
C Pithonelloideae
Up until now, there are few observations on which to base definitive statements.
4 Ecological Aspects
In the Recent, calcareous dinoflagellate cysts are known from a wide climatic
range of neritic and pelagic marine environments [Wall and Dale (1968a):
latitude 9°_42° N; in the Arctic and Antarctic oceans also: Dale (1983);
Akselman and Keupp (in press)]. The relatively rare neritic cysts prefer tropical
rather than temperate regions (Dale 1975, p 49), while oceanic forms (in the
Recent, they dominate the pelagic dinoflagellate assemblages) including the
thoracospheres, reach into the subarctic regions (Dale 1986). Calcareous cysts
are probably known since the Late Triassic (Orthopithonella and Obliquipi-
thonella: lanofske 1987). They existed during the Mesozoic in both Mediter-
ranean and Boreal Realms. Mesozoic calcispheres have been found
predominantly in sediments of open shelf to slope environments, sometimes of
the inner shelf also (Villain 1981). This palaeogeographic distribution corre-
sponds with the increasing importance of cyst formation in neritic
dinoflagellates in general (Reid 1978).
After the extinction of the Pithonelloideae at the Cretaceous/Tertiary
boundary, a migration of calcareous dinoflagellates into oceanic environments
seems to have occurred. This change presupposes new survival strategies for
pelagic species. Thoracospheres created a coccoid non-motile (planktonic)
vegetative life phase, instead of the flagellate thecate stage present in all other
dinoflagellates (Dale 1986). Some Recent pelagic calciodinellaceans (e.g.
Rhabdothorax, Calciodinellum) favour controlled sinking strategies in their
calcareous cysts: motile stages leave the cyst during its suspension in the water
column before it sinks to a depth from which the emerging cell can not re-enter
the photic zone (Tangen et al. 1982; Dale 1986).
The morphogenetic reactions to some environmental conditions by the
calcareous dinoflagellates listed above, in particular those of the Obliquipi-
thonelloideae, make this fossil group of considerable use for palaeoecological
analysis. The calcisphere assemblages of the Lower Cretaceous moderate
Boreal Realm (NW Germany, North Sea Basin) allow palaeooceanographic
and palaeogeographic interpretations, as well as ecostratigraphic correlations,
of regional value. This should explained by a few paradigms already published:
I. Periodic connections between the Tethyan and Boreal Realms via the
"Polish Street" to the east during the early Hauterivian and the "Paris
Street" to the west during Late Barremian to Aptian enabled the warmer
water faunas and floras to influence the Boreal basins (North Sea Basin,
Lower Saxonian Basin, etc.), which are partly separated by swells and
islands (Michael 1979). Through these migration two modifications oc-
curred within the calcisphere assemblages: (l) new species immigrated; (2)
the more-or-less cosmopolitan cyst species reacted phenotypically. Con-
sidering only the second point, nearly congruent contemporaneous vertical
sequences of temperature governed morphotypes of obliqui- and ortho-
pithonelloideans described above (two- or single-layered calcareous walls,
small or large crystals, ovoid or spherical cyst shapes) within different
sections of the Lower Saxonian Basin as well as in the North Sea Basin
(Keupp 1981, Figs. 22-24; 1987, Fig. 3). Despite the rare occurrence of
locally restricted morphotypes (see following paradigm), the calcispheres
can be used here for stratigraphic correlations. Large-scale ecological
alterations appear to have been responsible for the synchronous
modifications of cyst morphologies.
2. On the other hand, the local distribution patterns of Obliquipithonella
morphotypes in late Aptian sediments confirm the hypothesis of a coastal
upwelling within the Lower Saxonian Basin at this time (see above, Keupp
1982).
3. In the above-mentioned example of vertical calcisphere succession of the
D-Beds from Speeton Clay during a regressive-transgressive cycle (Keupp
and Mutterlose 1984), the morpho types obviously correlate to the different
salinities. The increasing freshwater influence seems to have modified the
cyst morphology in the same way as decreasing temperatures.
5 Phylogenetic Aspects
The phylogeny of the Peridiniineae sensu Bujak and Davies (1983), including
the cyst family Calciodinellaceae, is still obscure. The earliest certain peridi-
niinean tabulations are reported inside the organic phragms of the Lower
Cretaceous calcispheres Obliquipithonella patriciagreeleyae and Obliquipitho-
nella cf. thayeri as described by Keupp (l979c, 1981, PI. 1817). On the other
hand, the number and arrangement of tubercles modifying the cyst surface of
the Oxfordian Bicarinellum jurassicum (Deflandre 1948) also reflect a pe-
ridiniinean pattern (Keupp 1984 p. 20). Therefore, the Peridiniineae can be
traced back at least to the Late Jurassic. With the proviso of a peridiniinean
affinity to non-para tabulated cysts of Late Triassic age, which correspond to the
wall-structure of calcareous dinoflagellate cysts from the Jurassic and Cre-
taceous (Janofske 1987), the roots of the Peridiniineae suborder may reach into
the Triassic. The conservative Calciodinellaceae seem to be a monophyletic
family according to their unique paratabulation and their corresponding ar-
cheopyle position. Probably they represent an ancestral group of the suborder
Peridiniineae.
282 H.Keupp
6 Prospects
Despite some findings concerning morphology, wall structures palaeobiogeo-

graphy, and palaeoecology of calcareous dinoflagellate cysts over the last 20
years, our knowledge of this important group remains poor. The outline of
possibilities and interpretations in calcisphere research given here describe
qualitative trends and feelings rather than objective statements. The paradigms
used here have still to be proved through further investigations in otherlocalities
and stratigraphic positions. Furthermore, we need to understand the biology of
the Recent calciodinelloideans, including their different life cycles, conditions
for the formation of their calcareous shells, and the environmental conditions
controlling their morphology, before objective systematics of fossil forms and
definitive palaeoecological analysis will be possible.
Acknowledgements. I am grateful to Karl Tangen, who sent me photographs and information on the
new species of Recent Ensiculi[era (Figs. 14-17). I am indebted to Barrie Dale for some stimulating
discussions concerning Recent calcareous dinoflagellates and to both Elisabeth Gierlowski-Kor-
desch and Robert Riding for reading the English manuscript.
Appendix
Wall and Dale (1968a) described for the first time some Quaternary calcareous
cysts. These authors named the Recent cysts of Scrippsiella trochoidea and of
Scrippsiella sweenyae according to their thecate stage. All other calcareous cysts
of which the motile stages are not known (except of Calciodinellum operosum),
have been documented in an open descriptive nomenclature. The so called
"oblate cyst type" could later be identified belonging to Ensiculifera cf. mex-
icana Balech 1967 (Wall et ai. 1970). Using a consistent mode of classification
independently for cysts and thecae, which is essential to correlate fossil and
modern dinoflagellates, the cysts have to be named separately.
The more or less spinose calcareous cysts of Scrippsiella trochoidea (see Wall
et ai. 1970) correspond with the genus Rhabdothorax (Gaarder and Heimdal
1973; FUtterer 1977, p. 718; Keupp 1987, p. 44). Kamptner (1937, 1958)
supposed the Recent species R. erinacea and R. nigra to be coccolithophorids
with affinities to Thoracosphaera sensu Kamptner 1928. Rhabdothorax now has
been classified within the subfamily Orthopithonelloideae due to the perpen-
dicular orientation of its calcitic wall crystals. The affinities of subjectively
distinguishable Rhabdothorax morphotypes to one or several motile species are
still uncertain (see Dale 1976, PI. 1/19; 1977).
For the calcareous cysts of Scrippsiella sweenyae described by Wall and
Dale (1968a, pp. 1401, 1403) I here introduce the new obliquipithonelloid genus
name Wallidinellum n.g.:
Name: In honour of David Wall

Diagnosis of the genus: Microgranular calcitic cyst showing an incomplete
ortho-hexa-tabulation expressed by external rounded ridges, dominated by the

precingular and antapical homologues. Typespecies: Wallidinellum dalei n.sp.
Holotype: Single grain preparation C216 (Woods Hole Oceanographic Insti-

tution) from surface sediments of the Western Arabian Area, figured by Wall
and Dale 1968a, PI. 172, Figs. 5-6.
Name: In honour of Barrie Dale
Diagnosis of the species: see genus -diagnosis
Description: For morphologic details see the original description given by Wall
and Dale (l968a). In contrast to the opinion ofthese authors, the seven nodular
plate areas near the equatorial zone seem to reflect the precingular plates rather
than the cingular plates, analogous to other calcareous cysts (e.g. Bicarinellum:
Keupp 1981, Fig. 11; 1987, Fig. 7). The zigzag margin of the operculum result
from a large apical archeopyle including both apical and intercalary plates.
Differential diagnosis: Wallidinellum shows similarities with Heptasphaera,
having a single layered microgranular wall and a more or less tabulate para-
tabulation expressed by edges (especially of the hypocyst) without a clear
paracingulum. But in contrast to the apical flattened Heptasphaera with ~mall
archeopyle (equivalence of3' only) the apical area of Wallidinellum is convex
and possesses a large attached operculum. Sliteria (Krasheninnikov and Basov
1983 is similar to Heptasphaera in cysts shape, para tabulation mode, and the
small archeopyle. It differs from both Heptasphaera and Wallidinellum by its
coarse wall structure.
The "oblate cyst type" of Wall and Dale (l968a) recognized as a calcareous cyst
of Ensiculifera cf. mexicana by Wall et al. (1970) is here given the new
obliquipithonelloid genus name Pentadinellum n.g.:
Name: According to the pentagonal outline in polar view
Diagnosis of genus: A strongly oblate autophragm is surrounded by a bica-
rinate calcareous wall of microgranular structure.
Typespecies: Pentadinellum oblatum n.sp.
Holotype: Single grain preparation C721 (Woods Hole Oceanographic Insti-
tution) from Recent sediments of Bermuda, figured by Wall and Dale (1968a,
PI. 172, Fig. 20).
Name: (lat.) due to the oblate morphology.
Diagnosis of the species: see genus-diagnosis
Description: For details see Wall and Dale (1968a, p. 1406). Both circular keels
around the (?) ventrally tapered cyst are created by pre- and postcingular plate
homologues. The outline of the cyst in polar view is pentagonal, due to the
morphological dominance of the lower (= postcingular) keel. Similar reduc-
tions to a pentagonal cyst-groundplan (in contrast to seven precingulars),
conditional on morphological assimilation of the precingular equivalences on
the postcingulars, has been reported from other cysts (e.g. Praecalcigonellum
284 H. Keupp
polymorphum: Keupp 1980b; Carinellum SSp.: Keupp 1981, 1987; Gonellum

kurti Keupp 1987). In congruence with the genus Bicarinellum, the wide
transversal band seems to reflect the fused homologues of precingular and
cingular plates (Keupp 1987). The number of recognizable septa dividing the
transversal band into 7 poorly defined areas represent the sutures Of precin-
gulars rather than of cingular plates (compare: Calciodinellum operosum:
Keupp 1984, Fig. 8/6; Praecalcigonellumpolymorphum: Keupp 1980b, PI. 1517,
11). The apical archeopyle (operculum) shows a subcircular to polygonal
outline. Weak ridges of the more or less flattened hypocyst probably mark the
ventral sulcus and the two antapical plates.
Differential diagnosis: Among the Obliquipithonelloideae, Pentadinellum
show morphologically close similarities to the genus Bicarinellum, except for the
oblate autophragm. The pentagonal ground plan, the prominent carinae, and
the more or less flattened hypotract especially correspond with Bicarinellum
cristatum Keupp (1982) from sediments of the lowermost Barremian to early
Albian. But its spherical endocoel and lack of an operculum both distinguish it.
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III Algae and Stromatolites Through Time
Chapter 15
Archaean and Proterozoic Stromatolites
S.M. AWRAMIKI
Abstract
The approximately 3400-million-year history of the Archaean and Proterozoic Eons is rich in the
fossil remains of organosedimentary structures called stromatolites, built primarily by cyanobac-
teria. Stromatolites first appear 3500 Ma ago and argue well for the presence and therefore great
antiquity of cyanobacteria. The presence of cyanobacteria in such ancient rocks indicates that the
evolution of the major prokaryotic phyla had occurred by Early Archaean time.
Although rare in the Archaean and first 300 million years of the Proterozoic, stromatolites
undergo diversification and increase in abundance in the late Early Proterozoic due, in large part,
to the oxygenation of the atmosphere-hydrosphere system, permitting cyanobacteria to disperse,
colonize, and thrive in shallow continental shelf-like environments produced during earlier and
contemporaneous periods of cratonization. A second diversification occurred in the Early to Middle
Riphean (approximately 1500 to 1200 Ma ago), and might in some way be due to the appearance
of eukaryotes. A sharp drop in stromatolite diversity occurs during the Vendian (675 to 570 Ma ago)
and is probably due to the activity of metazoans. Two diversity plateaus occur, one separates the two
diversifications and the other occurs after the last diversification and before the Vendian decline.
Stromatolites are the products of the complex interactions of microbial, sedimentary, and
environmental factors. While stromatolites are not well understood from a biogeological per-
spective, they do provide valuable evidence for ancient life, they are useful for biostratigraphy and
palaeoecology, and their distributional and diversity patterns provide insight into the first 3 billion
years of the history of life on this planet.
1 Introdnction and Background
Stromatolites have an extremely long fossil record spanning some 3500 million
years (Ma) or 83% of Earth's history. All eons, eras, and periods of geological
time represented by the rock record contain stromatolites. Their great antiquity,
persistence, abundance (in particular during the Proterozoic), and their oc-
currence today in a wide variety of environments, make them a fascinating
subject for scientific inquiry.
Stromatolites occupy a curious position in studies of the history oflife on
Earth. Some researchers treat stromatolites as sedimentary structures rather
than as fossils. Others treat stromatolites as single paleontological entities,
giving them binomial names. At times some researchers appear to lose sight of
the fact that they are the product ofcomplex microbial-sediment interactions. Still
'Permanent address: Department of Geological Sciences, Preston Cloud Research Laboratory,

University of California, Santa Barbara, CA 93106, USA
Present address: Department of Earth and Planetary Sciences, Western Australian Museum,
Francis Street, Perth, Western Australia 6000
290 S.M . Awramik
other investigators attempt to understand stromatolites from a palaeobiological

point of view. All approaches yield interesting and useful results.
Along with microbial fossils, stromatolites represent the most ancient traces
oflife yet discovered on Earth. Excellent examples of both have been found in
sedimentary rocks approximately 3500-Ma-old in Western Australia (Lowe
1980b; Walter et al. 1980) and South Africa (Byerly et al. 1986). Although
stromatolites appeared in the Early Archaean, they are uncommon Archaean
fossils. They become increasingly abundant and diverse in the Early Prote-
rozoic, and by late Middle Proterozoic time (about 1000 to 900 Ma ago)
continuing into the early Late Proterozoic (up until about 700 Ma ago),
stromatolites achieve their maximum diversity and abundance (A wramik 1971 ;
Walter and Heys 1985; Figs. I, 2). This Proterozoic richness in form and
abundance is unequalled at any other time. Domes, simple to elaborately
branching columns, conical structures, conical structures with branches, wavy
00.065 U CENOZOIC
0 . 225
0N MESOZOIC
0
0: PALEOZOIC
w
o .!! z
«
0 .570 :I:
CAMBRIAN
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UPPER
RIPHEAN
1.0 1 . 0S0 z
« MIDDLE
til w RIPHEAN
u:I:
a:: -0..
« 1.350 u
0
0-
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LOWER
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>- 0 0: RIPHEAN
lJ..
0: W
1 .6S0 w .-
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0 « 0:
0..
:I:
....... 2 .0 0..
EARLY
co W
0 0:
0..
PROTEROZOIC
0:
X 0
2 .5
z
til «
Z Ii
0 m
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...J «
...J 3 .0 U
w
0:
In 0..
ARCHEAN
Z
W 3 .5 3 .5 FIRST S TOLITES
2
t- 3 .8 OLDEST TERRESTRIAL ROCKS
4 .0
HADEAN
NO ROCK RECORO
4 .5
4.8 ORIGIN OF EARTH
Fig. 1. Relative abund a nce of stromatolites plotted against tim e

Archaean and Proterozoic Stromatolites 291
190
170
150
130
'"
x
~
.~ 110
0
iii
E
e
t)
90
'0
Q; 70
.0
E
:J
Z 50
30
10
2500 2000 ~ R500 Ma

co ll)
Early I Late I Early I Middle I Late Iv E:
Late Early Proterozoic I Riphean Paleozoic
Archean Proterozoic Phanerozoic
Fig. 2. Diversity of stromatolite taxa (mainly forms) through the Proterozoic and into the Cambrian.
Redrawn and modified from Fig. 1 of Walter and Heys (1985)
laminated structures, and unattached spheroidal structures with encapsulating

laminae (oncolites) were common during the Proterozoic. A number of these
stromatolites have sufficiently distinctive, temporally restricted shapes that they
are used as index fossils (see review by Bertrand-Sarfati and Walter 1981).
During latest Proterozoic time, < 675 Ma ago, stromatolites underwent a
profound decline in diversity and abundance. Presumably this was the result of
grazing pressure and sediment disturbance by metazoans (Awramik 1971).
Although Phanerozoic stromatolite abundance is noticeably lower when
compared to the Proterozoic, they are still somewhat common in Cambrian and
Lower Ordovician strata. The great radiation of benthic marine invertebrates
beginning in the late Early Ordovician (Sepkoski 1979) brought about the
second major decline in stromatolite abundance (few data are available to
permit statements on Phanerozoic stromatolite diversity). From the Middle
Ordovician to the present, stromatolites are generally uncommon.
Although common and conspicuous in pre-Phanerozoic strata, stroma-
tolites have not contributed much to attempts at synthesizing the early history
of life. Microbial fossils, primarily those found permineralized in chert, have
292 S.M. Awramik
been the major focus of such studies (Cloud 1976a; Knoll 1985). Recently,
acritarchs (organic-walled microbial fossils mostly of uncertain taxonomic
position that are recovered from clastic rocks using palynological techniques)
have received considerable palaeobiological attention and are providing im-
portant information on the history of plankton and eukaryotes during the
Middle and Late Proterozoic (e.g. Vidal 1984). Nevertheless, stromatolites,
which are the products of the interaction of microbes with sediment, have the
potential to provide important palaeo biological information on the history of
life in the Archaean and Proterozoic. After all, stromatolites appeared very early
in the fossil record, they achieved complex geometric arrangements oflaminae
and columns, they persisted over the eons, they exhibit interesting patterns of
abundance and diversification through geological time, and certain geometries
and microstructures have time-restricted distributions in the pre-Phanerozoic
and early Palaeozoic. The problem, however, lies in the very nature of
stroma tolites: they are organosedimentary structures prod uced by communities
of diverse microbes which are rarely preserved in the stromatolites. The
stromatolite represents the as yet incompletely understood interplay of biotic
and physical factors. Morphological attributes of stromatolites ultimately will
be understood relating microbes, community ecology, and biotic-sediment
interactions within a microbial evolution framework. Only then will stroma-
tolites contribute far more to understanding the biogeology of the pre-Phane-
rozoic Earth.
A key generalization for the understanding of stromatolites is that most of
them are built by photosynthetic microorganisms. Many researchers consider
cyanobacteria to be sine qua non for stromatolite construction. The prominant
role of cyanobacteria in ancient stromatolite construction was recognized by
Charles Doolittle Walcott, who, in 1914, discovered filamentous microfossils
resembling cyanobacteria in stromatolites from the Proterozoic Belt Super-
group (Walcott 1914). Since then there have been several important papers that
helped to establish the link between stromatolites and cyanobacteria: (1) Black
(1933) described modern Bahamian microbial mats built by cyanobacteria; (2)
Logan (1961) concluded that cyanobacteria are the dominant microbe re-
sponsible for the Recent Shark Bay stromatolites; (3) Barghoorn and Tyler
(1965) described microbial fossils resembling cyanobacteria from stromatolites
of the Early Proterozoic Gunflint Iron Formation; (4) Monty (1967) described
the stromatolites and cyanobacteria of Andros Island, Bahamas; (5) Schopf
(1968) described fossil cyanobacteria preserved in stratiform stromatolites of
the Late Proterozoic Bitter Springs Formation; and (6) Golubic's (1973) review
of the relationship of cyanobacteria, carbonates, and stromatolites. Numerous
other microbes (e.g. diatoms, rhodophytes, chlorophytes, green non-sulphur
bacteria, and even fungi) can contribute to stromatolite construction (see
Golubic 1976 and Krumbein 1983). For instance, benthic diatoms have recently
been acknowledged to playa major role in the construction of some Recent
stromatolites (e.g. nonmarine stromatolites at Cuarto Cienegas, Mexico, Wins-
borough and Golubic 1987; subtidal Shark Bay stromatolites, Awramik and
Riding 1988). In such stromatolite communities with algae and other bacteria,
cyanobacteria are nevertheless an important component of the microbial com-

munity. Although the primacy of cyanobacteria in stromatolite construction
from the Archaean to the Recent is a matter of some debate (see Walter 1983
for a discussion on Archaean stromatolites), I view that cyanobacteria are
and were the most significant biological agents for stromatolite construction.
2 Definition of a Stromatolite
Although the pre-Phanerozoic was the time of stromatolites' greatest

development, the term 'stromatolite' originated from the study of a Mesozoic
nonmarine occurrence. In 1908 Ernst Kalkowsky coined the German term
"stromatolith" (from the Greek stroma meaning bed, mattress, or layer and
lith as meaning stone) for masses of laminated nonmarine limestone in the
Lower Triassic Buntsandstein of the Harz Mountains, Germany. He also
introduced the term "stromatoid" for individual growth structures with more or
less planar layers but with a tendency toward upward convex doming that
occurred within stromatoliths. A microbial origin was attributed to these
structures by Kalkowsky. Over the years, the anglicized term stromatolite
gained favor; however, the term was applied to individual, convex upward,
laminated structures.
A number of researchers prefer a purely descriptive, nongenetic definition
of stromatolite and restrict usage to " ... attached, laminated, lithified,
sedimentary growth structures, accretionary away from a point or limited
surface of initiation" (Semikhatov et al. 1979, p.993 but note difference of
opinion in this paper). It is rarely possible to demonstrate a biological origin for
ancient stromatolites, hence preference by some researchers for such a non-
genetic definition. Adherents to the descriptive definition, nevertheless, ap-
parently recognize that most of the structures they would call stromatolites
probably formed as a result of microbial activity. The descriptive definition also
includes numerous abiogenic structures such as some calcretes, stalagmites,
boiler crusts, and certain geyserites.
On the other hand, a widely used definition of stromatolite (and the one
preferred in this chapter) is genetic and defines a stromatolite as an organo-
sedimentary structure produced by the sediment trapping and binding activity
of microorganisms, primarily cyanobacteria (from Awramik and Margulis in
Walter 1976, p. 1). This definition does not specify lithification, the presence of
laminations, or convexity away from the region of attachment. It is a process-
oriented definition that reflects the corpus of knowledge that has accumulated
over the years, indicating that most structures commonly referred to as
stromatolites were built by photosynthetic microorganisms. The definition
eliminates the need for separate appellations, some multiple and/ or cumber-
some, such as algal stromatolite, cyptalgalaminite, potential stromatolite, and
thrombolite for microbially produced structures that are here grouped underthe
umbrella term stromatolite. A new term, microbialite, has been introduced by
Burne and Moore (1987, p. 241) for an organosedimentary deposit formed from
294 S.M. Awramik
the interaction between benthic microbial communities and detrital or chem-

ical sediments. The term is virtually identical to the genetic definition of
stromatolite.
3 The First Stromatolites
Two Early Archaean greenstone belts, one in the Pilbara region of Western
Australia and the other in the Barberton Mountain Land of South Africa,
contain stromatolites in rocks radiometrically dated asold as 3500 Ma. Although
not morphologically complex, the stromatolites are nevertheless well laminated
and range in sha pe from undulatory, stratiform structures to domical structures,
some with a pseudocolumnar geometry (Lowe 1980b; Walter et al. 1980; Buick
et al. 1981; Byerly et al. 1986; Fig. 3). Oncolites are also found (Lowe 1983).
These Early Archaean stromatolites are found preserved as chert, and the
Pilbara stromatolites might have originally been calcium carbonate (Buick et al.
1981). The overall shape and microstructure of the stromatolites are similar to
many Proterozoic and younger examples. Apparently, no true columnar
stromatolites have yet been found in Early Archaean terranes. Conically-
shaped, laterally-linked laminae occur in some Pilbara stromatolites (Lowe
1983). Near both the Western Australian and South African stromatolite
localities, filamentous and coccoidal microbial fossils have been discovered in
chert (Awramik et al. 1983,1988 ; Walsh and Lowe 1985; Schopf and Packer
Fig.3. Early Archaea n stromatolites from the 3500-Ma-old Warrawoona Group, Western Australia
1987}. Most of the Western Australian filaments occur within thinner, dark
laminae of wavy laminated light-dark couplets and the filaments are commonly
oriented parallel to lamination; an organization suggestive of stromatolitic
laminae (Awramik et al. 1983, 1988).
The occurrence of stromatolites in 3500-Ma-old, Early Archaean strata has
great palaeobiological significance. On one hand, most geologists and
palaeontologists would regard stromatolites as evidence for the presence of
primitive organisms. The construction of a stromatolite, however, requires
microbes that are metabolically, behaviorally, and morphologically quite so-
phisticated (Awramik 1986). Thus, the presence of stromatolites on the Earth
3500 Ma ago indicates that microbial evolution had already evolved to a rather
advanced grade of prokaryote. However, do these structures therefore indicate
the presence ofcyanobacteria ? At a minimum, the Early Archaean stromatolites
indicate the presence of cyanobacteria-like microorganisms in terms of their
phenotype (in the full biological sense ofthe term). The filamentous microfossils
from chert of the same age present permissive evidence for cyanobacteria
(Awramiketal. 1983}while the encapsulated pluricellularcoccoids described by
Schopf and Packer (1987) argue well for cyanobacteria. The carbon isotopic
data support the interpretation that autotrophs were present in the ecosystem
(Walter 1983). The cumulative weight of these observations and data on
microfossils, stromatolites, and isotopic geochemistry provides presumptive
evidence that cyanobacteria had evolved by Early Archaean time. (Whether or
not these 'cyanobacteria' where oxygen-releasing photoautotrophs is a separate
question because many modern mat-building cyanobacteria can use photo-
system I). Phylogenetic reconstructions of prokaryotes based on molecular
biological techniques indicate that cyanobacteria are one of the latest prokary-
otic phyla to evolve (Woese 1987) and this supports the widely held conclusion
based on more traditional lines of research that among prokaryotes, cyano-
bacteria are phylogenetically one of the last phyla to appear (e.g. see Margulis
and Schwartz 1982, p. 24). Therefore, the presence of cyanobacteria in 3500-
Ma-old rocks is not a trivial issue for it would imply that prokaryotes had
undergone a great deal of evolution prior to 3500 Ma ago and that most major
clades of prokaryotes had already appeared. Evolution of prokaryotes during
the remainder of the pre-Phanerozoic was at lower taxonomic levels in already
established major clades; some of these clades may have gone extinct.
4 Other Archaean Stromatolites
Although the Archaean represents 1440 Ma of geological time and stromatolites

appeared early in the Archaean, they are actually rare in the Archaean (Fig. 4).
Less than 20 occurrences are currently known; although this number has slowly
increased over the past five years, the overall number oflocalities will probably
remain relatively low when compared with Proterozoic examples. This paucity
may be due to geological factors rather than biological factors. In the Archaean,
sedimentary basins developed within greenstone belts and sedimentation was
296 S.M. Awramik
Fig. 4. Late Archaean domical stromatolites from the approximately 2700-Ma-old Steep Rock
Group. western Ontario. Canada. Large dome in cenler of picture is about 3 m across
characterized by rapid deposition of compositionally immature sediments in

subsiding basins or in unstable alluvial settings (Lowe 1980a). Stable, shal-
low-water environments not deluged with sediments were uncommon and/or
short-lived in greenstone belt environments. Carbonates apparently were scarce
(Cameron and Baumann 1972). Hence, even phototropic microbes that could
grow fast, reproduce quickly, and which possessed rapid motility would have
had difficulty maintaining themselves at or near the sediment-fluid interface,
influence sedimentation, and produce a stromatolite. The development of
sizable, stable, shallow water environments (i.e. continental shelves and
platforms) concomitant with Proterozoic-style tectonic regimes toward the
end of the Archaean and in the early Proterozoic resulted in a proliferation of
stromatolites (Cloud 1976b). The passive margins of continents, when in-
undated, provided relatively large, stable, shallow shelf environments con-
ducive to stromatolite formation. In addition, depending on climatic and
local tectonic conditions, such environments could also be major sites for
the production and accumulation of carbonate sediments. The great major-
ity of fossil and recent stromatolites are found as limestone or dolostone con-
structions. This relationship is not fortuitous. Cementation and lithification
of microbiologically accreted sediment are crucial for the preservation of
stromatolites and the carbonate geochemical system provides an excellent
means of fossilization . Also the uptake of CO 2 during photosynthesis would be
expected to induce calcium carbonate precipitation.
Most ofthe major morphological varieties of stromatolites had appeared by

the end of Archaean time. These include wavy laminated (stratiform), nodular,
domical, pseudocolumnar, columnar, conical, and oncolitic types. However, in
the Archaean, unlike the Proterozoic, columnar stromatolites are rare, few
branching columnar stromatolites are found and of these no complexly
branched varieties have been discovered. The known Archaean stromatolites
have shapes that are common throughout geological time; no distinctive
morphology unique to the Archaean has yet been recognized. The micros-
tructure of Archaean stromatolites is similar that found in many Proterozoic
stromatolites. Based on studies of Proterozoic and Recent stromatolites, our
current understanding on the development of microstructure suggests that
many of these Archaean stromatolites were built by filamentous cyanobacteria
(Walter 1983).
5 Proterozoic Stromatolites
Palaeontologically, the Proterozoic is known for its stromatolites. It was during

this long interval of geological time (some 1930 million years, from 2500 to 570
Ma ago) that stromatolites reached the acme of their diversity and abundance
and then underwent drastic reductions in diversity and abundance at the close
of the Proterozoic. Basic stromatolite architectures (cones, columns, domes, etc.)
were already established by Late Archaean time, and late in the Early Prote-
rozoic, conical, columnar, and columnar-branching stromatolites become
conspicuous components in stromatolitic deposits. Many Proterozoic stroma-
tolites are well preserved on both the macroscopic and the microscopic level, and
in a few cases, microfossils are found in silicified laminae. A useful way of
studying stromatolites is to consider them from all observational levels, from the
large metre-sized bioherm or biostrome groupings of individual structures
down to the microscopic level of microfossils and microstructure (Fig. 5). A
conspicuous component of Proterozoic stromatolites that separates this time
interval from the Archaean and Phanerozoic is the presence of numerous
columnar-branching and conical stromatolites (Figs. 6, 7). Following the ex-
ample of James Hall (1883) in giving stromatolites binomial names and tax-
onomic descriptions, geologists and palaeontologists who describe stromatolites
and use them in biostratigraphy have devised several techniques for their study.
The serial sectioning - gra phical reconstruction method of Krylov (1963) is the
most widely used technique for studying the morphology of individual
stromatolites and in the naming of stromatolites. Studies on the biostratigraphic
distribution of stromatolites for the Proterozoic has produced workable schemes
for the subdivision of Proterozoic time (see reviews by Semikhatov 1976;
Bertrand-Sarfati and Walter 1981).
A brief analysis of data and the patterns of diversification (using primarily
the stromatolite taxonomic rank ofform; Fig. I) presented in Walter and Heys
(1985) and an analysis of stromatolite information presented elsewhere for
Proterozoic stromatolites results in the following observations [Note: (a) Va-
298 S.M. Awramik
megascopic
Bioherm or Biostrome
Column. Dome. Club. and

11cm
Branching Column
Lamina Ilm m~
nA
Microstructure
Microbiota \II~ microscopic
Fig. 5. Levels of observations and organization in stromatolites
Fig. 6. Bait'alia maurilanica, a columnar branching stromatolite from the Late Proterozoic Atar
Group, Mauritania
Fig. 7. Weathered. longitudi-

nal section of Conophyton jac-
queti from the Late Proterozoic
Atar Group. Mauritania
riation between the points on the graph of Fig. 2 is impossible to determine

because of the imprecise age determinations and difficulties with the taxonomy
of stromatolites. Only the broadest or sketchiest of patterns are possible to
elucidate; (b) Only data used in Walter and Heys (1985) are used. Taxa
described from India are not well represented in these data and several new
monographs have been published (most notably Zhu et al. 1987, Early Prote-
rozoic stromatolites) since 1985. These new additions are currently being
compiled): (I) Only six taxa of stromatolites have been described from the first
300 Ma of the Early Proterozoic (2500 to 2200 Ma ago). (2) Two diversifications
occurred, one during the late Early Proterozoic, 2200 to 1650 Ma ago, and the
other during the Middle Riphean, 1350 to 1050 Ma ago. (3) Two diversity
plateaus occurred, one during the late Early Proterozoic and into the early Early
Riphean, around 1350 Ma ago, and the other from Middle to Late Riphean,
around 1050 Ma ago. And (4), a decrease in stromatolite diversity occurred
beginning in the Vendian, about 676 Ma ago, and continuing into the Cambrian.
300 S.M. Awramik
1. The small number (six) of stroma tolite taxa from 2200- to 2500-Ma -old Early
Proterozoic strata reflects lack of detailed taxonomic treatment and insufficient
study before 1985. This relatively low diversity (and low abundance) of
stromatolites (Walter and Heys 1985) may be real and reflect low diversity of
cyanobacteria, oxygen deficient conditions, and the small number of
sedimentary basins as noted in Walter and Heys (1985). The small number of
basins may reflect what Goodwin (1981) recognizes as a transitional phase
between the Archaean greenstone belt-type tectonics and the macro-plate
tectonics characteristic of Middle Proterozoic through Phanerozoic time when
ensialic basins developed in an intra-plate tectonic setting.
Oxygen-deficient conditions could have influenced cyanobacteria during
the earliest Proterozoic and Archaean. Many modern cyanobacteria, when
operating with photo system II (oxygen-releasing photosynthesis) are
microaerophilic; that is, they are adapted to low oxygen levels. Extending this
observation into the past, it can be speculated that the oxygenation of the
atmosphere beginning about 2300 Ma ago (see review by Cloud 1983), could
have been pivotal in the evolution and spread of cyanobacteria. Prior to this
time, many cyanobacteria might have been restricted to oxygenated environ-
ments largely of their own creation; localized regions where cyanobacteria were
growing and producing O2 , Microaerophily could make cyanobacteria 'pre-
adapted' to the oxygenated conditions that were to begin about 2300 Ma ago,
and they could easily disperse and readily colonize newly oxygenated shallow
water habitats.
The diversity of earliest Proterozoic microfossils found in stromatolites is
low when compared to Middle-Late Proterozoic examples. My reading of the
Early Proterozoic microfossil record indicates that all the major orders of
cyanobacteria were already present; however, diversity within these orders
appears to be low. The low diversity could have been the result of competition
with other prokaryotes. This low diversity of such microbes might have been a
contributing factor to the low diversity of stromatolites (this assumes that there
is a strong relationship between microbiota and shape in stromatolites; a
relationship that remains to be demonstrated).
2. The first diversification occurred during the late Early Proterozoic (2200 to
1650 Ma ago) when the number of stromatolite taxa increased to 97 (Walter and
Heys 1985). This large diversification represents the first significant adaptive
radiation of stromatolites and probably the microbes that built them. Macro-
plate tectonics began to dominate producing more 'basins' (26 noted in Walter
and Heys 1985). Continued oxygenation of the atmosphere and shallow hy-
drosphere occurred and allowed for greater cyanobacterial dispersal. Cyano-
bacteria probably underwent a major adaptive radiation. The appearance of
new cyanobacteria (species, genera, families within existing orders) in the
mat-building community with improved abilities to accrete sediment and form
cohesive fabrics, could have resulted in new geometric arrangements in the
stacking oflaminae. For example, change in the consistency of extracellular gel
(sheath and envelope), greater motility, and faster growth could alter the
sediment-microbe plexus in such a way that the interaction of the mat with
factors of the physical environment (water depth, currents, wave action,
sediment supply, temperature) would result in a new style of branching or some
other change in the shape of the stromatolite. The microbial fossil record of
cyanobacteria during this interval of time is not good. What few data are
available tend to support this speculation.
The second diversification noted above which occurred during the Middle
Riphean, approximately 1500 to 1200 Ma ago when the diversity increased from
98 taxa to 182 taxa, is much more problematical. It follows a diversity plateau
(see Fig. 2) and there is no suggestion in the available data of a decrease in
taxonomic diversity preceding this diversification. In view of the lack of any
evidence to suggest otherwise, atmospheric level of O2 was probably increasing
monotonically. Phanerozoic-style plate tectonics were well established, but no
clear suturing or breakup of major land masses has yet been determined for this
interval of the Proterozoic. In addition to or aside from another phase of
cyanobacterial diversification [the microfossil record in stromatolites, although
not great, does show an increase in diversity (Schopf 1977), but not the
pronounced increase one might associate with the profound increase in
stromatolite diversity], the only possible major change in the Earth-biosphere
system that might have had some effect on increased stromatolite diversity was
the appearance of eukaryotes. Although the subject of considerable debate, the
oldest eukaryotes are pro ba b ly photoa utotrophic plankters that appeared a bou t
1500 Ma ago (Vidal 1984). Possible benthic algal eukaryotes appear in the
record about 1300 Ma ago (for instance Beltina from the Greyson Shale; see
review by Hofmann 1985). The competition between benthic eukaryotic algae
and cyanobacteria might have resulted in a further cyanobacterial
diversification giving rise to an increase in stromatolite taxa. The role that
eukaryotes play in stromatolite construction today is not well understood;
however, it is possible that benthic, microbial eukaryotes took part in stroma-
tolite construction at this time, contributing to the production of new groups and
forms of stromatolites. A problem with this hypothesis is that there is no clear
indication in the microfossil record as preserved in stromatolites that eukaryotes
participated in stromatolite construction. Fossil cyanobacteria are the major
fossil microbes found. The causes for the doubling of diversity from Early to
Middle Riphean times remain obscure.
3. The diversity plateau during the late Early Proterozoic through Early Ri-
phean and during the Middle to Late Riphean is another curious feature of the
stromatolite diversity curve. The increase in diversification followed by a
tapering off (the plateau) resembles the sigmoidal growth phase curve that
increasing populations of various organisms follow and it resembles the curve
of the rapid diversification of animals at the beginning of the Phanerozoic (see
Sepkoski 1979). A plateau or tapering off in diversification curves is usually
explained as a result of filling of available ecological zones by the clades
(Bambach 1985; Valentine and Walker 1986). Employing Bambach's (1985,
p.247) explanation, this would mean that the adaptive range of the stroma-
302 S.M. Awramik
tolite-building microbes, primarily cyanobacteria, with its existing phenotypic

attributes, reached a diversity balance related to the ecological space the
organisms are able to utilize.
4. The sharp decrease in stromatolite diversity during the Vendian and con-
tinuing into the Cambrian has been noted and discussed elsewhere (Awramik
1971; Monty 1973; Pratt 1982; Walter and Heys 1985; Cao 1988). The cir-
cumstantial evidence still implicates grazing and burrowing by macrofauna and
meiofauna that probably appeared in the Vendian. Also, the advent of
biomineralization in the Tommotian, which produced coarse-grained car-
bonate sediment, changed the nature of the sediment available for stromatolite
accretion (Awramik and Riding 1988). Metazoans effectively outcompeted the
microbes that formed microbial mats/stromatolites in shallow marine envi-
ronments and brought about the stromatolite crash and prevented re-estab-
lishment of major stromatolite-building activities.
6 Summary and Conclusions
Although stromatolites received their name from a Phanerozoic example, it was

during the pre-Phanerozoic that stromatolites achieved their greatest
development. Until recently, the understanding of Proterozoic and Archaean
stromatolites has proceeded along lines comparing these ancient structures to
modem stromatolites and microbial mats forming in some of today's shallow
marine environments. Hoffman (1973), however, pointed out that the present
may not be an accurate analogue for the pre-Phanerozoic Earth. Many
stromatolites known from Proterozoic strata have no known morphological
counterparts in the Recent or even the Phanerozoic. In addition, most of Earth's
history was animal-free; oxygen levels were much lower in the Late Proterozoic
and earlier; shallow water, carbonate accumulating environments in the pre-
Cambrian (environments where stromatolites have thrived) lacked the abun-
dant bioclastic material found in later times; and the microbial ancestors of
today's stromatolite builders were possibly phenotypically as well as geno-
typically different.
Acknowledgements. Numerous cups of coffee at Nicoletti's with l.W. Valentine helped to sharpen
some of the ideas expressed in this paper. Thanks to David Pierce for comments on the manuscript
and to Karen Griffin and David Crouch for help with the figures. NATO Research Grant
RG84/0176 to R. Ridingand S.M. Awramik, and NSF Grant EAR 87-21192 provided funds to help
support the research on stromatolites. This is a contribution of the Precambrian-Cambrian
Transition Consortium and IGCP Project 261 "Stromatolites".
Archaean and Protero,zoic Stromatolites 303
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Chapter 16
Cambrian Calcareous Cyanobacteria and Algae
R. RIDING l
Abstract
The Cambrian calcareous flora is dominated by cyanobacteria which include the Angulocellularia.
Botomaella. Girvanella and Obruchevella groups. The important generic groups based on Renalcis
and Epiphyton are also probably cyanobacterial. Proaulopora is possibly a cyanobacterium. No
calcified red algae are known during the period. The earliest calcified dasycladaleans may be
represented by the very rare fossils Yakutina (Middle Cambrian) and Seletonella (Upper Cam-
brian). The sudden appearance of heavily calcified cyanobacteria near the base of the Cambrian
may reflect a change in environmental conditions which enhanced CaCO, precipitation rates in the
sea. Groups such as Angulocellularia. Epiphyton and Renalcis are important in creating reef
limestones and they are often so abundant that they impart a dendritic fabric to the rock (dendrolite).
The more common of these fossils appear to have a worldwide distribution but work has been
heavily concentrated in the USSR. on Siberian material and on the Lower Cambrian. Only recently
have other workers started to make use of the Soviet results, and many genera have not been reported
outside the USSR and Mongolia. although Cambrian floras are known in Europe, North America,
Australia and Antarctica. The outstanding problem which remains is of confidently interpreting the
affinity of many groups. More also needs to be known concerning the ranges of all the genera.
particularly in the Middle and Upper Cambrian. and of their spatial and ecological distributions.
1 Introduction
The Cambrian is significant not only for the earliest abundant remains of animal
skeletons and shells, but also for the first occurrence of well-calcified algae and
cyanobacteria. Biomineralization was therefore acquired by a very wide
spectrum of the organisms on Earth in a short space of time. In the case of
cyanobacteria, in particular, this was a striking development because this group
has a long Precambrian history. It is possible to explain the relatively sudden
appearance of skeletons in metazoans as being a result of rapid evolutionary
diversification from the late Proterozoic to the Cambrian. Indeed, this could also
apply to algae and cyanobacteria. However, the latter group is widely regarded
as being conservative, and it is known that a range of cyanobacteria comparable
with extant forms existed in the Proterozoic. Cyanobacteria do not closely
control their calcification but are dependent upon environmental conditions
favouring precipitation of CaCo3 . It is for these reasons that the abrupt ap-
pearance of well-calcified cyanobacteria near the Precambrian-Cambrian
boundary (Riding and VOfonova 1982a) has been interpreted to possibly reflect
a change in an environmental factor such as seawater chemistry (Riding 1982).
'Department of Geology, University of Wales College of Cardiff. Cardiff CF I 3 YEo UK

306 R. Riding
The Cambrian algal and cyanobacterial calcified flora is quite well known
in terms of taxonomic diversity, but there are numerous problems of affinity
remaining. These make it difficult to confidently generalize concerning the
sequence and timing of appearance of major groups such as chlorophytes,
rhodophytes, and so on. Earlier workers, for example Korde (1973), were ofthe
opinion that red algae were particularly important in the Lower Cambrian.
However, these fossils, such as Epiphyton, Botominella, Bija, etc. could well be
cyanobacteria. In fact, at present it seems quite likely that: (1) calcified cyano-
bacteria overwhelmingly dominate the Lower Cambrian flora; (2) no bona
fide red calcareous algae occur in the Cambrian; (3) dasycladalean green algae
are rarely represented in the Cambrian (by the problematic genera Cam-
broporella in the Lower Cambrian, Amgaella and Yakutina in the Middle
Cambrian, and Seletonella and Mejerella in the Upper Cambrian); and (4) no
bona fide udoteacean green algae occur in the Cambrian. However, such
generalizations are dangerous given the imperfect state of current knowledge of
the systematic positions of many of the fossils involved.
Shallow marine limestones of Cambrian age often contain abundant cal-
careous cyanobacteria and algae. Commonly they are rock-formers, especially
in reefs. Although Lower Cambrian bioherms have often been regarded as
dominated by archaeocyaths, in reality genera such as Angulocellulara, Epi-
phyton and Renalcis are often more important and are actually equally con-
spicuous because they impart a macroscopic dendritic fabric to the rock
(dendrolite, Riding 1988). However, the small size of most of these fos-
sils makes them susceptible to diagenetic alteration. Consequently, they
are best studied in cratonic areas where the sequences are relatively un-
disturbed.
The details of the earliest a ppearance of the calcified flora near the base of
the Cambrian are only recorded at a few localities around the world and they
have been mainly studied in Siberia, which has more than its fair share of Lower
Cambrian limestone sequences. Indeed, current knowledge of Cambrian cal-
careous cyanobacteria and algae is largely the result of painstaking work in the
Soviet Union, mainly since 1930. However, only recently have these results
started to be applied and evaluated by workers outside the USSR. The strength
of the Soviet contribution has been in taxonomic analysis, and this has been
combined with generally good stratigraphic control. There has been some
tendency to proliferate taxa. However, the major problems, still unresolved in
many cases, concern affinity. There has been conspicuous instability in the
assignment of generic groups to higher taxonomic categories, particularly at
division (red algae, green algae, etc.) level.
The most reliable guide to affinity lies in recognition of modern analogues.
Many of the more common Cambrian genera range beyond the Cambrian and
high into the Palaeozoic. Some, all of them cyanobacteria, possess Recent
analogues. In the absence of analogues it is difficult to make progress in
systematic evaluation. However, it is possible to rationalize taxonomy by
sidestepping problems of affinity and concentrating on purely morphological
groupings (Riding and Voronova 1985). This is clearly only a partial approach
Cambrian Calcareous Cyanobacteria and Algae 307
to the problem, but it serves to refine the descriptive base for systematic
classifications which will follow.
It is possible to hypothesize that, following the crisis for stromatolites
resulting from the appearance and rapid diversification of metazoans in the Late
Precambrian, calcification provided a new and more secure mode of life for
cyanobacteria. Animals disturbed, grazed and competed for space with
stromatolites (Garrett 1970). Calcification provided physical strength and
protection for cyanobacteria to resist disturbance and grazing, and it also gave
them the possibility of a reef-building role. Cambrian reefs can be seen as
habitats in which, in many cases, cyanobacterial calcification provided a firm
substrate for sessile animals such as archaeocyaths. This assumes that the
conspicuous dendrolite-forming genera (A ngulocellularia, Epiphyton, Renalcis)
are all cyanobacteria. Angulocellularia has a Recent analogue in the cyano-
bacterium Schizothrix calcicola (Riding and Voronova 1982b); Epiphyton and
Renalcis are now often assumed to be cyanobacterial. The small pore spaces
between the individual filaments of these genera may well have localized early
cementation under the stimulus of photosynthetic metabolism, and this would
have provided a further stimulus to reef formation.
It could also have been difficult for stromatolite-building cyanobacteria to
cope with the coarse bioclastic sediment which animal biomineralization was
beginning to produce (Pratt 1982). Again reef-building, by raising the substrate,
would have provided an elevated habitat which reduced the possibility of
cyanobacteria being drowned by particulate sediment. Thus, for a variety of
reasons, calcification by cyanobacteria at the beginning ofthe Cambrian can be
seen as a successful response to metazoan and parazoan diversification and
biomineralization. However, whether or not this was the prime stimulus for
cyanobacterial calcification remains to be seen.
A number of important algal groups has long been thought to be repre-
sented by calcified fossils in the Cambrian. Johnson (1966, p. 1) noted "the high
degree of evolutionary development exhibited by the Cambrian flora and its
great variety and richness of form". Johnson had in mind all algae, not only
calcified ones, but he was without doubt influenced by Soviet workers, such
as Vologdin (1962a), who had ascribed a variety of genera (including
Razumovskia, Renalcis, Epiphyton, Chabakovia, Angulocellularia, and Bija) to
the Rhodophyta. However, there is no clear evidence that any of these is likely
to be a red alga and the same applies to many other genera which Korde (1973)
placed in the Rhodophyta. Solenopora is an example. A Solenopora-like or-
ganism reported by Priestley and David (1910) from Antarctica, proved to
belong to Epiphyton (see Gordon 1920, p. 684). However, Maslov (1937) and
Vologdin (1955) later reported Solenopora from the Upper Cambrian of the
Angara River and the Cambrian of the Tien Shan in Kirgizia. Nevertheless,
Solenopora has often been confused with other genera, and at present no definite
reports of Solenopora, as it was originally described by Dybowski (1878), are
known outside the Ordovician. Bija, which resembles so-called Solenopora (see
Johnson 1966, p. 15), is actually quite distinct and more closely resembles
Hedstroemia which is probably a rivulariacean cyanobacterium.
308 R. Riding
Apart from calcified cyanobacteria, the only significant Cambrian group

which must be considered here is the Seletonellaceae (Korde 1971, quoted in
Korde 1973, p: 239) which Bassoullet et al. (1979) regard as the earliest of the
three dasycladalean families, and which has representatives mainly in the
Middle and Upper Cambrian.
2 History of Research
Cambrian calcareous algae and cyanobacteria have been studied for little more
than 100 years. The main advances have been made during the past 50 years as
a result of studies in Russia of Siberian material. Of the 74 genera discussed here
(Table 1),69 have been originally described in the Soviet Union and few ofthese
Table 1. Cambrian genera mentioned in the text
Acanthina Korde 1973 Kyzassia Korde 1973

Amgaella Korde 1957 Lebedevaella Vologdin 1958
Amgaina Korde 1973 Lukaschevella Vologdin 1958
Amganella Reitlinger 1959 Marenita Korde 1973
A ngulocellularia Vologdin 1962 Mejerella Korde 1950
Bajanophyton Drosdova 1980 Mucilina Korde 1973
Batenevia Korde 1966 Nephelostroma Dangeard & Dore 1957
Bija Vologdin 1932 Nicholsonia Korde 1973
Bogutschania Korde 1965 Nubecularites Maslov 1937
Bostrychosaria Vologdin 1958 Obruchevella Reitlinger 1948
Botomaella Korde 1958 Palaeomicrocystis Korde 1955
Botominella Reitlinger 1959 Palaeonites Maslov 1956
Cambrina Korde 1973 Parachabakovia Korde 1973
Cambroporella Korde 1950 Potentillina Korde 1973
Cavifera Reitlinger 1948 Proaulopora Vologdin 1937
Chabakovia Vologdin 1939 Pseudoanthos Korde 1973
Chomustachia Korde 1973 Razumovskia Vologdin 1939
Cornutula Korde 1973 Renalcis Vologdin 1932
Dasycirriphycus Vologdin 1962 Sajania Vologdin 1962
Epiphyton Bornemann 1886 Seletonella Korde 1950
Epiphytonoides Korde 1973 Serligia Korde 1973
Erbina Korde 1973 Siberiella Korde 1957
Filaria Korde 1973 Solenopora Dybowski 1878
Fistulella Korde 1973 Sporinula Korde 1973
Flexurella Vologdin 1958 Subtifloria Maslov 1956
Foninia Korde 1973 Taninia Korde 1973
Gemma Luchinina 1982 Tarthinia Drosdova 1975
Girvanella Nicholson & Etheridge 1878 Tomentula Korde 1973
Globuloella Korde 1958 Tubomorphophyton Korde 1973
Glomovertella Reitlinger 1948 Tubophyllum Krasnopeeva 1955
Gordonophvton Korde 1973 Tuvaellina Vologdin 1958
Hedstroemia Rothpletz 1913 Umbellula Korde 1973
Kadvoya Korde 1973 Uranovia Korde 1958
Kenella Korde 1973 Vologdinella Korde 1957
Kordephrton Radugin & Stepanova 1964 Vologdinia Korde 1973
Korilophyton Voronova 1976 Yakutina Korde 1973
Kundatia Korde 1973 Zaganolomia Drosdova 1980
- - - - --~--- - - - -
have yet been reported from other parts of the world. Nonetheless, there is a
number of other regions where these fossils are known to occur, notably in
Antarctica, Australia, China, Europe, Morocco, and North America, but these
have not received close attention either because their carbonate sequences are
not as well preserved as those in Siberia, or simply because geologists have not
been available to do the work. At the time of writing (1988) the history of
research divides into pre-Soviet (1886-1932) and Soviet (1932-1988) periods
(Fig. I). As far as description of new genera is concerned, only 5 of the 74 genera
listed here (Table I) were originally described outside the USSR, and since 1932
Nephelostroma is the only new genus described from the rest of the world.
Vologdin (1932, 1939) started the Soviet spate of geperic descriptions, and
Korde overwhelmingly dominates the period between 1950 and 1973. In fact,41
of the 74 genera in Table I are Korde genera.
However, the Soviet contribution to this research did not begin until the
1930s, some fifty years after Bornemann's (1886, pp. 16-17) discovery of Epi-
phyton, together with a new Con[ervites species, C. primordialis, which is
Epiphyton-like, from the Lower Cambrian of south-western Sardinia. Epi-
phyton is a widespread and abundant fossil and it was subsequently recognized
in Cambrian limestones from Antarctica (Chapman eta!. 1914; Chapman 1916)
where it had already been reported as a "Solenopora-like organism" by Priestley
and David (1910). Gordon (1920) described a new species, E. grande, also from
Antarctica which Korde (1973) was l ater to use to erect the genus
Gordonophyton .
Girvanella was originally described from the Ordovician (Nicholson and
Etheridge 1878) but it is common in the Cambrian. Blackwelder (1907) and
Yabe and Osaki (1930) repQrted it from China, Chapman (1907-1908) and
Mawson and Madigan (1930) from Australia, Hazzard and Crick may (1933)
from California, and Mercier (1934) from France.
30
20
10
1870 1890 1910 1930 1950 1970 1990
Fig. 1. Publications of genera of calcareous algae and cyanobacteria reported from the Cambrian.
Synonyms and incertae sedis included. For list see Table I. Only 5 (shown in black) of the 74 genera
shown were originally described outside the Soviet Union : Solenopora 1877 (reported , but not
actually present. in the Cambria n). Girvanella 1878. Epiphyton 1886, Hedstroemia 1913, Ne-
phelostroma 1957. Five of the 6 genera described during the 1930s were by Vologdin (V), and 10 of
the 21 genera described in the 1950s were by Korde (K) . All but 2 of the 31 genera described in the
1970's were in Korde's (1973) monograph (K)
3\0 R. Riding
The third well-known Cambrian genus, which ranks in importance

alongside Epiphyton and Girvanella, is Renalcis, which Vologdin described from
the Altai Mountains in 1932. Following the First Five Year Plan (1928-1932),
development of Siberia moved forward quickly. Cambrian and associated
Lower Palaeozoic carbonate sequences are extensively exposed in Siberia, and
are also present in the Urals to the west and Altai Mountains to the south (Fig.
2). The largest and best preserved outcrops are on the Siberian Platform (which
approxima tes to the Central Siberian Platea u), particularly in the Ana bar region
in the north and the middle reaches of the Lena River and its tributaries in the
south-east, but also along the Yenisei River on the western side of the Platform
and in the Angara River area near Irkutsk. Of all these the prime region is that
ofthe middle Lena above Yakutsk together with its tributaries such as the Amga,
Botoma and Sinyaya. This area is more accessible than the remote Anabar
region and it has been intensively sampled.
The presence of these extensive and well-preserved Lower Palaeozoic
carbonates, and the need to subdivide and correlate them for survey purposes,
resulted in the discovery of abundant calcareous algae and cyanobacteria, along
with stromatolites and archaeocyaths. All received attention. The two principal
early workers were Vologdin and Maslov. Vologdin's chief interest at this time
lay in archaeocyaths, but in his papers he also described the algae and cyano-
bacteria which he found associated with them. These included first descriptions
of Bija and Renalcis in material from the Altai Mountains south-east of
Krasnoyarsk (Vologdin 1932) and the first description of Chabakovia from
Kuvandyk in the southern Urals (Vologdin 1939).
As remote areas became more accessible, palaeontologists often joined
survey teams, but before World War II and sometimes afterwards too, it was
normal for field geologists making reconnaissance studies, often along rivers, to
provide material for study by palaeontologists back in Moscow or Leningrad.
Some of these collectors gave their names to the fossils they helped discover; for
example N.K. Razumovsky and E.E. Razumovskaya to Razumovskia Vologdin
(1939). In other cases the river provided the name: the Botoma both to
Botominella Reitlinger (1959) and to Botomaella Korde, the Amga to Amgaella
Korde (1957b), and so on.
The area of the rivers Aldan, Amga, Botoma and middle Lena upstream
from Yakutsk is the largest single outcrop of Lower Cambrian carbonate rocks
in Siberia, and probably in the world. It is comparable in size to England. Maslov
(1937) studied Lower Palaeozoic strata from this general region of the southern
Siberian Platform and described the new genus Nubecularites (a junior synonym
of Renalcis) from the Middle Cambrian and Epiphyton and Girvanella from the
Lower Cambrian. Subsequently this area was to prove a fruitful source of
material for Korde and later workers. However, World War II temporarily
halted this geological research in the USSR.
Elsewhere, progress in the study of Cambrian algae and cyanobacteria was
slow. The main activity in this field during the 1940s was in North America, but
it was largely limited to routine reports of Girvanella from various localities,
including the Grand Canyon (McKee and Resser 1945); Caborca, Mexico
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Fig. 2. Locali ties of Camb rian calcareous cyanobacteria and a lgae in Siberia and adjacen t areas
312 R. Riding
(Cooper and Arellano 1946; later Johnson 1952 also worked here), and the latest
Cambrian-Early Ordovician Ellenburger Group of Texas (Cloud and Barnes
1948; subsequently, Ahr (1971) described the Girvanella and Renalcis bioherms
of the underlying Wilberns Formation in detail).
Following World War II, intensive work on Cambrian calcareous algae and
cyanobacteria in the USSR by Korde resulted in a series of publications during
the 1950s. Korde (1950a) described the unusual and very rare dasycladalean-
like fossils Seletonella (named after the River Seleta) and Mejerella from the
Upper Cambrian on the southern edge of the West Siberian Plain, within
northern Kazakhstan; and also the somewhat similar Cambroporella from the
Lower Cambrian of Tuva, south of the Altai Mountains (Korde 1950b). But
subsequent work concentrated on Botomaella, Epiphyton, Renalcis and as-
sociated fossils from the Lower Cambrian of the middle Lena region (Korde
1955, 1957a, 1958a,b, 1959), and on more Seletonella-like genera (Amgaella and
Siberiella) (later named Yakutina by Korde 1973) from the Middle Cambrian
of the same area (Korde 1957b). About this time Reitlinger (1959) produced an
atlas of Siberian microfossils which included Obruchevella from the Upper
Cambrian or Ordovician.
This period of Korde's work culminated in an important monograph on
"Cambrian algae of the south-eastern Siberian Platform", i.e. the middle Lena
region (Korde 1961). In this publication were listed no less than 67 species of
Epiphyton, 62 of them being new ones. Shortly afterwards, in 1963, Korde and
Maslov co-authored sections on rhodophytes, cyanophytes and chlorophytes in
the Russian fossil treatise, Osnovy Paleontologii.
Meanwhile, Maslov himself, who had worked widely throughout the
Phanerozoic, had produced his impressive book on "Fossil calcareous algae of
the USSR (Maslov 1956), before turning his attention to stromatolites (Maslov
1960). Vologdin's work was also moving in that direction and he illustrated
much new material from the Sinian (Upper Proterozoic), Cambrian and Or-
dovician of the Angara (near Irkutsk) and Lena regions, emphasizing
stromatolitic microfabrics (Vologdin 1962a). In the same year he published a
study of Cambrian archaeocyaths and algae from the Baikal Platform (Vologdin
1962b).
There was nothing happening in the world outside the Soviet Union to
compare with this level of innovative research on calcareous algae, cyanobac-
teria and stromatolites in general, and particularly on those from the late
Precambrian and Early Palaeozoic. Dangeard and Dore (1957) in France
described Nephelostroma (yet another junior synonym of Renalcis) together
with Epiphyton and Girvanella from the Cambrian of Normandy (see also Dore
1962). In America, Rezak (1957) discussed the stratigraphic distribution of
Girvanella. 1961 saw the publication of Johnson's influential book
"Limestone-building algae and algal limestones" , but this failed to mention, not
only Epiphyton, but also Renalcis and all the other Cambrian genera described
from the Soviet Union. However, this serious omission was redressed when
Johnson reviewed the Cambrian algae (Johnson 1966), drawing heavily on
Soviet work: more than halfthe illustrations in this publication being taken from
either Korde, Maslow or Vologdin. Hill (1964) described Epiphyton and Re-
nalcis associated with archaeocyaths from the Beardmore Glacier, Antarctica.
Back in the USSR, Korde was preparing her book on Cambrian algae
(Korde 1973) which brought together nearly 50 genera in a comprehensive
taxonomic treatment which included detailed, cellular-level reconstructions of
Epiphyton and other fossils (many being interpreted as red algae, for which five
orders and seventeen new families were introduced) and a detailed evolutionary
scheme for cyanobacteria. However, a new generation of workers was emerging,
with interest focusing on the Precambrian-Cambrian boundary on the Siberian
Platform (Voronova 1969, 1976, 1979; Stepanova and Luchinina 1982) as well
as continuing with work on the Lower Cambrian of the Siberian Platform in
general (Luchinina 1975). Drosdova (1980) extended this to a study of algae in
Lower Cambrian bioherms in western Mongolia, where algae and cyanobac-
teria close to the boundary were also described (Voronova in Voronin et al.
1982). Some reassessment of earlier work, such as Korde's proliferation of
Epiphyton species, took place (Gudymovich 1967) and Luchinina's (1975)
classification marked an important new approach by suggesting that all the
principal genera belong to the cyanobacteria. But, in general, Soviet work of the
1970s applied and extended preceding lines of research, albeit with stratigraphic
refinements and a rather more critical eye for problems of affinity.
Cambrian calcareous algae and cyanobacteria still did not arouse the same
level of interest elsewhere, and much of the Soviet work went unutilized;
nevertheless, reports of occurrences ofthese fossils became more common from
localities as scattered as western Canada (Handfield 1971: Epiphyton and
Renalcis; McIlraeth 1977: Epiphyton), SouthAustralia (Haslett 1976: Renalcis;
Brasier 1976: Renalcis), southern Spain (ZamarreflO 1977: Epiphyton, Renalcis),
Labrador (James and Kobh.lk 1978: Angulocellularia identified as Renalcis or
Renalcis-like; Kobluk and James 1979: Epiphyton, Girvanella, Renalcis, Ser-
ligia); Mali (Bertrand-Sarfati 1979: a Bija-like fossil of Cambrian aspect from
presumed Vendian); Nevada (Kobluk 1981, Rowland 1981a,b: Epiphyton,
Renalcis); Newfoundland (James 1981: Epiphyton, Girvanella, Renalcis); Vir-
ginia (Pfeil and Read 1980: Epiphyton, Girvanella and Renalcis); Markello and
Read 1982: Girvanella and Renalcis); and Antarctica (Rees et al. 1989).
Not until the 1980s was the Soviet work on systematics closely scrutinized.
Riding and Voronova (1982b,c, 1984) discussed the affinities of genera such as
Angulocellularia and Tubomorphophyton, and concluded (Riding and Voro-
nova 1982a), in agreement with Luchinina (1975), that most Early Cambrian
"algae" were actually likely to be cyanobacteria and reflected enhanced
calcification relative to the Late Precambrian. Riding (1982) suggested that this
was due to a change in oceanic carbonate chemistry. Pratt and James (1982) and
Pratt (1984) proposed that Epiphyton and Renalcis might be diagenetic products
of coccoid cyanobacteria. Riding and Voronova (1985) reviewed the taxonomy
of Cambrian calcareous algae and cyanobacteria from the point of view of
morphology rather than affinity, and recognized 6 major groups with 19 genus
groups, and discussed the morphological series which exist in several ofthe more
common of these.
314 R.Riding
In the book Fossil calcareous algae (Ch uvashov et al. 1987) Luchinina (1987,
pp. 12-38) provides a classification of calcified cyanobacteria in which notable
Cambrian genera are included (pp. 23-24). However, the section by Chuvashov
(pp. lO9-139) includes Gordonophyton and Tubomorphophyton and some other
Epiphyton-like genera in the rhodophytes, which follows Korde's (1973)
classification, and Batenevia and Botominella (which show some resemblances
to Girvanella) are figured as early representatives of solenoporaceans
(Chuvashov et al. 1987, Fig. IV.22). Clearly, discussion of the systematics of
these fossils has not finished. Rowland and Gangloff (1988), in a review of Lower
Cambrian reefs, suggested that Renalcis and Epiphyton were facultative het-
erotrophs which could live either heterotrophically or autotrophically and that
this accounted for their occurrence in reef cavities. Luchinina and Tikhomirova
(1988) show SEM photographs of Epiphyton, Renalcis and Subtif/oria. It is this
loind of new information, combined with Recent analogues, which offers the
most promising way to clarify the affinities of Cambrian calcareous algae and
cyanobacteria.
3 Systematics
Ideally, morphology should indicate affinity, and both should be reflected in

classifications of these fossils. However, the interaction of these two approaches
has resulted in muddled schemes. Two reasons stand out: firstly, parallelism of
organization in these organisms results in misleading similarities between quite
different groups; secondly, workers often appear to have sought to clarify
taxonomic difficulties by accepting likely explanations of affinity for a few
genera and then attempting to pull the rest into conformity. As a result, these
fossils are notorious for changes of attributions of affinity. Just a slight perusal
of the literature reveals how Epiphyton, Renalcis and other genera have been
switched about between red algae and cyanobacteria. Proaulopora even has the
distinction of being moved between greens (Vologdin 1962a, p. 546), reds
(Korde 1973, p. 224) and cyanobacteria ~Luchinina in Chuvashov et al. 1987,
p.28).
Important classificatory schemes are compared in Table 2, which shows
generic groups, but for clarity omits orders, families, etc. into which they have
been placed. In some instances in Table 2 the sequence of groupings has been
changed to facilitate overall comparison of the schemes, but on the whole the
classifications are shown as they are published. Johnson's (1966) classification is
largely based upon the work of Korde (1961) and Vologdin (1962a) and,
following the Russian authors, places genera such as Renalcis, Angulocellularia,
Epiphyton and Bija in the red algae, and Praulopora and Seletonella in the
greens. Vologdin (1962a, p. 309) placed Girvanella in the cyanobacteria, but
Johnson (1966) preferred to also keep a chloroph yte option open for poro-
stromates. Korde's (1973) classification was more comprehensive, notably in
including spherical objects such as Bogutschania, Foninia, and A canthina. Some
ofthese were placed in cyanobacteria, but most of the genera she described were
put in the red algae. This included not only Girvanella-like taxa such as
Nicholsonia and Botominella, but also Proaulopora.
Riding and Voronova (1985) temporarily abandoned affinity in an attempt
to reappraise the classification of these fossils on grounds of morphology alone.
The result was recognition of six principal groups: (1) spherical (calcispheres,
Uranovia, etc.); (2) botryoidal (Renalcis, Tarthinia, etc.); (3) dendritic
(Angulocellularia, Chabakovia, Epiphyton, etc.); (4) tubiform, a) fans of tubes
(Botomaella, Hedstroemia, Solenopora, etc.), b) tangled, coiled, and straight
tubes (Girvanella, Obruchevella, Proaulopora, etc.); (5) tuberous (Seletonella,
etc.); and (6) cups (radiocyathids). The cup group is omitted from Tables 2
and 3.
Chuvashov et al. (1987), in a comprehensive review of Phanerozoic fossil
calcareous algae, attempted again to classify according to affinities, with the
work divided between Chuvashov (red algae), Luchinina (cyanobacteria) and
Shuysky (green algae). Shuysky safely places the Seletonellaceae in the greens,
but there appears to be some conflict in the areas of reds and cyanobacteria.
These of course are the two groups that previous workers have chiefly used in
attempting to place these fossils systematically. The main difficulties in
Ch uvashov et al. 's (1987) treatment of Cambrian genera concern the following:
1. Epiphyton group; Luchinina (in Chuvashov et al. 1987, p. 23) places Epi-
phyton in the cyanobacteria, but Chuvashov (in Chuvashov et al. 1987,
p. 127) puts the very similar genera Gordonophyton, Tubomorphophyton,
Epiphytonoides, and Filaria, in the red algae.
2. Girvanella group; Luchinina places Girvanella in the cyanobacteria, but
Chuvashov appears to place Nicholsonia, a junior synonym of Girvanella,
along with other kenellaceans in the red algae.
3. Batenevia group; Luchinina places Batenevia and Subtiftoria in the cyano-
bacteria, whereas Chuvashov (in Chuvashov et al. 1987, Fig. 22, p. 139)
places Batenevia and Botominella (junior synonym of Subtiftoria) in an
evolutionary line with solenoporacean supposed red algae.
Despite these continuing problems of affinity (Table 4), which are only
likely to be finally resolved by discovery of modern analogues, the comparison
of classifications in Table 2 reveals an emerging consensus regarding major
groupings. In this paper I follow Riding and Voronova's (1985) scheme based on
morphological details, but here I add comments on possible affinities (Table 3).
Omitted from detailed consideration here are the spherical and cup-like groups
of Riding and Voronova (1985), neither of which are likely to include well-
defined algae or cyanobacteria. Of the members of the spherical group,
Uranovia are small (30 /tm) micritic or hollow bodies within stromatolites;
Palaeomicrocystis is larger (250 /tm) and micritic with a clotted microfabric;
Bogutschania is a small (500 /tm) ooid; Foninia is large (2.75 mm) and may be
an ooid; some specimens attributed to Acanthina multiformis (e.g. Korde 1973,
PI. V) appear to be small recrystallized ooids; Marenita is micritic 50-300-/tm
diameter objects, which may be part of ooids or stromatolite fabrics. In contrast,
the cup-like group contains radiocyathids which are large, well-defined fossils
Table 2. Comparison of classification schemes. showing generic groupingsa
RIDING
KORDE 1973 I & 1985 G CHUVASHOV etai. 1987 THIS PAPER
I; ; / ; ; ; ; ; J ; ; ; ; ; ; ; ; ; ; .. _ _ _ _ _ _ _ VORONOVA }-U~~NINA pp 23-24 CHUVASHOV pp Lv,",
, SPHERICAL - - - - - .. ; ; ; ; ; ; ; ; ; ; A; ; ; 7 } ; ; ; ; ; 71 w
I 0-
Foninia I
• Marenita •
• Globuloella • Globuloella
• BOTRYOIDAL' Palaeomlerocystis
x
x
x
x Razumovskia 1
x ~N~~~~X.H.X~~ ~~~l~l~ : ~~~~iV~~,u,afla ~ m:~:'~~~la
x Pseudoanthos 7 x x)( ... Gemma I
x
Acanthina 7 ) ( I -: Pseudoanthos 7 J I
~,
./; /; /; /; /; /; /; /; /; /; 1b/; /; /; /; /; /; /; /; /; /; /1j Mueilina 7 97777777777V7777777777x Aeanthln~ 7 17?V/???/?'1
~ Angulocellularia Angulocellularia Kundatia 7 x DENDRITIC .:: MUelllna "';r,:
..."""..'"....
"" -'""...~""..,.~:..,~-""'-"-I
I x Angulocellulana " Kundatla 7 Angulocellularla
x Cornutula 8 ~ - - - - - - _.J
x
x Razumovskia 1 x ~~??
Kordephyton x Kordephyton Ix ~
. Kadvoya 9 x (x,.
x Chabakovla Chabakovia Serligia 9 x I X - :7
x Taninia . = Taninia .:. x ?
x Chabakovla x Chabakovia xl I
= . n" . . Chomustachia x Chomustachia -:: ) I
x sa,.a 18 SaJanla Parachabakov ia x Parachabakovia I< X? ?
~ Eplphyton Epiphyton A~gaina KO~ilophyton KOriiOPhyton': :.
Eplphyton
x x
=Eplphyton Epiphyton f' Xi Epiphyton
iI
x Cambrtna l O x Cambrina 10 i: X- Korilophyton
~ Gordonophyton : Gordonophyton Gordonophyton x? Gordonophyton ?
x Tubomorphophyton x Tubomorphophyton C TU?OmOrphOPhyton:- Tubomorphophyton I
x x t< Eplphytonoides t6 ~ I
x x l Filaria 16 x? ?
x Dasycirriphycus 2 Dasycirriphycus 2 x TUBIFORMI BIJa 3 t< - - - - - -
~ --;
x Razumovskia 1 x BaJanophyton I I .,;t. XI Botomaella
x x Flstulella t2 (pars) Zaganolomia 15 .. ~ Hedstroemia3 I
x Bija 3 Bija 3 Bija 3 = Hedstroemla 3 L
=z:j;;z;;;;,r::%z::::j'~4 >cI Bajanophyton I
x Solenopora x S~lenopora notomaella t< S o l e n o p o r a : ' I
fJ'~x.A~:,x~~x~x~~~~~~~~~~J''':o~><O~ :I B~Ja3 I Hedslroemia 3 r XI
~. I x Glfvanella TUBIFORM Girvanella .. N'1C h i ' 11 x-· •
I Girvanella I§ Girvanella oXNichoisonia11 x Nicholsonia 11 III K ~Isonla
It XI Glrvanella I
• !=Kenella ~ ~~I~~Y~"e"at3 • Razumovskia 1 t< ;:. Batenevia
F~s~~le~la12 I
! Nephelostroma 4 !x Fistulella 12 xSubtifloria. 1 • Obruehevella 1:. ,.. I
• Nubeeularites 4 'x Botominella 13 x BRaZtUmovlslkla K • Batenevia • Botomlnella 13 xl Subtlflofla t3 I
/ ~ oX x 0 omae a enella S b "fl . -X .
~~~~
~
~ ~L? ?X Batinevia 14 x Obruchevella Cavifera I u II orla I< Batenevla :. Obruchevella I
000 0 oL 000000000000 OX x Glomovertella I JIC._;::-_..,-_ _ _ _ _>L- ---.II
o OX x Proaulopora P.roaulopora • Proaulopora ..- - - - - - - - .,
o Proaulopora P I 0" Proaulopora x Amganella 6 ' It )(I ?
o roau opora OX Volo din ella 6 x Palaeoniles I Vologdinella 6 XI Proaulopora i
00 0" 9 Tubophyllum
x Tubophyllum Ix "? ? ?"
0.00 o"o"b"o"o"o"o"o"o"o"o"cI'o"~gdinella6 ~~~~--~-~Mxxxxxxo"o"ox~? ? ?-.J ;:0
Amgaella Amgaella 6 o
0 Amgaella TUBEROUS 0 0 00 SHUYoS'ky p~ 603~6o/ 00 00 0 "7'e'0'0'e'?b" ~ 0.:
o Cambroporella Cambroporella 0 0 Amgaella 00 0 5'
~
• • o MeJerelia MeJerelia ~ MeJerella ~ C_ambroporella o? ?
o Seletonella Seletonella 0 Seletonella 0 Mejerella :
~
~
~
~ ~ Sibenelia 5 Yakutma 5 ~ 0 Seletonella 00 Seletonella 0
//~ Vologdlnella 6 0 0 if! Yakutina ?
- - - - - - _ _ _ ~ _ _ _ _ _ ~nn_~_nnnnnnn ~nnnnnn~nnnnnnnnnnn~nnnnnnn~~nn?nn~n?nnn~
Table 3. Morphological groups of Cambrian calcareous cyanobacteria and algae (adapted from
Riding and Voronova 1985), with possible affinities added
Morphology Group Other genera Affinities
Botryoidal Renalcis Chabakovia, Probably

Chomustachia, Gemma, cyanobacteria
Parachabakovia,
Taninia, Tarthinia
Bushlike A ngulocellularia Filamentous
cyanobacteria
such as
oscillatoriaceans
Dendritic Epiphyton Gordonophyton, Probably
Korilophyton, cyanobacteria,
Tubomorphophytorf possibly
stigonema taleans
Tubiform branched Botomaella Bajanophyton, Bija, Filamentous
Hedstroemia, cyanobacteria
Kenella such as
rivularieaceans
tangled Girvanella Batenevia, Filamentous
Nicholsonia, cyanobacteria
Subtifloria such as
coiled Obruchevella Glomovertella, oscillatoriaceans
Lukaschevella
straight Proaulopora Possibly
filamentous
cyanobacteria
Tuberous Seletonella Yakutina ?Dasycladaleans
a And probable synonyms of Epiphyton (see No. 10 of Footnote, Table 2) .
....
a Higher taxa are omitted for clarity; as are non-calcified and obscure genera and genera based on
stromatolitic microfabrics. 1 There is some doubt concerning diagnostic features of Razumovskia
(see Riding and Voronova 1985, p 65), and consequently it is mobile between classifications. 2
Dasycirriphycus somewhat resembles Renalcis. 3 Bija may be ajunior synonym of Hedstroemia. 4
These genera are morphologically misplaced here, Nephelostroma is a junior synonym of Renalcis.
Nubecularites may be a catagraph (see Riding and Voronova 1985, p 60) although it has also been
tentatively placed in synonymy with Renalcis (Reitlinger 1959, p \3).5 Yakutina replaces Siberiella
Korde 1957 (non Radchenko 1955). 6 Vologdinella and Amganella are junior synonyms of
Proaulopora (see Voronova 1976, p 79). 7 Pseudoanthos and Acanthina are irregular wispy micritic
patches; A. multiformis appears to be part of spherical objects resembling ooids (see Korde 1973, PI.
5). Mucilina is somewhat similar. Kundatia is stick-like with traces of rectangular to ovoid chambers.
8 Cornutula is ooid-like and about 350 /Lm in diameter. 9 Kadvoya and Serligia are apparently
tubular and are not similar to other members of the Cambrinales. 10 Described by Korde (1973),
but not listed here, are the following cambrinalean genera, all of which, together with Cambrina, are
likely to be junior synonyms of Epiphyton: Kyzassia, Filaria, Umbellula, Sajania, Vologdinia,
Potentillina, Erbina, Tomentula, Sporinula and Epiphytonoides.11 Nicholsonia is a junior synonym
of Girvanella (see Danielli 1981, p 96).12 The type-species of Fistulella belongs with Botomaella (see
Riding and Voronova 1985, p 63).13 Botominella is a junior synonym of Subtifloria (see Luchinina
1975, p 28). 14 Correct spelling is Batenevia (see Voronova 1976, p 68). 15 Zaganolomia resembles
Praesolenopora Tsao. 16 Epiphytonoides and Filaria are likely to be junior synonyms of Epiphyton.
Also see 10. xxx Encloses red algae; dashes enclose cyanobacteria; circles enclose green algae.
Combinations of these symbols, and question marks, reflect the views of the original author(s). Note
that Riding and Voronova (1985) gave no affinities. See Table 4 for a clearer outline of attributed
affinities.
Table4. Affinities attributed to generic groups shown in the comparison of classifications (Table 2). Riding and Voronova (1985)
gave no affinitieS'
RIDING ....
VOLOGDIN 1962 I JOHNSON 1966 KORDE 1973 & 1985 C H UVASHOV et al. 1987
THIS PAPER ;;
VORONOVA
SPHERICAL
CYAN a
BOTRYOIDAL
DENDRITIC
I CVAN
RED
CYAN I ?CYAN
RED
I CYAN
CYAN GREEN l
OR
CYAN
---J
TUBEROUS SHUYSKY pp 63-64
GREEN
?"
GREEN I? GREEN I ~
er
(JQ
which show some similarities with both receptaculitids and archaeocyaths (see
Riding and Voronova 1985, p. 75). They do not closely resemble any ofthe other
groups considered here. They are probably not algae.
4 Morphological Series
Riding and Voronova (1985), in describing major morphological groups of

Cambrian calcareous algae, recognized the presence of morphological series
within a number of them: "In some cases the series are disjunct; in others they
are transitional and present serious difficulties in establishing diagnoses and
separating taxa. The presence of transitional series shows that confusion in
classification of these fossils is not due solely to subjective argument concerning
affinity" (Riding and Voronova 1985, p. 57).
Series are most clearly seen in the Renalcis, Angulocellularia, Epiphyton,
Botomaella, and Girvanella groups. The Girvanella-Obruchevella-Proaulopora
series of unbranched or rarely branched tubiform fossils is discontinuous and
does not present serious taxonomic difficulties. The Botomaella-Hedstroemia-
Solenopora series (Riding and Voronova 1985, p. 71, pp. 74-75) is continuous
and therefore more difficult, but elucidation of problems regarding Solenopora
and its allies will greatly reduce these problems. As has already been noted
(Introduction, above), reported Cambrian Solenopora are not congeneric with
Solenopora Dybowski.
The Renalcis-Epiphyton ring of Riding and Voronova (1985, p.71,
pp. 73-74) can be simplified for Cambrian specimens (Devonian material may
be more difficult) into an Epiphyton series (Korilophyton-Epiphyton-
Tubomorphophyton-Gordonophyton) which is partially disjunct, and a Renal-
cis-Angulocellularia series (Fig. 3). The latter is the major problem for taxonomy
because it shows apparently clear transition along its entire length. The
significance of this for interpretations of affinity is still unclear. Saltovskaya
(1975) has regarded Renalcis and Epiphyton as members of a single group, and
Pratt (1982) has argued that these genera, and their variants, are products of
diagenetic calcification of coccoid cyanobacteria. But A ngulocellularia com-
pares closely with calcified sheaths of modern Schizothrix (Riding and Vo-
ronova 1982b) which is a filamentous cyanobacterium. And yet it grades via
Tarthinia to Renalcis, which may well be coccoid in origin (Hofmann 1975).
Recognition of morphological series does not solve these difficulties, but it does
help us to define the problems more precisely .
...
• C YA N Cyanobacteria. Note that the most consistent attribution of affinity has been for genera
regarded as green algae. In contrast, many of the Cambrian genera regarded as red algae by
Vologdin (1962a) and Korde (1973), are now considered more likely to be cyanobacteria.
320 R. Riding
Fig. 3. Morphological series shown by the A ngulocellularia, Epiphyton and Renalcis groups (based
on Riding and Voronova 1985, Fig. 16). 1 Angulocellularia anisotoma Vologdin (Riding and
Voronova 1982a, Fig. 2c); 2 Korilophyton inopinatum Voronova (Voronova 1976, PI. 17, Fig. 5);
3 Epiphyton altum Korde (Korde 1961, PI. 17, Fig.4); 4 Tubomorpophyton botomense Korde (Riding
and Voronova 1982b, PI. 96, Fig. 5) ; 5 Gordonophyton distinctum Korde (Korde 1973, PI. 40, Fig.4a);
6 Taninia tomentosa Korde 1973, PI. 25, Fig. 3a); 7 Renalcis seriata Korde (Korde 1961, PI. 4, Fig.
3); 8 Renalcis erbinatus Korde (closely resembles R. granosus Vologdin) (Korde 1973, PI. 2, Fig. 4) ;
9 Gemma inclusa Luchinina (Riding and Voronova 1985, Fig. JOc); 10 Tarthiniagelatinosus(Korde)
(Korde 1961 , PI. 4, Fig. 5). All specimens are drawn at the same magnification. Angulocellularia (1)
is 750 p,m in longest dimension
5 Renalcis Group
Vologdin (1932) described Renalcis from limestone believed to be of Middle

Cambrian age in the Altai Mountains. The type locality is the Kandalep River,
south-east of Krasnoyarsk. The type species, R. granosus (Fig. 4f) shows clusters
of thick-walled botryoidal chambers with the larger ones enveloping smaller
ones. As with so many Soviet Cambrian fossils, the type-specimen appears to be
lost and it is unlikely that the type-locality can be accurately pinpointed.
Consequently, the original illustration by Vologdin (1932, PI. I, Fig. I) is all that
we have , apart from the description, to diagnose the genus. Renalcis is one ofthe
best known of Cambrian calcified cyanobacteria and algae. It is very common
in Lower Cambrian bioherms, often in association with Epiphyton and ar-
chaeocyaths. Indeed, Vologdin (1932, p. 79) states "The limestones from the
rivers Kandalep, Karagan and Lebed often contain in association with the
Archaeocyathinae considerable quantities, nearly equalling those of Epiphyton,
of small vesicular remains referred by the author to a special genus of algae,
Renalcis gen. nov.".
Fig.4a-h. Original illustrations of some Cambrian calcified cyanobacteria and algae. a Subtifloria
delicata Maslov (1956, PI. XXVII, Fig. 4). The genotype, Lower Cambrian, Tuva. x27; b An-
gulocellularia anisoloma Vologdin (I 962a, PI. 2, Fig. 2). The genotype, Lower Cambrian, River
Yangud , Baikal. x 15; c Tarthinia rOlUnda Drosdova (1980, PI. III, Fig. 5). Lower Cambrian, western
Mongolia. x40; d BOlOmaella zelenovii Korde (1961, PI. I, Fig. 7). Lower Cambrian, River Botoma.
x47; e Taninia tomentosa Korde (1973, PI. XXV, Fig. 3a). The genotype, Lower Cambrian,
Kuznetskii Alatau. x50; f Renalcis granosus Vologdin (1932, PI. I, Fig. I). The genotype. Middle
Cambrian. River Kandalep, Altai Mountains. x7; g Gordonophylon dislinclUm Korde (1973 , PI. XL,
Fig. 4a). The genotype. Lower Cambrian, Tuva. x36; h Korilophyton inopinatum Voronova (1976,
PI. XVII. Fig. 6). Nemakit-Daldyn Horizon, River Kotui. x36
322 R. Riding
This association between Renalcis, Epiphyton and archaeocyaths has often

been called the algal-archaeocyath association, even though it is more likely that
both Renalcis and Epiphyton are cyanobacteria ra ther than algae. The confusion
arises because cyanobacteria have hitherto long been termed blue-green algae.
Although the illustration of the type-specimen does not show it, Renalcis often
has short irregular branches composed of the broad, inflated, thick-walled
chambers. A number of somewhat similar genera occur higher in the Palaeozoic,
particularly in the Devonian, which have been confused with Renalcis. These
include Izhella Antropov and Shuguria Antropov. Riding and Brasier (1975), in
suggesting a foraminiferal affinity for Renalcis, were actually describing a
Devonian specimen of Shuguria. Differences in wall thickness and morphology
distinguish these genera from Renalcis: Izhella has very thick walls with internal
V-shaped clefts; Shuguria has thin walls. In addition, Shuguria has more
numerous chambers than either Izhella or Renalcis. Luchinina in Chuvashov et
al. (1987, p. 37) shows Shuguria ranging from the Middle Cambrian to Car-
boniferous, although I personally know of no firm records of Shuguria (or
Izhella for that matter) prior to the Devonian.
Nevertheless, other Renalcis group genera such as Taninia, Tarthinia and
Chabakovia do occur in the Cambrian, and Taninia closely resembles Shuguria
as do two other Cambrian genera: Chomustachia and Parachabakovia. The
internal taxonomy of this group requires refinement, but it forms a distinctive
class of chambered fossils with an overall botryoidal to stubbily branched
appearance. Two morphological series occur in relation to this group (Fig. 3):
(1) botryoidal fossils (e.g. R. granosus) grade towards branched fossils (e.g.
Taninia, Fig. 4e); (2) chambered fossils (the entire group) grade via Tarthinia
(Fig. 4c) towards solid micritic fossils (e.g. Angulocellularia, Fig. 4b). Tarthinia
includes T. gelatinosus, originally described by Korde (1961, p. 57) as a Renalcis
species (see Riding and Voronova 1984, p. 206), but the genus was described by
Drosdova (1975) from the Lower Cambrian of western Mongolia. Latham and
Riding (1988) recognized Tarthinia, together with Renalcis, forming the
macroscopic clots of thrombolites in probable Lower Cambrian of Morocco,
and this was the first report of Tarthinia outside Mongolia and Siberia.
A distinctive additional genus which can tentatively be assigned to the
Renalcis group is Gemma. In published reports it is only known from the remote
Anabar region of north-central Siberia (Luchinina in Zhuravleva et al. 1982,
p. 100) and from western Mongolia (Riding and Voronova 1985, Fig. lOc). It has
a very thick wall composed of numerous micritic spherules.
The affinities of Renalcis and similar genera are a perennial problem. The
simplicity, and variability, of form, coupled with relatively small size, suggest
cyanobacteria. Although Vologdin (1962a) and Korde (1973) placed Renalcis in
the Rhodophyta, Korde (1961, p. 55) did refer it to the cyanobacteria and in
Maslovand Korde (1963, p. 51) suggested a rivulariacean affinity. Saltovskaya
(1975), concentrating on Devonian material, also placed Renalcis, along with
Epiphyton, in the filamentous cyanobacteria (Hormogonophyceae), and Lu-
chinina (1975) working wi th Cam brian rna terial, also fa voured a cyanobacterial
affinity for Renalcis. Hofmann (1975) suggested that Renalcis represents
diagenetic calcification of chroococcalean colonies, an idea developed by

Poncet (1976; who actually illustrated Devonian Izhella) to include a rivula-
riacean affinity. Pratt (1984) also supported both a coccoid cyanobacterial and
a diagenetic origin for Renalcis (and also Epiphyton) but concentrated on
non-Cambrian material. Luchinina (in Chuvashov et al. 1987, p. 24) also
favours a chroococcalean affinity for Renalcis.
Overall, then, opinion supports a cyanobacterial (chroococcalean or
rivulariacean) affinity for Renalcis, and Hofmann's (1975) suggestion of periph-
eral calcification of colonies most plausibly accounts for the appearance of
this enigmatic fossil. The problem that remains is that Renalcis and its allies
appear to have a stratigraphic range limited to the Palaeozoic. Is this significant,
and do modern analogues exist for Renalcis? These questions need to be
addressed in future research.
6 Anguiocellularia Group
This genus, characterized by being bushy in form and densely micritic in

structure, is much more common than published reports indicate. Vologdin
(1962a, p. 484; Fig. 4b) described it from the Lower Cambrian of the River
Yangud, in the Baikal region. Riding and Voronova (1982b) emended the
diagnosis to exclude chambers (which are not visible in Vologdin's photograph
of the type-specimen although he gives a drawing showing them (Vologdin
1962a, Fig. 4)), and described a Recent analogue of A ngulocellularia in calcified
sheaths of the oscillatoriacean Schizothrix calcicola.
Angulocellularia shows transition towards Tarthinia in the Renalcis group,
and towards Epiphyton. For example, Korilophyton angus turn (Voronova 1976,
p. 84), which shows some similarities with Epiphyton, has been placed in
A ngulocellularia (Riding and Voronova 1984, p. 206). Korilophyton itself, based
on Epiphyton inopinaturn, remains a distinct genus within the Epiphyton group.
However, it is towards Korilophyton, which shows short finger-like branches
but is less dendritic than Epiphyton, that Angulocellularia shows transition
(Fig. 3).
7 Epipbyton Group
Epiphyton and its allies constitute perhaps the most important single group of
Cambrian calcareous algae or cyanobacteria. Epiphyton is so named because
Bornemann (1886) found it epiphytically attached to archaeocyaths. Epiphyton
and its relatives, and also Renalcis, can be so abundant that despite their
individually small size they form macroscopic dendritic masses (dendrolite,
Riding 1988) which may dominate Cambrian reeffabrics.
There are too many species of Epiphyton, most of them described by Korde
(1961) from the Lower Cambrian of the middle Lena River and the Amga River
south-west ofYakutsk, as well as from the Kuznetsky Alatau mountains midway
324 R. Riding
between Novosibirsk and Krasnoyarsk. The species are difficult to distinguish

from one another because they are often based on obscure or subtle details of
microstructure or branching angle. There has also been a proliferation ofgenera
within the group, again by Korde (1973): Cambrina, Kyzassia, Filaria, Um-
bel/ula, Sajania, Vologdinia, Potentillina, Erbina, Tomentula, Sporinula and
Epiphytonoides are all likely to be junior synonyms of Epiphyton. These are all
based on Lower Cambrian material, and are all from the Altai Mountains
(except for Erbina from Kuznetsky Alatau); most (the exceptions are Kyzassia,
Potentillina and Epiphytonoides) being from the River Bazaikha in the eastern
Altai.
Nevertheless, it is not so difficult to recognize several important genera.
These are Epiphyton (solid micritic, dendritic), Korilophyton (solid micritic, but
with short branches), Gordonophyton (large size, regular discoidal or lenticular
transverse chamberlike gaps in the filaments, and often showing gradual distal
expansion of filaments), and Tubomorphophyton (tubiform or chambered). Of
these Epiphyton and Gordonophyton appear to be most widely distributed.
Gordonophyton was first described as Epiphyton by Gordon (1920) from the
Cambrian of Antarctica, and then created by Korde (1973, p. 198) using
material (G. distinctum) from Tuva (Fig. 4g). It also occurs in the Upper
Cambrian of Newfoundland (Pratt 1984, Fig. 5b). Korilophyton and
Tubomorphophyton are not yet known from outside Siberia and Mongolia (see
Riding and Voronova 1984). Korilophyton (Fig. 4h) is based on material from
the Koril Member near the base of the Tommotian in Anabar, which was
originally referred to Epiphyton (see Voronova 1976, pp. 83-84). Tubomor-
phophyton too, is based on the originally Epiphyton species, E. botomense, from
the middle Lena River (Korde 1973, p. 204).
The idea that Epiphyton is a red alga stems from Korde's (1959) report of
cells preserved within the filaments, but this observation has never been
substantiated. Earlier workers suggested a chlorophyte (Bornemann 1886) or
cyanophyte (Pia 1927, p. 39) affinity. Saltovskaya (1975) and Luchinina (1975)
both regarded Epiphyton as a filamentous cyanobacterium. Riding and Vo-
ronova (1982c) compared Tubomorphophyton with both red algae and cyano-
bacteria and concluded that a cyanobacterial affinity is the more likely of the
two and that some stigonemataleans, in particular, closely resemble epiphy-
taceans. Pratt (1984) suggested that Epiphyton and Renalcis are diagenetic
microfossils produced by coccoid cyanobacteria. However, the filamentous
form of Epiphyton, which is often very well-defined, does not seem to be
consistent with such an origin. In particular, it has not been explained how
coccoid cells would create such a regularly branched dendritic pattern of
filaments, nor how diagenetic calcification (as opposed to calcification during
life) would preserve the morphology so well. Luchinina (in Chuvashov et al.
1987, p. 23) places Epiphyton and Korilophyton in filamentous cyanobacteria,
but Ch uvashov (in Ch uvashov et al. 1987, p. 127) places Tubomorphophyton and
Gordonophyton in the Rhodophyta. Recent analogues are required to resolve
these continuing uncertainties concerning the affinities of this important group
of Cam brian microfossils.
8 Botomaella Group
Botomaella (Fig. 4d), named by Korde (l958b) after the Botoma River, and
Kenella have delicate tubular branched filaments characteristically arranged in
fan-like radiating clusters. Hedstroemia (originally described from the Silurian
by Rothpletz 1913) is similar in general form, but has filaments which are both
wider and more variable in width. In the Cambrian, Botomaella and Hed-
stroemia typically occur as small isolated specimens and do not appear to be
associated with reefs. Whereas Botomaella has been reported from several areas
of central Siberia (see Riding and Voronova 1984), and also from western
Mongolia (Drosdova 1980, p. 18), Hedstroemia has hitherto mainly been de-
scribed in the Cambrian as Bija or Rothpletzella Wood. Bija is difficult to clarify
because the original material was only illustrated in transverse section
(Vologdin 1932, Fig. II), but it could be a junior synonym of Hedstroemia.
Rothpletzella igarcaensis and R. flabellata described by Voronova (1976, p. 82)
are not congeneric with Rothpletzella Wood, but resemble irregular Hed-
stroemia (see Riding and Voronova 1984, p. 206). Bajanophyton, described by
Drosdova (1980, p. 43) from the Lower Cambrian of western Mongolia, appears
to be intermediate between Botomaella and Hedstroemia.
Despite these confusions, the affinities of Botomaella and Hedstroemia can
be clarified by comparison with Recent calcified cyanobacteria. These modern
analogues (Riding, unpublished) show that the two genera are closely related,
both being very similar to species of Rivularia (Roth) Agardh. Consequently,
and rather surprisingly, we have an example here of species within a single
extant genus being more readily separated as fossils into two morphogenera.
Normal expectation would be the reverse: that fossil taxa tend to combine
several extant ones, as is prooably the case with Girvanella (Riding 1977, p. 43).
9 Girvanella Group
Girvanella is probably the best known, by name, of all these Cambrian fossils,
mainly because of its long post-Cambrian range. But it is also a very widely
reported genus within the Cambrian, particularly so in North America (see Sect.
2), although it figures less prominently in Soviet work. Originally described from
the Ordovician of Scotland by Nicholson and Etheridge (1878), Girvanella
characteristically occurs as loosely or densely tangled masses of fairly thin-
walled unbranched tubes. It also commonly forms the cortex in oncoids and
occurs as crusts within reefs.
Korde (1973, p. 212) distinguished Nicholsonia from Girvanella and named
it after H.A. Nicholson using Lower Cambrian specimens from Tuva. But
Danielli (1981, p. 96) placed it in junior synonymy with Girvanella. However, it
should be noted that one of Korde's (1973) Nicholsonia species, N. grandis, has
tube diameters of 50 /lm which is large by Girvanella standards.
Two genera which can be included in the Girvanella group are Batenevia
(original spelling Batinevia, but see Voronova 1976, p. 68) and Subtifloria (Fig.
326 R.Riding
4a) (senior synonym of Botominella, see Luchinina 1975, p. 28). In contrast to

Girvanella their tubes, although similar in size to Girvanella and also un-
branched, are aligned in thick cable-like strands, and Batenevia has ovoid spaces
within the strands. Danielli (1981, p. 96) placed Batenevia and Botominella in
synonymy with Girvanella, but their distinctive arrangement of tubes make it
useful to continue to recognize them as separate genera.
Another distinctive variant of gross morphology within this group is the
development of delicate, wispy prostrate threads which turn upwards at one end
and which may be Razumovskia (see Riding and Voronova 1985, p. 65).
Cambrian Girvanella has been regarded as a cyanobacterium (Vologdin
1962a), a cyanobacterium or chlorophyte (Johnson 1966) and, in the case of
Nicholsonia, a rhodophyte (Korde 1973) (see Table 4). However, a Recent
analogue has been reported in the calcified sheaths of Plectonema from fresh-
water pools in Aldabra Atoll (Riding 1977), and it is likely that members of the
Girvanella group as a whole have an oscillatoriacean affinity.
10 Obruchevella Group
Another group of tubiform microfossils, whose dimensions suggest that they

could be cyanobacterial sheaths, exhibits coiling. Typical of these is Obru-
chevella which Reitlinger (1948) originally described as a foraminifer from the
Lower Cambrian of the River Sinyaya, tributary to the middle Lena, although
later (Reitlinger 1959) she considered it to be of uncertain systematic position.
Elias (1954, p.52, footnote) regarded it as an alga. Similar genera include
Cavi[era and Glomovertella, also described by Reitlinger (1948), from the
middle Lena area, and Bostrychosaria, Flexurella, Kordeella, Lebedevaella,
Lukaschevella, and Tuvaellina from the Lower Cambrian of Tuva (Vologdin
1958). Further details are given in Loeblich and Tappan (1964, p. C787-C789)
who regarded the systematic position of these genera as uncertain. The spiral
arrangement of these fossils, particularly well seen in Obruchevella and Lu-
kaschevella, may be compared with filamentous cyanobacteria, including
rivulariaceae such as Dichothrix (see Rippka et al. 1979, Fig. 63) and oscilla-
toriaceans such as Spirulina (Luchinina 1975, p. II; Luchinina in Chuvashov et
al. 1987, Fig. 11.17, and see Fritsch 1959, p. 832).
Obruchevella is a calcareous fossil (see Reitlinger 1948; Riding and Vo-
ronova 1984, p. 206) but phosphatized (e.g. Spirellus Jiang, see Peel 1988) and
silicified fossils with similar appearance are also known and may occur in the
Precambrian, thus complicating evaluation of both the taxonomy and range of
these organisms.
11 Proaulopora Group
Proaulopora was originally described by Vologdin (in Krasnopeeva 1937) from

the Lower Cambrian of the western Altai Mountains. In contrast to the
Obruchevella and Girvanella groups, the tubes of Proaulopora do branch and are
straighter and stick-like in appearance. It is also larger, generally having a

diameter of 50-100 /Lm. A distinctive feature is the presence of whorl-like
external collars which extend from the outer surface at an acute angle to the
stem. Occasionally, Proaulopora occurs complete as branched masses (see
Korde 1973, PI. 53, Fig. 2), but more commonly it is broken into short fragments,
some of which have received separate names (e.g. Amganella). Luchinina (1975,
p.26) regarded Amganella, Palaeonites, Tubophyllum, and Vologdinella as
junior synonyms of Proaulopora. As with Obruchevella, Proaulopora does not
appear to be a very common fossil although it is locally conspicuous as fragments
in grainstones.
Proaulopora has the distinction, rare. even among Cambrian fossils, of
having been referred at various times to the green algae, red algae, and
cyanobacteria (Table 4). Luchinina (in Chuvashov et ai. 1987, p. 28) compares
it with Calothrix gypsophyla, a rivulariacean species with similar collar-like
projections ofthe sheath, although she notes that Proaulopora is more than twice
the diameter of Calothrix.
12 Seletonella Group (Fig. 5)
Seletonella is a rare fossil which may be a dasycladalean alga. It has a hollow,

tuberous, form with numerous closely packed branches. Seletonella is up to 20
mm in size and thus is relatively large when compared with most Cambrian
calcareous algae and cyanobacteria. Korde (l950a) described Seletonella and
Mejerella from the Upper Cambrian of northern Kazakhstan, south of Omsk.
Cambroporella comes from the Lower Cambrian of Tuva (Korde 1950b), and
Amgaella and Siberiella from the Middle Cambrian of the Amga River (Korde
1957b). The name Siberiella was preoccupied, and consequently it was renamed
Yakutina (Korde 1973). In 1971 Korde (quoted in Korde 1973, p. 239) united
these genera, which she regarded as dasycladaleans, in the Seletonellaceae.
They are all very rare. Korde (1961, p. 63) described a second Seletonella species,
S. grandis, from the Ordovician of northern Kazakhstan.
Elias (1954) reinterpreted Cambroporella as a probable bryozoan and
compared it with Coeloclema Nickles & Bassler. No one seems to have seriously
questioned the dasycladalean affinity postulated for the other genera, although
the polygonal elements in Amgaella (see Korde 1961, PI. 5, Fig. 2) seem atypical
and Mejerella (see Korde 1961, PI. 6, Fig. 6) has unusual long flexuous branches.
Bassoullet et ai. (1979, p. 432, Footnote) expressed reservations concerning the
dasycladalean nature of these Cambrian fossils but nevertheless included them
in the Seletonellaceae, which they regarded as one of the three families (the
others being Oasycladaceae and Aceta bulariaceae) of Oasycladales.
However, Seletonella and Yakutina may more plausibly be dasycladaleans.
These fossils from the Upper and Middle Cambrian respectively must therefore
be seriously considered to be not only the oldest known calcified dasyclads, but
also the oldest calcified green algae because Proaulopora seems unlikely to be a
chlorophyte, and at present there are no other serious contenders for this affinity
among Cambrian calcareous fossils.
328 R. Riding
Fig. 5. Selelonella group. Recon-

structions of S elelonella mira
Korde (55). Mejerella vamosa
Korde (56). Cambroporella luvensis
Korde (57). Yakutina aciculata
Korde (58), and Amgaella amgan-
ensis Korde (59). Reproduced
from Korde (1973 , p73)
13 Conclusions
There are some 20 well-defined, senior-synonym genera of Cambrian cal-

careous cyanobacteria and algae of a total of more than 70 which have been
described. Seventeen of these twenty have originally been recognized in the
USSR and most have yet to be reported from elsewhere. Affinities are a major
problem, but progress is being made in narrowing down the possibilities. At
present it seems likely that all the fossils considered here are either definitely or
probably cyanobacteria with the exception of Proaulopora, which may only
possibly be a cyanobacterium, and Seletonella and Yaku/ina which could well
be dasycladalean green algae. No calcified red algae are known from the
Cambrian. This apparent domination of the flora by cyanobacteria is a striking
feature, which is enhanced by the scarcity of algae. It seems reasonable to

conclude that calcareous algae, sensu stricto, are not at all common prior to the
Ordovician, and that the Cambrian, particularly the Lower Cambrian, repre-
sents the major time in the entire stratigraphic record of calcification of marine
cyanobacteria. This event may have been switched on by a change in seawater
chemistry at the end of the Precambrian which promoted CaC0 3 precipitation.
Information concerning the Lower Cambrian flora and the first appear-
ances of genera are quite well documented. The appearance of heavily calcified
cyanobacteria at the base of the Cambrian is no less abrupt than the synchronous
appearance of diverse invertebrate shelly fossils (Fig. 6). Much of this knowl-
edge comes from work in the Soviet Union on material from Central Siberia
and from the Urals and the Altai Mountains. All the major genera, with the
exception of Epiphyton and Girvanella, have been defined in the USSR. The
upward limits of the ranges of many genera are poorly known, and there is
relatively little detailed information concerning the Middle and Upper Cam-
4-------+-------+--------tt~ Renalcis
Tarthinia
Gemma
~ _ _________________________ ---------------- . . Angulocellularia
_______+ _______+ _______ +_~ Epiphyton
Korilophyton
Gordonophyton
Tubomorphophyton
Botomaella
______________ --------------- ----- ---------- . . Hedstroemla
-
I-------__ ~------ __
Bajanophyton
I _ - - - - - - _ _ i. . Girvanella
Batenevia
Subtifloria
-- ....-- Obruchevella
Proaulopora
Seletonella
Yakutina
I T I A I B'T
NO Lower Middle Upper
Fig. 6. Ranges of principal genera of Cambrian calcareous cyanobacteria and algae. Based on
Riding and Voronova (1984) and other references cited in the text. Note that the precise ranges of
Yakutina and Seletone/la, within the Middle and Upper Cambrian respectively, are not known.
ND Nemakit Daldyn; TTommotian; A Atdabanian; B Botomian; TToyonian
330 R.Riding
brian (Fig. 6). Cambrian floras have long been known in other parts of the world,
particularly Antarctica, Australia, Europe and North America, but none of
these has received the detailed study given to Soviet material. Considerable
progress, particularly with regard to palaeobiogeography, can be expected when
work in the rest of the world catches up with that in the Soviet Union.
The principal groups can be defined by simple morphological categories:
botryoidal (Renalcis group), dendritic (A ngulocellularia and Epiphyton groups),
tubiform (Botomaella, Girvanella, Obruchevella, and Proaulopora groups),
tuberous (Seletonella group). Discontinuous morphological series occur in
several groups and the Renalcis and Angulocellularia groups show continuous
series of morphological variation which make taxonomic definition difficult.
Individually, all of these fossils are essentially microscopic, with the exception
of the Seletonella group. However, members of the Renalcis, Angulocellularia
and Epiphyton groups commonly occur in concentrated masses, forming reefs in
association with archaeocyaths, which impart a macroscopic dendritic fabric to
the rock which can then be termed dendrolite. These reef-forming genera,
together with Girvanella, are the most widespread and abundant members ofthe
flora. Members of the Botomaella, Obruchevella and Proaulopora groups are less
common, and members ofthe Seletonella group are extremely rare, often only
being known by single occurrences. The Cambrian represents the first occur-
rence of a heavily calcified flora, indicating that biomineralization in algae and,
particularly, cyanobacteria was broadly synchronous with that in invertebrate
groups.
Acknowledgements. This research was strongly supported by The Royal Society and the USSR
Academy of Sciences, through their jointly operated exchange programme, and also by the Natural
Environment Research Council. I am very grateful to Nina Drosdova, K.B. Korde, Veronica
Luchinina and, particularly, Larisa Voronova for advice and discussion. A. Yu. Rozanov and L.
Tatarinov helped facilitate my visits to the Palaeontological Institute in Moscow, without which this
work would not have been accomplished.
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Zamarreno I (1977) Early Cambrian algal carbonates in southern Spain. In: Flugel E (ed) Fossil
algae. Springer, Berlin Heidelberg New York, pp 360-365
Zhuravleva IT, Meshkova NP, Luchinina VA. Kashina LN (1982) Late Precambrian and Early
Cambrian biofacies of the Anabarian Sea. In: Environment and Life in the geological past,
palaeolandscape and biofacies. USSR Acad Sci. Siberian Sec, Inst Geol Geophys. Novosibirsk,
Trans 510:74-103 (in Russian)
Chapter 17
Ordovician Algae and Global Tectonics
A. Roux l
Abstract
The evolution of faunal provinces during the Ordovician shows a progressive reduction of the
endemisms resulting from the progressive closing of the Proto-Atlantic ocean. The geographic and
stratigraphic distributions of some representative algal groups support this interpretation and
suggest some modifications for the proposed positions of the landmasses, particularly for North
America, the Baltic shield and the Kazakhstan-Tadjikhistan area. The algal migrations seem to be
related to the surface currents.
1 Introduction
Most palaeogeographic reconstructions are based on palaeomagnetic data and

faunal and floral data are relatively little used, at least not for worldwide
reconstructions.
Works using Palaeozoic algae for these research topics are very scarce and
limited to some titles concerning mainly the Upper Palaeozoic: Mamet (1972)
has shown the striking coincidence of the distribution of the Visean microflora
with the Tethyan belt. Mamet and Roux (1977) and Jansa et al. (1978) have
shown that the worldwide distribution of the Devonian-Carboniferous
microflora conflicts with the reconstructions which admit a physical contiguity
between North-America and Euroafrica, suggesting the presence of a Proto-
Atlantic ocean during Carboniferous. Roux (1979, 1986) and Mamet et al.
(1987) have shown that the Permo-Carboniferous microflora of the Sverdrup
Basin was not equatorial but warm temperate to subtropical, suggesting a
migration towards the north and a rotation of the North American block. Mamet
and Roux (1983) have confirmed the position of Australia during the Devonian
and Carboniferous previously proposed by Keppie (1977). Bildgen and Roux
(1 982a,b ) have shown that the hypothesis of an equatorial palaeoclimatic belt,
proposed on the basis of the distribution of karst bauxites, may be confirmed by
palaeobotanical data (see a synthesis in Roux 1985). Finally, new positions of
the North American block during the Lower Carboniferous and Permian have
been proposed (Roux 1986).
This study is a new attempt at reconstruction of the position of the
continental blocks (North America and Eurasia) during the Upper Ordovician
based on the stratigraphic and geographic distribution of algal microfloras.
151 Avenue de Verdun, 77610 Fontenay-Tresigny, France

336 A. Roux
2 Ordovician Palaeogeography and Faunal Data
According to Scotese et al. (1979), most of the continents occupied the same
latitudinal belt during the Late Cambrian and Middle Ordovician: during the
Upper Cambrian, Laurentia, Siberia and the southern portion of Gondwana
were in equatorial latitudes; China and Kazakhstania are also supposed to have
been within 30° of the equator. According to palaeomagnetic data, Baltica was
located at relatively high latitudes. As a consequence, the polar oceans were of
wider extent than at any subsequent times. From the Late Cambrian through
Middle Ordovician, Laurentia and Gondwana rotated counterclockwise, while
Baltica and Siberia moved northward.
The Early Ordovician coincides with a wide transgression over several
European and North African regions. The worldwide distribution of trilobites
and graptolites has shown a faunal provincialism suggesting that the extreme
northwestern part of Europe was part ofa Greenlando-Canadian plate, sepa-
rated from the Gondwano-European area by a Proto-Atlantic ocean called
Iapetus. More exactly, the northwestern margin oflapetus includes Greenland,
part of Newfoundland, northwestern Ireland, Scotland, and Norway. The
study of trilobite and brachiopod faunas has suggested the existence of two
provinces over Europe: a northern province (Baltic area) and a southern
province, separated by a wide and deep ocean (see a synthesis in Johnson 1977).
The definition of a Prototethyan realm during this time is problematic; a
Prototethys or a Prototethyan realm are intended in this work only to
differentiate the North-American realm.
Boucek (1972) has already proposed a probable position of the Lower
Ordovician (Tremadoc-Llandeilo) equator, which crossed the North American
block from north to south through Hudson Bay and Europe from Finland to the
Bay of Bengal. On the basis of that position of the equator, the author distin-
guished two provinces:
A Pacific tropical northern province including North America, western
Ireland, Norway (Trondheim area), the northern part of USSR (Taimyr), China
and New Zealand.
A southern province known in Europe, North Africa, the western part of
Asia and South America, and divided into:
A tropical southern or Anglo-Scandinavian subprovince including the
British Isles (western Ireland excepted), Scandinavia, the Baltic region, part of
the USSR (Ural region and Kazakhstan) and the western regions of South
America.
A temperate to cold, Mediterranean subprovince gradually linked with the
former and approximatively limited by Syria, Florida and Alabama, Wales,
Belgium, Bohemia, Poland, Morocco, Sahara and the eastern part of South
America.
During the Late Ordovician, a progressive homogeneization of the faunas
becomes evident in Europe; during the Lower Ordovician, evolution in each
province seems to have been independent; during the Upper Ordovician, the
Ordovician Algae and Global Tectonics 337
progressive merging of provincialism suggests the disappearance or a reduction

of physical barriers, that is, the closing of the Proto-Atlantic ocean.
The progressive closing of Iapetus accounts for more precise faunal ex-
changes between North America and Baltic Europe. The reduction of en-
demism seems to be the result offaunal (and probably algal) migrations from
North America to Europe.
On the basis of stratigraphic breaks observed in the Ordovician of northern
Estonia, Roomusoks (1972) has demonstrated several changes in the faunas: a
renewal of bryozoan, brachiopod and trilobite faunas appears at the Volk-
hov-Kunda Formations boundary (Lower Ordovician); among the immigrants
are some representatives of earlier faunal communities of Bohemia and North
America.
Near the end of the Middle Ordovician, numerous new genera of bra-
chiopods, bryozoans and trilobites appear in the Keila-Ondu Formation (Viru
Group) with a large number of immigrants from North America and Britain
among the brachiopods.
During the Upper Ordovician (Ashgillian, Pirgu Formation, Harju Group),
a large number offorms appears, including genera characteristic of Middle and
Upper Ordovician deposits of North America.
3 Distribution of Some Representative Algal Groups
Vermiporella
Figure I shows the worldwide distribution ofthe species of Vermiporella: during

the Middle Ordovician, the genus is only known in North America (Canadian
Arctic, Hudson Bay, Saint Lawrence Lowlands, New York State) and Scotland,
and is unknown in the 'Prototethyan' realm (Scotland belongs during this time
to the North American realm). The genus is represented by five species: V.
borealis, V. eisenacki, V. fragilis, V. inconstans and V. wesenbergensis. It is
interesting to note that these species were originally described from Upper
Ordovician deposits of the 'Prototethyan' realm.
During the Upper Ordovician, the genus is cosmopolitan, distributed from
Kazakhstan to the Canadian Arctic and represented by all the previously
mentioned species with, however, some species such as V. acerosa and V.
diffluens which are only known in Kazakhstan, and V. canadensis which is only
known in North-America (Newfoundland, Hudson Bay, Canadian Arctic).
Although the genus becomes cosmopolitan during the Upper Ordovician, the
former species are 'Prototethyan' and the latter, 'American'.
The presence of V. borealis, V. eisenacki, V. fragilis, V. inconstans and V.
wesenbergensis in North America and Scotland during the Middle Ordovician
and the presence of the same species in the 'Prototethyan' realm during the
Upper Ordovician, may be explained by migrations from North America and
especially the Saint Lawrence Lowlands and the New York State area to the
338 A. Roux
LOwER MIDDLE ORDOVICIAN UPPER ORDOVICIAN
:>
~ 9 :>
sa
~
:>:l
~ ~
~ C'i
H
C) H
~
t'l
H ~ F=
5
wesenbergensis ~
I
N
:>
acero~a_ ""
:I:
I'
I
U>
~ -l
1iffluens :>
z
I inconstat;ls - ~
U> wesenbergensis ..... c5
z
H
fragilis :>
fragilL ~
t;;l
A
I \
U>
;' \ I
borealis
~
V
eisenacki
borealis
I a
inconstans ~
:>
-<
I
I z
canadensis _
~
I
.
elsenac )
1 _
:>
::i
§
U>
~ ::l
:2 borealis ?'i ;::;
~ eisenacki (/)
~ >-I r
fragilis :>
Q inconstans ;j r
:I:
borealis_
I '"
:>
-<
c:
t:l
U>
fanadenSij - ~
I I
canadensis _
-H
:> £2
9'"
Z
". :>
;" t:l
00 wesenbergensis H
...., ()
S;
I I
Fig. 1. Geographic and stratigraphic distribution of species of the dasycladacean VermljJOrelia
'Prototethys' through Scotland and Norway, which belong to the North Ame-
rican realm (maps of Johnson 1977; Keppie 1977; Scotese et al. 1979).
These migrations were possible because of the relative proximity of the
North American and the Eurasian blocks, due to the more-or-less complete
closing ofthe Proto-Atlantic ocean during the end ofthe Middle Ordovician and
the Upper Ordovician.
Rhabdoporella
Figure 2 shows the worldwide distribution of the species of Rhabdoporella: the
genus is mainly Silurian, but occurs in Sweden and Anticosti Island during the
> ;g
~
~
~
Ul ~
~ i;Ii §
~
.....
f=: § ~ r'
.....
~ ~
>
'"
.....
00 :c
>
~ S+P
~
I
Ul
I;" "
~a
~
Ul S+P -- r -
....
""tl
I;" "
S+P ~
~
t;;1
1\
I \
\ I
."
00 . ---
." ~
>
-<
I;" "
..........~
......
§ ~ S+P .... -- --
- B
~
.....
8
~
.....
~
f;l
.....
~ S ....
Ul
~.
P F
Fig.2. Geographic and stratigraphic S+P '2 ~

g) '"
distribution of species of the dasy-
cladacean Rhabdopore/la I fi ~
340 A. Roux
Upper Ordovician (Ashgillian); the Norwegian references are not clear. During
the Silurian, the genus is cosmopolitan in North America (Anticosti Island, New
York State, Quebec, Saint Lawrence Lowlands) and in the 'Prototethys' (Wales,
Gotland, Sweden).
It is difficult to establish the pattern of migrations because of the simul-
taneous occurrence of the species R. stolleyi and R. pachyderma in Sweden and
Anticosti Island during the Upper Ordovician. According to other works
(Roomusoks 1972), the migrations may have taken place from Anticosti Island
to the 'Prototethys' through Sweden (and possibly Norway) during the end of
the Upper Ordovician or the basal part of the Lower Silurian, because of the
closing of the Proto-Atlantic ocean which had eliminated or reduced the
physical barrier of Iapetus.
Udoteaceae
The stratigraphic and geographic distributions of important genera of erect
udoteaceans are shown in Fig. 3: during the Middle Ordovician, the species
Dimorphosiphon rectangulare is known in the 'Prototethyan' and North Amer-
ican realms (Kazakhstan and Norway and the Saint Lawrence Lowlands
respectively). The same species occurs in Scotland and remains present in
Kazakhstan during the lower part of the Upper Ordovician (Caradocian).
Dimorphosiphon diadromum appears in Kazakhstan during the Upper Or-
dovician and occurs in northern Quebec during the Silurian as D. cf. diadromum.
Dimorphosiphon magnum seems to be cosmopolitan from Kazakhstan to
northern Quebec during the Silurian.
Among the genus Palaeoporella, the species P. opuntiformis appears in
Tadjikhistan during the upper part of the Upper Ordovician (Ashgillian) and
occurs in Anticosti Island (as P. cf. opuntiformis) during the Lower Silurian. P.
recta occurs in Kazakhstan during the lower part of the Upper Ordovician and
in northern Quebec during the Upper Silurian (as P. cf. P. recta). P. variabilis
occurs in the North American and 'Prototethyan' realms (Norway and Sweden)
during the Upper Ordovician.
The simultaneous occurrence of D. rectangulare in Kazakhstan and Scot-
land seems to be indicative of a relative proximity of the North American and
north Eurasian blocks during the lower part of the Upper Ordovician
(Caradocian). The same scheme is indicated by the simultaneous presence of P.
variabilis in Sweden and Norway.
The genera Dimorphosiphonoides and Lowvillia are strictly 'American' and
restricted to the Middle Ordovician.
Other Algae
Among the dasycladaceans, it is interesting to note that the genus Kazakh-
stanelia, originally described from the Ordovician of the USSR (central
Kazakhstan, Lake Ajakmalajsor and Bajanoul and Pribalkhash regions) is also
present in the Middle Ordovician of the Saint-Lawrence Lowlands (Quebec and
Ontario). Originally described from the Caradocian of Scotland, Intermurella
MIDDLE OROO. UPPER OROO. SILURIAN
r- Q .. r- ."
~'"
~ );
~
Cf)
:I: ~ c
0
§'"
..
z
~
H
z
0
[T1
~ "~ 8< § r-
'"z 5 ~
..::
z
>
;., ~ P. opuntiformis ...,,..
~ E
;;;
.
:I:
...,
H
Cf)
..2
;g ~ P. ? ramigera ~
§ ~ P. recta
N
~
Cf)
D. diadromum
D. rectangulare
D. magnum .....,
Cf)
z
I
~ P. variabilis
...
S1
/1
/ \
\ I
1/ Cf)
D. rectangulare
~z
I 0
P. variabilis
I
D. rectangulare (I!e~goen) I ..3'"
~
..::
D. rectangulare (Trondheim)
P. cf. opuntiformis
3
~ ~
~
8
Cf)
;j
~
Q
DlMORPHOSIPHONOIDES + LOWVILLIA Cf)
I ~!""
I
D. r~tangulare
I f=
I ) P. cf. P. recta I D
c~
D. magnum + D.
I
diadr,mum
I ~:i!
Fig. 3. Geographic and stratigraphic distribution of Udoteaceae
342 A. Roux
has been recently reported from the Ordovician of the Hudson Bay and
Canadian Arctic. Originally described from the Caradocian of Scotland,
Novantiella has been recently reported from the Silurian of Hudson Bay and
Canadian Arctic.
On the other hand, other recently described Ordovician genera such as
A nticostiporella and A rcticella are till now only known from their type-locali-
ties; Anticosti Island and Canadian Arctic respectively (see Korde 1957; Elliott
1972; Gauthier 1981; Poncet 1986).
Among the rhodophytes, it is of interest to mention the simultaneous
occurrence of Solenopora spongioides, originally described from Estonia, in
Scotland, Quebec and New York State. However, making use ofrhodophytes
for palaeogeographic reconstructions remains difficult; in fact, the Lower
Palaeozoic solenoporaceans need a complete revision.
4 Sedimentary and Palaeoclimatic Data
Middle Ordovician calcareous algae of Quebec, Ontario and New York State
are abundant but undiversified in shallow-water limestones; the Middle Or-
dovician of the Saint Lawrence platform is largely dominated by porostromates
and codiaceans-udoteaceans; the role of dasycladaceans is accessory.
The microfloras of the Upper Ordovician-Lower Silurian carbonates of
Anticosti Island show the same characteristics and suggest neither equatorial
nor tropical, but warm-temperate or subtropical waters (Gauthier 1981;
Gauthier-Coulloudon and Mamet 1981; Mamet et al. 1984).
The scarcity of dasycladaceans seems to be a characteristic of the Or-
dovician carbonates of the eastern margin of the North American continent,
except in the bioherms of the Chicamanga Group in Alabama (Stock and
Benson 1982).
The North American Ordovician is characterized by an important
development of bioherms and carbonate mud-mounds. During the Lower
Ordovician, the organic buildups occur in the calcareous facies of the external
part of the Lower Palaeozoic North American craton: the Great Basin, Texas,
Oklahoma and Newfoundland. During the Middle Ordovician (Chazy Group),
organic buildups are numerous and diverse in New York, Vermont, Quebec
(Day Point, Crown Point and Valcour Formations), Tennessee (Carters and
Holston Formations), Virginia (Effna Formation) and Newfoundland (Black
River Group, Lourdes Formation). According to Wilson (1975), Middle Or-
dovician buildups developed on a shallow-water shelf westward of the Ap-
palachian Belt in more-or-less agitated waters (see a synthesis in Roux 1985).
The Ordovician microflora of Hudson Bay and Canadian Arctic, which
contains five genera of dasycladaceans and three genera of codiaceans-
udoteaceans suggests a tropical to subtropical palaeoclimate (Poncet 1986).
In central Sweden, the Upper Ordovician (Ashgill, Boda Limestone) shows
an important reef development in which massive beds consist almost entirely of
mats of Palaeoporella variabilis. Corals and bryozoans occur mainly near the
margins of the reef and stromatolites are dominant in the massive limestone of
the center. The knoll-forming stromatolites grew on a substrate of corals and
were overgrown by Wetheredella. The dasycladacean Rhabdoporella (with R.
stolleyi and R. pachyderma) formed mats on sandy marls in the off-reef facies
(lux 1966a,b).
In Anticosti Island, Rhabdoporella is abundant in the Upper Ordovician
(Gamachian, Ellis Bay Formation). Coccoid algae played an important role in
the formation of bioherms in the upper part of the Ellis Bay Formation. The
substrate of the mud-mounds is constituted by well-sorted packs tones with
abundant Rhabdoporella; the core of the mud-mounds is made up of a Wethe-
redella-coral-bryozoan association bound by coccoid algae. The mud-mounds
of the Ellis Bay Formation occur at the maximum of the Upper Ordovician
regression (Gauthier 1981).
In Estonia, the Ashgillian is characterized by continuous decrease in water
depth. The Early Ashgillian (base of the Pirgu Formation) shows a massive
appearance of dasycladaceans ('Dasyporella') and the presence of bioherms. The
Upper Ashgillian (Porkuni Formation) shows widespread distribution of
bioherms on a stable shallow-water shelf (Roomusoks 1972).
Judging from this, the palaeogeographic conditions seem to have been
relatively the same in Anticosti Island, central Sweden and Estonia during the
Upper Ordovician. The effects of the Late Ordovician glaciation are discussed
below.
5 Palaeogeographic Significance
Empirical determination of an equatorial belt

Figure 4 shows the Upper Ordovician distribution of the studied algal groups,
plotted on a present-day map of the world.
In North America, the southernmost localities occur in Newfoundland and
Anticosti Island, at about 50 N. The northernmost localities occur in Canadian
0
Arctic Archipelago (Ellesmere Island) at about 80 N. If we consider as a first

0
approximation that the dasycladacean-udoteacean association flourished in

warm equatorial to tropical or subtropical waters, an assumed position of the
equator should have been on the median part of a belt limited by the extreme
latitudes of 50 oN and 80 N, that is, about 65 ON. The palaeoequatorran through
0
the southern part of Baffin Island.

In northwestern Europe, the southernmost localities occur in Estonia,
Scotland and southern Sweden, that is, between 55 and 60 N, and the 0
palaeoequator should have passed at about 70-75 oN, but more probably 5 to
10° northward off Estonia, Scotland and southern Sweden.
In Kazakhstan and Tadjikhistan, the localities occur between 35 and 45 °N,
and the palaeoequator should have passed at latitudes 50 to 60 N.
0
According to this assumed position of the Upper Ordovician equator,

Anticosti Island should have been located in a subequatorial position at about
15°S; Scotland, southern Sweden and Estonia should have been located at a
344 A. Roux
• VERHlPOREllA
r1 * PAlAEXlPOREUA
U e DlHORPHOSIPHON
A.R.87
Fig. 4. Upper Ordovician localities plotted on a present-day map of the world. The assumed
position of the equatorial belt is indicated by broken lines
latitude of 10° S, also in a subequatorial position. Kazakhstan and Tadjikhistan

should have been located at latitudes IS to 200S in a tropical position.
It is important to note that these conclusions are based only on a comparison
of the occurrences of Upper Ordovician localities plotted on a present-day map
of the world, but serve as a basis for further investigations concerning par-
ticularly the movement of the continents. It is of interest to compare these
conclusions with former reconstructions.
6 Comparisons with Former Reconstructions
On the Lower Ordovician map of Smith et al. (1973), the palaeoequator runs
through North America and particularly the Canadian Arctic Archipelago
which is located between 100S and lOON. Estonia and southern Sweden are
located at a latitude of about 20° S in tropical position; Scotland is located on the
western margin of the Proto-Atlantic ocean at a latitude of about 200S.
In the previously assumed position of the palaeoequator, the North
American continent is located between 200N and 4SoS, that is, within a belt of
6So . On the reconstruction of Smith et al. (1973), the North American block is
located between 30 0N and 30° S, also within a belt of 60° ; the global latitudinal
position of North America seems to have been relatively constant.
In northwestern Europe, the palaeoequator of Smith et al. (1973) runs
slightly off northern Scandinavia (10°) and Kazakhstan and Tadjikhistan are
located at about 30 0S in subtropical positions. Kazakhstan, Tadjikhistan and

northern Scandinavia are located in a belt of about 20°.
The reconstruction of Scotese et al. (1979, Fig. 13, Mercator projection)
shows a map of the world during the Middle Ordovician, from the Llandeilian
to the Early Caradocian. Present-day latitude-longitude is shown on each
palaeocontinent by a grid of five degrees apart, which can be used to plot data.
The North American block is located within a belt between 30 0N and 30 0S.
The equator runs across Canada from near the USA-Canada border
(Vancouver Island, present-day latitude 50 0N) to the Hudson Bay (present-day
latitude 60° N) and the southern part of Greenland (present-day latitude 60° N).
The extreme localities of the distribution of the microflora plotted on a pre-
sent-day map of the world (Fig. 4, Anticosti and Ellesmere Islands) are located
on the reconstruction of Scotese et al. (1979) respectively at about 15 °Sand
200N. Scotland is located between 0 and IO S. Kazakhstan (and probably
O
Tadjikhistan, the position of which is difficult to plot precisely on the recon-

structions) is completely separated from the Baltic shield (Baltica) and located
between 0 and 10° S. The Baltic shield is located far southward between 60 and
70 S.
0
7 An Upper Ordovician Map of the World
Figure 5 shows a likely position of the North American continent, the Baltic
shield and Kazakhstania-Siberia during the Upper Ordovician, proposed on the
basis of the distribution of algal microfloras.
The Proto-Atlantic ocean is nearly completely closed. To be in accordance
with the preceding remarks and particularly the sedimentary and
palaeoecological data, it seems necessary to rotate the North American block
clockwise, in order to draw Anticosti Island into a warm temperate to sub-
tropical position at about 25 to 30 0S. Scotland and Norway are then located at
nearly the same latitudes, about 20 and 25°S.
The position of Kazakhstan and Tadjikhistan remain the same as on the
map ofScotese et al. (1979), that is, about 10° S, but they are united to the Baltic
Shield because of the similarity of the microfloras.
The most important modification concerns the Baltic Shield which must be
moved northward and located between 10 and 40° S. Estonia and southern
Sweden are then located at about 25°S within the same latitudinal belt of
Scotland and Anticosti Island.
It is interesting to note that these positions are relatively similar to those of
the map of Johnson (1977, Fig. IA). Ellesmere Island is also located in an
equatorial position and the centre of Hudson Bay is at about 15°S. The
Canadian Arctic and Hudson Bay are then loca ted in a warmer position than are
Anticosti Island and Scotland and Norway.
Cocks and Fortey (1982, Fig. 5) have proposed a configuration of European
and neighbouring plates for the Late Ordovician to the Lower Silurian:
346 A. Roux
50
~
40
30
1 ......--.:t----1
Fig. 5. Upper Ordovician ma p of the world (Mercator projection) showing a likely position of
Laurentia . Baltica and Kazakhstania-Siberia, adapted from Scotese et al. (1979). The Middle
Ordovician position of Baltica is shown by broken lines
Tornquist's Sea separating the Baltic Continent from the 'Gondwana' Con-
tinent has disappeared. Northern Europe, including the southern part of the
British Isles and eastern Newfoundland, is separated from the 'Gondwana'
Continent by the Rheic Ocean; 'the Iapetus Ocean was evidently no longer the
barrier for faunal interchange it had been earlier'; the Bahamitic type of the
Baltic limestones 'and the similarity offaunas between Britain and Scandinavia
would suggest that both regions were within the tropics'.
On the map of the authors, Anticosti Island is situated at about l8°S.
Estonia within 15-20° S and Scotland within 6-11 oS. Such a configuration may
be explained by the effects ofthe important Late Ordovician glaciation, causing
a general regression and 'marked by a temporary cessation of limestone
deposition in all but the most equatorial regions'.
The concept or the effects of glaciations must, however, be used with
wariness: for example, limestone deposition in Anticosti Island is continuous
from the Upper Ordovician to the Lower Silurian; the Late Ordovician is even
marked by the development of bioherms. In Estonia, the Upper Ashgill shows
a widespread distribution of bioherms.
Was the Late Ordovician general cooling sufficiently intensive to create
subtropical or warm-temperate climatic conditions in the equatorial regions?
Was the cooling insufficient to modify significantly the distribution of the
climatic belts over the Earth? The debate remains open.
In the first case, the reconstruction of Cocks and F ortey will be correct and
then the position of Anticosti, Scotland and Baltica fits relatively well with my
Fig. 4. In the second case, the proposed position of the land masses of my Fig.
Ordovician Algae and' Global Tectonics 347
5 is probably correct and fits relatively well with the Ashgill base map ofScotese
(1984, reproduced in Beadle and Johnson 1986).
If one considers that the global direction of oceanic surface currents has not
changed during geological times, the microfloral exchanges between North
America and the 'Prototethyan' realm may be explained by such surface
currents moving counterclockwise.
8 Conclusions
The global palaeogeographic distribution of algal microfloras are closely related

to transgressive-regressive cycles and the position of continental blocks; there-
fore, fossil algae may be useful tools for palaeogeographic reconstructions.
Such studies are, however, still limited for many reasons: an empirical
determination of an equatorial belt is restricted by the number of known
localities and consequently stratigraphic and geographic distributions require
more information; sedimentary and ecological data are often lacking in the
literature; finally, the stratigraphic data are not always accurate from one region
to another or even from one author to another.
As with any new research, this contribution is certainly incomplete and must
be regarded as an introductory study. Such investigations need more careful and
extensive taxonomic, sedimentological and ecological studies within accurate
stratigraphic frameworks. Nevertheless, the first results are promising.
References
Beadle SC, Johnson ME (1986) Palaeoecology of Silurian Cyclocrinitid algae. Palaeontology 29, pt
3:585-601
Bildgen P, Roux A (l982a) Correlations entre la repartition des Algues et des bauxites de karst au
Paleozoique superieur. In: 107th Congr Nat Soc Sav, Brest, vol 3, pp 163-171
Bildgen P, Roux A (1982b) Algues, bauxites de karst et tectonique globale au Devonien-Carbon-
ifere. In: 9th Reun Annu Sci Terre, Paris, p 55
Boucek BV (1972) The paleogeography of Lower Ordovician graptolite faunas: a possible evidence
of continental drift. In: 24th Congr Geologie Int, Montreal, Sec 7, pp 256-272
Cocks LRM, Fortey RA (1982) Faunal evidence for oceanic separations in the Palaeozoic of Britain.
J Geol Soc London 139:465-478
Gauthier L (1981) Paleoeologie des Algues ordoviciennes et siluriennes de rIle d'Anticosti, Quebec.
M Sc Thesis, Univ Montreal
Gauthier-Coulloudon L, Mamet B (1981) Algues et sedimentation carbonatee dans I'Ordovicien et
Ie Silurien de I'lle d'Anticosti. In: Lesperance PJ (ed) Subcommiss Silurian stratigraphy, Field
Meet Anticosti-Gaspe, Quebec, vol 2: Stratigraphy and paleontology, pp 87-90
Elliott GF (1972) Lower Palaeozoic green algae from southern Scotland, and their evolutionary
significance. Bull Brit Mus Nat Hist 22, 4:357-376
Jansa LF, Mamet B, Roux A (1978) Visean limestones from the Newfoundland shelf. Can J Earth
Sci 15,9: 1422-1436
Johnson GAL (1977) Paleozoic accretion of western Europe. Ann Soc Geol Nord
96(1976)4:347-352
Jux U (l966a) Palaeoporella im Boda-Kalk vo Dalarne. Palaeontogr B 118,4-6: 153-165
348 A. Roux: Ordovician Algae and Global Tectonics
Jux U (1966b) Rhabdoporella im Boda-Kalk sowie in Sandsteinen Dalarnes (Ashgill, Schweden).

Paleontogr B 188,4-6:166-183
Keppie JD (1977) Plate tectonic interpretation of Palaeowic world maps. N Scot Dep Mines Pap
77-3
Korde KB (ed) (1957) Nouvelles representantes des Algues siphonees. In: Donnees pour les
'principes de paleontologie'. Akad Nauk SSSR 1:67-75 (in Russian)
Mamet B (1972) Un essai de reconstitution paleoclimatique base sur les microflores algaires du
Viseen. In: 24th Congr Geologie Int, Montreal, Sec 7, pp 282-291
Mamet B, Roux A (1977) Algues rouges devoniennes et carboniferes de la Tethys occidentale, pt 4.
Rev Micropaleontol19, 4:216-266
Mamet B, Roux A (1983) Algues devono-carboniferes de l'Australie. Rev Micropaleontol 26,
2:63-131
Mamet B, Roux A, Shalaby H (1984) Role des Algues calcaires dans la sedimentation ordovicienne
de la Plate-forme du Saint-Laurent. Geobios Mem Spec 8:261-269
Mamet B, Roux A, Nassichuk W (1987) Algues carboniferes et permiennes de l' Arctique canadien.
Geol Surv Can Bull 342
Poncet J (1986) Les Algues calcaires du Paleozoique inferieur de la Baie d'Hudson et de I'Archipel
arctique canadien. Bull Centres Rech Exp10r Prod E1f-Aquitaine 10,2:259-282
Roomusoks A (1972) Stratigraphic breaks in the Ordovician of northern Estonia and the corre-
sponding changes in the fauna. In 24th Congr Geologie Int, Montreal, Sect 7, pp 605-611
RouxA (1979) Algues du Paleowlque superieurdu Bassin de Sverdrup, Archipel arctique canadien.
D Sc Thesis, Univ Libre Bruxelles, 354 pp
Roux A (1985) Introduction Ii I'etude des Algues fossiles paleowiques (de la bacterie Ii la tectonique
des plaques). Bull Centres Rech Explor Prod Elf-Aquitaine 9, 2:465-699
Roux A (1986) Microflores algaires paleowiques (tendances evo1utives, repartitions) et tectonique
globale. Bull Centres Rech Explor Prod Elf-Aquitaine 10,2:555-563
Scotese CR, Bambach RK, Barton C, van der Voo R, Ziegler AM (1979) Paleowic base maps. J Geo1
87,3:217-277
Smith AG, Briden JC, Drewry GE (1973) Phanerozoic world maps. In: Hughes NF (ed) Organisms
and continents through time. Spec Pap Paleontol, vol 12. Palaeontol Assoc, London, pp 1-42
Stock CW, Benson DJ (1982) Occurrence and distribution of fossils within and adjacent to
Middle Ordovician bioherms in the southern Appalachians of Alabama. In: 3rd North Amer-
ican Paleontology Convention, Montreal, Proceedings, vol 2, pp 517-524
Wilson JL (1975) Carbonate facies in geologic history. Springer, Berlin Heidelberg New York
Chapter 18
Ordovician to Devonian Marine Calcareous Algae
A. Roux!
Abstract
After the Cambrian. when calcified algae are rare or poorly known. the Ordovician-Devonian
microfloras are largely dominated by chlorophytes, and particularly the filamentous (or nodular)
codiaceans and the erect udoteaceans.
Long-ranging and non-diagnostic forms are numerous and most taxa have little stratigraphic
value. It is possible, however, to determine characteristic associations or assemblages, for each
period. In addition, erect udoteaceans and the dasycladaleans show an original pattern of
development and are characteristic of systems.
1 Introduction
The first calcareous algae appear by the Lower Cambrian times with Girvanella
and probable microbial forms such as Renalcis and Epiphyton which dominate
the microftora. According to Wray (1977), the first solenoporaceans appear by
the Middle Cambrian and the first udoteaceans (Palaeoporella) by the Upper
Cambrian. The most important Cambrian event is, however, the appearance of
the first dasycladaceans (Seletonellaceae), represented by several mor-
phologically advanced genera such asA mgaella and Siberiella (Korde 1957) and
Cambroporella, Mejerella and Seletonella (Korde 1950).
2 Ordovician
The Ordovician is characterized by the appearance or the diversification of

numerous groups, particularly the filamentous codiaceans and the erect udo-
teaceans. These two groups largely dominate the microftora, but the dasyclads
also diversified.
Filamentous Codiaceans
Originally created from the Lower Carboniferous of England, Bevocastria
conglobata Garwood first occurs in the Middle Ordovician of the Saint-Law-
rence Lowlands (New York State, Quebec, Ontario) and the Upper Ordovician
of Anticosti Island (Guilbault and Mamet 1976).
'51 Avenue de Verdun, 77610 Fontenay-Tresigny, France

350 A. Roux
Ortonella Garwood emend. Mamet and Roux is represented by 0. aequalis

(H0eg) in the Middle Ordovician of the Saint-Lawrence Lowlands, New-
foundland, and Norway. O. kershopensis Garwood, which is cosmopolitan
during the Dinantian, also already occurs in the Middle Ordovician of New
York State.
Garwoodia Wood, which is cosmopolitan from the Ordovician to the
Permian, first occurs in the Middle Ordovician of New York State with G.
gregaria (Nicholson).
H edstroemia Rothpletz is known by two species: H. halimedoidea Rothpletz
in the Middle Ordovician of the Saint-Lawrence Lowlands; H. biftlosa in the
Middle Ordovician of Qut!bec and New York State.
Questionable codiaceans ? or cyanophytes ? are represented by two species
of Sphaerocodium Rothpletz emend. Wray (= Rothpletzella Wood auct., =
Coactilum Maslov): S. gotlandicum Rothpletz emend. Wood, and S. monilife-
rum (Seely). Both appear in the Middle Ordovician.
Erect Udoteaceans
The erect udoteaceans are represented by Dimorphosiphon H0eg (tribe
Dimorphosiphoneae Shuysky), Dimorphosiphonoides and Lowvillia, both of
Guilbault and Mamet (1976) (tribe Lowvillieae Shuysky), and Palaeoporella
Stolley. According to Shuysky (1987b), Palaeoporella belongs to the Dasy-
cladaceae, tribe Palaeoporelleae Shuysky. The stratigraphic and geographic
distributions of these genera are given elsewhere (see Roux, this Vol.).
Dasycladaleans
Although relatively diversified, the Ordovician Dasycladales are represented by
simple vermiporellid aspondylous forms, without regular arrangement of the
branches, thus belonging to the Seletonellaceae. The most important or com-
mon genera are Dasyporella Stolley, Kazakhstanelia Korde, Rhabdoporella
Stolley, Vermiporella Stolley, and Novantiella and Intermurella, both of Elliott
(1972), etc.
H is important to mention the discovery by Poncet (1986) of the genus
Arcticella in the Ordovician of the Canadian Arctic Archipelago (Devon
Island), which displays euspondylous branches, crowded in successive alter-
nating verticils. A rcticella is the first known true dasycladacean and the ap-
pearance of the euspondyly must be considered as a very early character in the
evolution of the dasyclads (Fig. 2).
Solenoporaceans
Although numerous and diversified, the Ordovician solenoporaceans need
careful revision; only important or common taxa are listed below: Solenopora
spongioides Dybowski is largely distributed in Estonia, Scotland, Saint Law-
rence Lowlands. New York State. S. ouareauensis Fritz, emend. Sinclair is also
abundant in the Middle Ordovician of Quebec and New York State and the
Ordovician to Devonilln Marine Calcareous Algae 351
Upper Ordovician (Gamachian) of Anticosti Island. S. trentonensis Brown is

present in Estonia (described as S. compacta by Nicholson and Etheridge 1885),
the Saint Lawrence Lowlands, New York State and Anticosti Island (Upper
Ordovician). S. canadensis (Foord), emend. Sinclair is known in the Middle
Ordovician of the Saint Lawrence Lowlands. Parachaetetes embrunensis
(Wilson) has been reported in Estonia, Ontario, Quebec and New York State.
Problematic Algae
Although known from the Silurian, Wetheredella silurica Wood occurs in the
Upper Ordovician (Ashgillian) of Sweden (as Rhabdoporella glomerata Jux) and
Anticosti Island (as Wetheredella tumulus Copper). The genus Wetheredella is
known as early as Lower Cambrian in Labrador (Kobluk and James 1979).
Halysis H0eg and Flabellia Shuysky (Codiaceae ?) play an important
sedimentological role. Halysis occurs in the Ordovician of Norway, the Saint
Lawrence Lowlands, and Anticosti Island. Flabellia occurs in the Ordovician of
the Saint Lawrence Lowlands and Anticosti Island (Guilbault et al. 1976). These
two genera are generally reported to the Hormogonophyceae by Russian
authors.
The problematic microorganism Nuia Maslov (bacterial or cyanobacterial
growth form ?) is globally distributed and ranges up to the Silurian (Nuia
distincta Ischenko). Gnoli and Serpagli (1980, Fig. 4) have illustrated the
Ordovician distribution of Nuia; see also Vachard and Tellez-Giron (1986).
Numerous species of Nuia reported in the Devonian (N. devonica and N.?
minuta Shuysky) do not belong to the genus.
Nuia is an important rock-builder, notably in the Upper Cambrian-Lower
Ordovician of Texas and the Middle Ordovician (Chazy Group) of the Lake
Champlain area and southern Quebec. Although the carbonate mud-mounds
are mainly built by coelenterates, sponges, bryozoans or tabulates, Nuia has
played an active role in the building of the mounds: this form was able to baffle
and cause local accumulations of carbonate mud. Nuia may, exceptionally, be
very abundant and cause the formation of bafflestones. The associated algae
mentioned by the authors are only Girvanella or Renalcis, with solenoporaceans
and Sphaerocodium. In most cases, Nuia is associated with Girvanella in the same
biotope. In other environments, Girvanella and Nuia are mutually exclusive.
Renalcis has been encountered in the mud-mounds of the Arbuckle Group
(Oklahoma) without Nuia; at least in this case, Nuia and Renalcis seem to be
mutually exclusive.
Other genera of Ordovician incertae algae are Guttoporella, Crinitella,
Palmatoporella, all ofGnilovskaja (1972), probable Tharama, etc. An important
event is the first occurrence of proba ble algal cysts (Radiospheres). Remnants of
Cambrian flora are the proauloporales Proaulopora Vologdin and Setula
Luchinina.
Cyclocrineae may be green algae and are very abundant in the Ordovician
with Cyclocrinites (= Cyclocrinus) Eichwald, Mastopora Eichwald, Coelo-
sphaeridium Roemer, Striola Gnilovskaja, and Apidium Stolley. Diversoporella,
352 A. Roux
Sinuatoporella, and Dolioporella, all of Gnilovskaja (1972) are reported by the

author as siphonales, together with Dimorphosiphon and Palaeoporella.
Reported as 'red algae' by Gnilovskaja (1972) but very doubtfully so, are the
Moniliporellaceae M oniliporella, Contexta, A nsoporella, Furcatoporella, Plexa,
Texturata, and Villosoporella.
Characteristic associations
The Middle Ordovician of North America is characterized by the
Dimorphosiphon-Dimorphosiphonoides-Lowvillia-Nuia assemblage with the
species D. rectangulare, D. lesperancei, L. multipora, L. raripora, and L. grandis.
The general lack of Nuia in the Black River and Trenton Groups is a local index
of the uppermost part of the Chazy Group. The Solenopora spongioides-S.
ouareauensis-S. canadensis-So trentonensis assemblage seems to be character-
istic of the Ordovician.
3 Silurian
In addition to the very long-ranging and ubiquitous Girvanella, the

Silurian microfiora is still dominated by filamentous codiaceans and erect
udoteaceans.
Filamentous codiaceans
Bevocastria conglobata occurs in the Silurian of Estonia and in the Wenlockian
to Pridolian of Quebec.
Ortonella aequalis is present in the Wenlockian-Ludlovian of Quebec (Baie
des Chaleurs), with Garwoodia gregaria and is also known from the Pridolian of
Podolia (Ischenko 1985). Although created from the Visean, Garwoodia zonata
Derville is known in the Wenlockian of Quebec (Baie des Chaleurs) as G. aff.
zonata. Ortonella rigida Lewis occurs in the Llandoverian ofW ales and remains
present in the Wenlockian-Ludlovian of Quebec. Ortonella humilis and O.
stricta have been erected from the Wenlockian and Pridolian of Podolia by
Ischenko (1985).
Likewise, Hedstroemia halimedoidea and H. biftlosa remain present in the
Silurian of Gotland. the Pridolian of Podolia (Ischenko 1985) and the Upper
Silurian of the Siberian platform (Luchinina 1987). H. halimedoidea and H.
bernierensis Mamet and Roux are present in the Wenlockian-Ludlovian of
Quebec. Hedstroemia pulchra Ischenko is reported from the Pridolian of
Podolia.
Among the questionable codiaceans? or cyanophytes? Sphaerocodium
munthei Rothpletz. emend. Wood appears in the Lower Silurian and remains
present in the Wenlockian to Pridolian (Gotland. Quebec. Anticosti Island.
Japan). S. gotlandicum remains present in the Silurian of Gotland. Sweden.
Podolia, Czechoslovakia. Great Britain, Quebec. California.
Ordovician to Devonian Marine Calcareous Algae 353
Erect Udoteaceans
The genus Litanaia Maslov appears in the Upper Silurian of Quebec (Lake
Matapedia) as L. matapediensis Mamet and Roux. In Podolia, the tribe Li-
tanaiae Shuysky is also represented by Maslovina Obrhel (Maslovina meyenii
Obrhel).
Dimorphosiphon magnum Gnilovskaja remains present in the Lower
Silurian of Kazakhstan and the Upper Silurian (Wenlockian to Pridolian) of
Quebec (Baie des Chaleurs). A new species, D. pusillus, was erected by Ischenko
(1985).
Likewise, Palaeoporella remains present in the Upper Silurian
(Ludlovian-Pridolian) of Quebec (Baie des Chaleurs) as P. sp. cf. P. recta
Gnilovskaja. The genus disappears in the Upper Silurian.
Dasycladaleans
The Silurian dasycIadaleans are represented by abundant Rhabdoporella and
Vermiporella (see Roux, this volume). Litopora Johnson occurs in the Lud-
lovian-Pridolian of Quebec (Baie des Chaleurs). Sokolella Ischenko is 'hybrid'
between Vermiporella and Wetheredella.
Solenoporaceans
Solenopora sensu stricto (S. spongioides Dybowski) is illustrated by Ischenko
(1985) from the Late Silurian of Podolia.
Parachaetetes embrunensis remains present in the Lower Silurian of On-
tario, the Upper Silurian of Gotland and the Wenlockian-Pridolian of Quebec
(Baie des Chaleurs). Although mainly a Lower Carboniferous taxon, Para-
chaetetes palaeozoicus (Maslov) is known in the Upper Silurian of Quebec
(Wenlockian to Pridolian).
Pseudochaetetes filiformis (Nicholson) and Petrophyton kiaeri H0eg are
present in the Silurian of Japan, the Wenlockian-Ludlovian of Quebec (Lake
Matapedia), and the Pridolian of Podolia (Ischenko 1985).
Charophytes
First representatives of the Sycidiceae and Trochiliscaeae are encountered in the
Ludlovian-Pridolian (Sycidium Sandberger, Praesycidium Ischenko, Primo-
chara Ischenko and Saidakovsky).
Problematic algae
Wetheredella silurica persists in the Silurian of Gotland and the Wenlockian-
Pridolian of Quebec (Baie des Chaleurs). The other species of the genus, W
pachytheca Mamet and Roux is so far only known from the Wenlockian-
Ludlovian of Quebec (Lake Matapedia).
354 A. Roux
Halysis and Flabellia are reported in the Silurian of Ufa and Podolia
(USSR) and the Lower Silurian of Anticosti Island.
The first record of Nostocites Maslov is from Ludlovian-Wenlockian of
Podolia (as 'Globochaete').
The S. gotlandicum-S. munthei-Dimorphosiphon magnum-Rhabdoporella as-
semblage seems to be characteristic of the Silurian.
4 Devonian
The Devonian microflora is relatively quite prolific. Dasycladaleans are still

distinctly subordinate to cyanophytes, rhodophytes, filamentous codiaceans,
and erect udoteaceans (Fig. 1).
Filamentous codiaceans
Bevocastria conglobata remains abundant particularly in the Tethyan Realm
(Russian platform, Afghanistan, France, Belgium, etc.).
Ortonella balbinia (Poncet) occurs in the Lower Devonian (Siegenian) of
France and O. tenuissima Garwood, described from the Carboniferous (C, ) of
Scotland, is known as O. sp. afr. tenuissima in the Famennian of New South
Wales, Australia.
Hedstroemia is represented by H. australe Johnson in the Lower Devonian
of Australia.
Sphaerocodium munthei is reported in the Devonian ofKuzbass, the Urals,
and France and S. gotlandicum persists in the Devonian of Belgium, France, the
USSR, Australia and Morocco. The Upper Devonian is characterized by the
appearance of S. exile Wray and S. magnum Wray in the Upper Devonian
(Frasnian-Famennian) of Australia (Canning and Bonaparte Gulf
basins), where they are often associated with Bevocastria and Ortonella
(Wray 1967).
Erect Udoteaceans
The erect udoteaceans show an acme during the Lower Devonian, a decline in
the Middle Devonian, and a strong reduction in the Frasnian.
During the Lower Devonian, five species of Abacella Maslov (tribe
Abacelleae Shuysky) are present: A. corn uta Shuysky (Emsian, Urals); A.
delicatula Johnson (Lower to Middle Devonian, Australia. Lower Devonian,
Urals); A. dentata Shuysky (Lower Emsian. Urals); A. pert usa Maslov (Lower
Devonian, Kuzbass, Lower Emsian. Urals); A. pustulata Shuysky (Lower
Emsian, Urals).
Litanaia Maslov (tribe Litanaiae Shuysky) is represented by eight species:
L. anirica Maslov (Lower Devonian. K uzbass. Lower Emsian. Urals - L. anirica
o
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o
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ABACELLA
CLlBECA
p...._ _ _ _ _ _ _ _ _ _+ _______ - - --- - - -l LANCI CULA
F PONCEfEllINA 1-- LlTANAIA

PARALITANAIA
1-_ _ _ _....jPSEUDOPAlAEOPOREI LLA
I
RATIO GENERA 5/24 (0 .20) 3/4 (0 . 75) 4/5 (0 .80)
SPECIES
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Fig. 1. Stratigraphic distribution and species diversity of Devonian erect udoteaceans. Note : Lancicula includes the new genera ofShuysky V>
(\985)
356 A. Roux
is chosen by Shuysky (1987) as type of the genus Praelitanaia Shuysky, tribe

Dimorphosiphoneae Shuysky); L. bella Pantie (Lower to Middle? Devonian,
Yugoslavia); L. clivosa Shuysky (Lower Emsian, Urals); L. cracens Johnson
(Lower to Middle Devonian, Australia); L. grandis Pantie (Lower to Middle?
Devonian, Yugoslavia); L. ? maslovi Palla (Lower Devonian, Carnic Alps); L.
mira Maslov (Lower Devonian, Kuzbass, Urals); L. robusta Johnson (Lower
Devonian, Australia - L. robusta is partly transferred by Shuysky (1987) to a
new genus: Semilancicula Shuysky).
Lancicula Maslov (tribe Lanciculeae Shuysky) is represented by one
species, L. alta Maslov (Siegenian, Lower and Upper Emsian of the Kuzbass
and the Urals, cited but not figured from the Lower Devonian of Greece, Chio
Island). The other species previously described as Lancicula are transferred as
follows in new genera by Shuysky (1985): Lanciculella gortanii (Palla 1966)
Shuysky (Lower Emsian, Carnic Alps); Lepidolancicula lancea (Shuysky 1973)
Shuysky (Siegenian-Lower Eifelian, Urals); Planolancicula plana (Shuysky
1973) Shuysky (Siegenian-Emsian, Urals); Lanciculella sagittata (Shuysky
1973) Shuysky (Siegenian-Emsian-Eifelian, Urals); Quasilancicula sergaensis
(Shuysky 1973) Shuysky (Siegenian-Emsian, Urals); Lepidolancicula valeriae
(Pantie 1973) Shuysky (Lower Devonian, Yugoslavia); Quasilancicula wolfi
(J ohnson 1964) Sh uysky (Lower to Middle Devonian, Australia, Lower Emsian,
Carnic Alps and Urals); Lanciculina yugorica (Shuysky 1973) Shuysky (Lower
Emsian, Cap Blanc Nez area, USSR).
The genus Circella'Schirschova, 1984 in press' is related to Litanaia but
remains a nomen nudum at this moment.
Clibeca devoniana Poncet (tribe Clibeciae Shuysky) occurs in the Devonian
of France.
The subtribe Voycarellinae Shuysky includes the genera Voycarella
Shuysky and Schirschova (V. magna Shuysky and Schirschova), Cauculicula
Shuysky and Schirschova (c. venusta Shuysky and Schirschova), Semilancicula
Shuysky with S. robusta (Johnson), pars.
Also related are the following taxa:
The tribe Ampulliporeae Shuysky with Ampullipora Shuysky (A. subita
Shuysky); Litanaella Shuysky and Schirschova (L. constricta Shuysky and
Schirschova); Praelitanaia Shuysky (P. anulata Shuysky).
The tribe Botryelleae Shuysky with Botryella Shuysky (B. spinosa Shuysky
and Schirschova).
The genus Funiculus Shuysky and Schirschova (F. venosus Shuysky and
Schirschova) does certainly not belong to the tribe Boueneae Shuysky and is a
synonym of Pseudopalaeoporella Mamet and Pn!at.
During the Middle Devonian, all species of Abacella and most of those of
Lancicula and Litanaia disappeared, excepting Lepidolancicula lancea (Lower
Eifelian, Urals), Quasilancicula wolfi (Middle? Devonian, New South Wales),
Litanaia cracens (Middle? Devonian, New South Wales), and L. grandis
(Middle? Devonian, Yugoslavia). Litanaia? bijagodensis Chuvashov (Givetian,
Urals) has been withdrawn from the genus (see below). The Eifelian Litanaia
radiosa Shuysky probably does not belong to the genus.
Palaeoporella was extinct by this time (P. lummatonensis Elliott being

withdrawn from the genus, see below).
From the Lower Givetian of Belgium, Mamet and Preat (l985a) have
described two genera representing a renewal of the Devonian udoteacean-flora:
Paralitanaia (type-species P. baileuxensis), including Litanaia? bijagodensis
Chuvashov (type-species of Bijagodella Chuvashov).
Pseudopalaeoporella (type-species Palaeoporella lummatonensis) is
monospecific and occurs also in the Upper Couvinian (C0 2d ) of Belgium (the
taxon of Elliott has been described from the Givetian of England (= Funiculus
Shuysky and Schirschova).
Described from the Frasnian of Australia, Litanaia perisseia W ra y is closely
related to P. baileuxensis (Mamet and Preat).
The genus Poncetellina Mamet and Roux (described as Poncetella by
Mamet and Roux 1983) occurs from the Lower Devonian (Lower Siegenian of
France) with P. erecta (Poncet), described as Solenopora, to the Upper Devonian
(Frasnian-Famennian of Australia, New South Wales) with P. veeversi (Mamet
and Roux 1983; Fig. 1).
Dasycladaleans
Devonian dasyclads are unevenly represented; for instance, only a questionable

vermiporellid and Litopora spaciosa are reported from Australia (Upper
Devonian of Canning Basin and Lower Devonian of New South Wales
respectively).
Forms described as Rhabdoporella, such as R. melekesensis Kulik and R.
uralica Shuysky must be withdrawn from the genus.
Other forms, however, certainly represent very important stages of the
evolution of the Palaeozoic dasycladaleans:
Unella roquellensis Poncet (Upper Siegenian of France) displays an eu-
spondyl alternating disposition of the branches (R I). This taxon is another
example of euspondyly within the Palaeozoic dasyclads.
Diplopora praehexaster Poncet (Middle Siegenian, France) and especially
Diplopora constantini Poncet (Upper Siegenian, France) are metaspondyl and
vestibular. The thallus is unsegmented, the vestibule is simple and the branches
not divided (R 1). Thus, the first known occurrence of metaspondyl vestibular
Dasycladaceae dates as early as the Lower Devonian.
Givetianella described by Mamet and Preat (1982) from the Lower Givetian
of Belgium (G. tsienii) shows a very large ramose thallus and aspondylous
branches (R I) made up of a series of spheroids. From an evolutionary point of
view, this dasyclad recalls the Lower Palaeozoic vermiporellids in the shape of
the thallus, but the morphology of the branches announces the Carboniferous
and Permian Epimastopora-like forms with halter-shaped primary branches
(Fig. 2).
Other Devonian dasyclads are:
Ulocladia Shuysky and Schirschova with U. pilosa Shuysky and Schirs-
chova and U. calva Shuysky.
358 A. Roux
GIVETIANELlA - TYPE BRANQI EPIMASTOPORA - TYPE BRANQI

ASPONDYL - Rl ASPONDYL - Rl
[GIVETIAN] [CARBONIFEROUS & PERMIAN]
DIPLOPORA CONSTANTINI - TYPE BRANQlES

METASPONDYL VESTIBUlAR - Rl [SIEGENIAN]
UNELlA - TYPE BRANQ-lES

EUSPONDYL ALTERNATING - Rl [ SIEGENIAN]
ARCfICELlA - TYPE BRANCHES

EUSPONDYL ALTERNATING - Rl [ORDOVICIAN]
Fig. 2. Shape of thallus, type and arrangement of branches of important Lower Palaeozoic
dasycladaleans (partly after Poncet 1965. 1974b, 1986 and Mamet and Preat 1982)
Fig. 3a-f. Some representative genera and species of questionable codiaceans and nodular or
filamentous codiaceans. a Sphaerocodium monili!erum (Seely) 1904. Originally described as
Slromalocerium . Middle Ordovician. C hazy Group. USA. Vermont. (Seely 1904). x35. b Sphae-
rocodium gO I/andicllm Rothpletz 1908. emend Wood 1948. Lower Devonian. Australia. Note the
characteristic type of branching. (Johnson 1964). x50. c Garwoodia air. zonata Derville 1931.
Wenlockian, La Vieille Formation, Quebec, Baie des Chaleurs. (Bourque et al. 1981). x25. d,e
Orlonella rigida Lewis 1937. Wenlockian, La Vieille Formation, Quebec, Baie des Chaleurs.
(Bourque et al. 1981). x25 . f Hedslroemia halimedoidea Rothpletz 1913. Silurian, Quebec, Lake
Matapedia. (Heroux et al. 1977). x30
360 A. Roux
Scasyporella Shuysky with S. uralica (Shuysky).

Ivdelipora Shuysky and Schirschova (related to Albertaporella) with I.
gloriosa Shuysky and Schirschova.
Rotella Shuysky and Schirschova with R. coni[era Shuysky and
Schirschova.
Coticula Shuysky and Schirschova with C. binata Shuysky and Schirschova.
Thibia Shuysky with T. proninae Shuysky.
The genera Lulipora Shuysky (type L. schratovii) and Lopsiella Shuysky
(type L. mansica) are attributed to the Acetabulariaceae. This attribution is,
however, doubtful.
Solenoporaceans
Solenopora spongioides remains present in the Lower Devonian of the Urals
with S.? taravalensis Shuysky.
Pseudosolenopora Mamet and Roux occurs in the Frasnian-Famennian of
Australia (Canning Basin) with P.? geikiei (Wray).
Pseudochaetetes Haug, emend. Peterhans is well-represented by Pseudo-
chaetetes devoniensis (Delepine) in the Lower Devonian (Siegenian) of France
and the Upper Devonian of Turkey (Anatolia).
Parachaetetes johnsoni Maslov is cosmopolitan from the Upper Devonian
upwards (Frasnian ?-Famennian of USSR, Belgium, Australia, etc.). Likewise,
P. regularis Konishi is cosmopolitan from the Frasnian-Famennian upwards
(Belgium, Alberta, and Australia where the taxon is very abundant in the Upper
Famennian of the Bonaparte Gulf Basin). Parachaetetes palaeozoicus remains
present in the Upper Famennian of the Urals; P. concentrica and P. russiensis,
both of Maslov (1956) seem to be restricted to the Devonian of the Moscow area
(for the taxonomy, see the revision ofthe Devonian solenoporaceans in Mamet
and Roux 1977; Fig. 5).
In addition to common solenoporaceans, there are mainly puzzling but
highly characteristic red algae which seem to be restricted to the period:
The Demidellaceae (with Demidella Shuysky).
The Paralanciculeae (with Paralancicula Shuysky).
The Katavellaceae (with Katavella Chuvashov) .
...
Fig. 4a-f. Some representative genera and species of erect udoteaceans. a Dimorphosiphon rec-
tangulare H0eg, 1927. Ordovician, Norway. Note the medullary tubes and the subperpendicular
lateral or cortical filaments. (H0eg 1927). x5. b Palaeoporella variabilis Stolley 1893. Upper Or-
dovician (Ashgillian) Sweden. Note the medullary filaments and the dichotomously branching
lateral filaments. (lux 1966). x30. c Poncetellina veeversi (Mamet and Roux), 1983. Holotype.
Frasnian, Cockatoo Formation, Australia, Bonaparte Gulf basin. (Mamet and Roux 1983). x25.
d,e Litanaia bella Pantie, 1973. Lower Devonian, Yugoslavia, Bosnia. (Pantie 1973). x32. f Pa-
ralitanaia baileuxensis Mamet and Preat 1985. Givetian, Belgium. Note the sinuous medullary
tubes. (Mamet and Preat 1985"). x25
362 A. Raux
Palaeosiphonocladales (issinellids and palaeoberesellids)

A major event ocurs in the Givetian with the appearance of abundant issinellids
and palaeoberesellids: Issinella devonica Reitlinger, emend. Mamet and Roux,
Pseudoissinella Mamet and Rudloff, Pseudokamaena Mamet, Kamaena An-
tropov, and Parakamaena Mamet and Roux. In Upper Devonian times,
Kamaena is diversified by the appearance of K awirsi Mamet and Roux and K
delicata Antropov. At the same time, the cosmopolitan genus Palaeoberesella
Mamet and Roux appears, then in Late Devonian Exvotarisella Elliott with P.
lahuseni (von Moller) and E. index (Ehrenberg) respectively.
Issinella and Kamaena are cosmopolitan in the northern hemisphere from
the Middle Devonian onwards. Exvotarisella index, originally known from the
Visean to the Lower Namurian of the Tethys, is reported from the Devonian-
Carboniferous boundary in eastern Siberia and the Late Famennian of Aus-
tralia. Parakamaena and Pseudokamaena are cosmopolitan in the northern
hemisphere from the Middle Devonian onwards.
lansaella ridingi Mamet and Roux, described from the Devonian of Alberta
is an incertae algae, probably a chlorophyte.
The palaeoberesellids most probably originate from the issinellids. From
lansaella to the palaeoberesellids through the issenellids, one observes a pro-
gressive arrangement of the pores ('branches'): in lansaella, they are very thin
and numerous, straight and arranged without any order; in Issinella, they are
apparently less numerous, but perpendicular to the cortex; in the palaeobere-
sellids, the pores are again less numerous and located only in parts of thallus
between two consecutive 'septa'. The pores are simple in Kamaena,
Palaeoberesella, Pseudokamaena, Kamaenella and Parakamaena and dicho-
tomous with three orders of ramification in Exvotarisella (Fig. 6).
Kamaena, Kamaenella, Palaeoberesella, Parakamaena, and Exvotarisella
are included in the Palaeosiphonocladales by Shuysky (1985). The same genera
with lansaella and Pseudokamaena are attributed to the Dasycladales, Bere-
sellaceae Deloffre, tribe Palaeobereselleae Mamet and Roux, by Deloffre
(1987). Issinella and Pseudoissinella are attributed to the tribe Issinelleae
Deloffre (see Deloffre 1988).
Shuysky (1987b) includes in the Siphonocladales the genera Calcicaulis
Shuysky and Schirschova (c. vesiculosum Shuysky and Schirschova) and
...
Fig. Sa-f. Some representative rhodophytes. a Parachaetetes embrunensis (Wilson), 1945. Wen-
lockian, La Vieille Forma tion, Quebec, Baie des Chaleurs. (Bourque et al. 1981) x25. b,c Solenopora
ouareauensis Fritz 1941, emend. Sinclair 1956. Upper Ordovician, Ellis Bay Formation, Anticosti
Island. (Gauthier 1981). Note the presence ofa possible sporangia I structure (b) and the "septiform
processes" of the longitudinal partition of the tubes (c). x25. d Petrophyton kiaeri H0eg 1932.
Ordovician, Norway. (From H0eg 1932). x15. e Parachaetetes johnsoni Maslov 1962. Upper
Famennian, Belgium. Note the irregular diameter of the tubes and the horizontal partitions situated
at the same level from one tube to another. (Mamet and Roux 1977). x55. f Parachaetetes regularis
Konishi 1958. Upper Famennian, Australia, Bonaparte Gulfbasin. Note the very regular arrange-
ment of the cells. (Mamet and Roux 1983) x62
364 A . Raux
-" ...
,'"
.I'..
~% ,#-
.J
,.'...
-,
...tT•
...
,'4
\~~ \
. ~
J'
I~\.
.v": &11/".
,.....
.,f'
b ...r:.
. lP-
n
;
Parmacaulis Shuysky and Schirschova (P. hulgensis Shuysky and Schirschova).

These genera are, however, radically different from any known Siphonoclades.
Umbellids and charophytes

Charophytes, which find their roots in the Silurian, are very prolific in the
Devonian with representatives of Chovanellaceae Grambast, Pinnoputame-
naceae Wang and Lu, Sycidiaceae Karpinsky, emend. Peck, Trochiliscaceae
Karpinsky, and Karpinskyaceae Wang and Lu.
Many Russian authors include the umbellids among charophytes. They
have, however, a particular wall structure (a micritic inner layer and a finely
prismatic outer layer) unknown among trochiliscids. They appear in the Lower
Devonian, are extremely common in non-marine, brackish and littoral waters,
and become extinct in the Early Carboniferous.
Well documented Devonian genera are: Umbellina Loeblich and Tappan
(= Umbella 'Maslov' in Bykova and Polonova 1955); Elenia Pojarkov;
Eoumbella Platonov (= Costatumbella Berchenko); Planoumbella Platonov;
Spinoumbella Platonov; Quasiumbella Pojarkov; Biumbella Mamet (= Qua-
siumbelloides Berchenko); Kusjaella Chuvashov; Lagenumbella Mamet; Col-
lumbella Mamet and Roux.
Problematic algae
Wetheredella silurica remains present in the Upper Devonian (Upper Frasnian)
of Belgium and Halysis and Flabellia are cited from the Lower Devonian of the
Urals.
It is important to mention the presence of Labyrinthoconidae in the
Givetian of Belgium, Germany and Moravia, whose wall structure and mor-
phology announce the calcified wall of the Carboniferous Stacheiinae, with,
however, a different morphology of the cell threads (Mamet et al. 1982).
Proninella Reitlinger in Menner and Reitlinger, and Asphaltina Mamet in
Petryk and Mamet are present. Calcispheres become very abundant in
...
Fig.6a-f. Some representative dasyciadaleans, paleoberesellids and problematic algae (renalcids
and microcodiacids). a Vermiporella acerosa Gnilovskaja 1965. Upper Ordovician, USSR, Tar-
bagatay Range. Note the dichotomy of the thallus and the simple branches, aspondyl Rl.
(Gnilovskaja 1965). x30. b Vermiporella wesenbergensis Moskalenko 1956. Upper Ordovician,
USSR, Tarbagatay Range. Note the dichotomy of the thallus and the very large central stem.
(Gnilovskaja 1965) x30.c Diplopora constantini Poncet 1965. Holotype. Siegenian, France. Note the
continuous thallus and the clusters of branches issuing from simple vestibules. (Poncet 1965). xiS.
d Kamaena delicata Antropov 1967. Upper Tournaisian, Great Britain. Note the shape of the
medullary cavities. (Mamet and Roux 1974). x135. e Renalcis granosus Vologdin 1932. Frasnian,
Cockatoo Formation, Australia, Bonaparte Gulf Basin. Note the transition Renalcis(R); Izhella(l);
Shuguria (S); Chabakovia (C). (Mamet and Roux 1983). x62. f Palaeomicrocodium devonicum
Mametand Roux 1983. Holotype. Frasnian, Cockatoo Formation, Australia, Bonaparte Gulf Basin.
(Mamet and Roux 1983). x36
366 A.Roux
Devonian lagoons and restricted environments (Calcisphaera, Parathuram-

mina, Palaeocancellus, Radiosphaera, etc.).
Renalcis and related forms

The Lower Palaeozoic microfloras contain long-ranging and non-diagnostic
taxa such as Girvanella, Renalcis or epiphytales of uncertain affinity (microbial?
forms).
Renalcis Vologdin, emend. Mamet and Roux is itself a growth-form,
including I zhella Antropov, Chabakovia Vologdin, and Shuguria Antropov (see
the revision of the genus Renalcis in Mamet and Roux 1983).
Epiphyton Bornemann is very common in the Lower and Middle Cambrian
and is often a rock-former. Although reported from the Devonian of USSR, no
Ordovician or Silurian forms are known.
A rela ted taxon, Paraepiphyton, is described by W ray (1967) from the Upper
Devonian (Frasnian-Famennian) of Australia. Paraepiphyton and Epiphyton
may be synonymous. Paraepiphyton is often associated with Renalcis.
Another form, Tharama, is described by the same author (Wray 1967) from
the Frasnian-?Famennian of Australia, where it is associated with Renalcis and
Epiphyton. Tharama is known in the Middle Ordovician ofN orth America (N ew
York State and Ontario).
Forms closely related to Frutexites are illustrated by Mamet and Roux
(1981, in Bourque et al.) from the Silurian of Quebec.
Frutexites was originally described without a type by Maslov (1960) from
Ordovician stromatolites of USSR. The 'genus' is known in Devonian
stromatolites in Australia and was probably an important rock-former in the
Devonian reefs in Alberta. Likewise, forms described as 'Epiphyton' are im-
portant rock-formers in the Devonian reefs of Belgium.
M icrocodiaceae
It is important to mention the discovery by Mamet and Roux (1983) of
Microcodiaceae Maslov in the Upper Devonian of Australia. Palaeo-
microcodium devonicum Mamet and Roux is common in marine back-reef
environments and often associated with Renalcis. Since then, Microcodi-
aceae have been found in Givetian-Frasnian (Mamet and Preat 1985b).
The Litanaia-Lancicula-Abacella association is typical of the Lower Devonian.
During the Upper Devonian, the S. exile-P. johnsoni-P. regularis-P. geikiei
assemblage seems to be characteristic.
5 Conclusions
Despite an abundant literature which is often confusing, the Lower Palaeozoic

microfloras remain poorly known (Riding 1979). Many problems remain to be
solved: the explosion of heavy calcified forms by Lower Ordovician times
implies the existence of Cambrian unca1cified stocks; the Silurian dasyc1ads are
poorly known and the affinities of many taxa are uncertain. Deciphering the
evolution of the Lower Palaeozoic algae requires further taxonomical and
stratigraphical studies.
Acknowledgements. Dr. B.L. Mamet has kindly read the manuscript and made useful comments.
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Geofiz Novosibirsk 674:38-109 (in Russian)
Sinclair GW (1956) Solenopora canadensis (Foord) and other algae from the Ordovician of Canada.
Trans R Soc Can L, Ser III, 4:65-81
Stolley E (1893) Uber silurische Siphoneen. N Jahrb Mineral Geol Palaontol2: 135-146
Vachard D, Tellez-Giron C (1986) EI alga Nuia en el Ordovicico de Mexico: hipotesis diversas. Rev
Inst Mex Petrol 18(2):12-25
Wood A (1948) Sphaerocodium, a misinterpreted fossil from the Wenlock Limestone. Proc Geol
Assoc 59:9-22
Wood A (1957) The type-species of the genus Girvanella (calcareous algae). Palaeontology I,
1:22-28
Wray JL (1967) Upper Devonian calcareous algae from the Canning Basin, western Australia. Prof
Contrib Col Sch Mines 3
Wray JL (1977) Calcareous algae; developments in palaeontology and stratigraphy. Elsevier,
Amsterdam, 185 pp
Chapter 19
Carboniferous Calcareous Algae
Abstract
This compendium records 139 critically assessed valid genera and 22 form-genera of Carboniferous
blue-green, green and red algae. It also includes some umbellinids, algal cysts and incertae sedis. For
each genus, it lists the valid species and assesses suprageneric attributions. It records stratigraphic
range, geographic distribution and abundance. The role in carbonate sedimentation is emphasized.
The vast majority of algae are very long-ranging and cannot be used for precise age
determination. Obvious exceptions are Calcifolium and Koninckopora.
1 Introduction
Carboniferous calcareous algae are diverse, abundant and controversial. The

purpose of this compendium is not to add fuel to disputable classifications but
to critically record what is securely known, and list valid generic and specific
taxa, with their known stratigraphic range, their role in sedimentation and their
observed geographic distribution. Suprageneric attributions remain tentative
and largely equivocal.
Numerous attempts at classifications have recently been published: Bas-
soullet et al. (1979) on dasycladales; Bassoullet et al. (1983) on udoteaceans;
Chuvashov, Luchinina, Schaikin and Shuysky edited by Dubatolov (1987); and
Deloffre (1988). But there is still little concensus on palaeobotanical attribu-
tions. Dasyclads are morphologically quite complicated and are therefore only
partially understood. On the other hand, simple cyanobacteria remain a maze
of puzzling and contradictory taxonomical classes and orders.
These taxonomic limitations admitted, algae are nevertheless of paramount
importance for understanding carbonate sedimentation and furnish some
useful stratigraphic observations. The purpose of this compilation is therefore
to condense taxonomic and stratigraphic information gleaned from numerous
articles together with unpublished personal observations.
2 Calcareous Algal Microftora
Nearly two hundred genera have been described in the literature (197 taxa). In
this compilation, 35 of these genera are regarded as synonyms. An additional 22
'Departement de geologie, Universite de Montreal, c.P. 6128, succursale A., Montreal, Quebec
H3C 317, Canada
Carboniferous Calcareous Algae 371
genera are "composite taxa", usually the result of biological activity of two or
three organisms (spongiostromids). This leaves us with a grand total of 139
genera and 330 valid species.
For the sake of simplicity, genera are divided into informal groups and
plotted on an idealized Carboniferous carbonate platform (Fig. I).
1. Cyanobacteria (or cyanophytes) are omnipresent, very long ranging

through the whole Paleozoic and represented in the Carboniferous by only
seven genera ofChabakoviaceae, Epiphytaceae and Girvanellaceae.
2. Twenty-two "genera" of spongiostromids (algal biscuits) are common in
Tethysian protected lagoons ("Chondrostoma", "Polymorphocodium",
"Pycnostroma") but need further study to establish their real identity.
3. Nodular codiaceans (?) or nodular cyanophytes (?) comprise 8 genera of
Garwoodiaceae. Bevocastria, Garwoodia, H edstroemia, M itcheldeania,
Ortonella, thrive in Palaeozoic restricted and semi-restricted
environments.
4. Carboniferous udoteaceans are poorly understood. Five genera, all
strongly endemic could possibly be attributed to the group: Cummingsella,
Halenopora, Orthriosiphon, Orthriosiphonoides and Taldykites.
5. Anchicodiaceae ("phylloid" algae in the restricted sense) comprise only 4
genera, but they can be very abundant and play an important role in
building Middle Carboniferous-Early Permian mounds (Anchicodium-
Eugonophyllum-Ivanovia-Neoanchicodium). Early marine cementation of
phylloid blades could indicate a cyanobacterial consortium.
6. The Palaeosiphonocladales are important for carbonate production. They
are erect, small, simple cylindrical thalli, with rather simple functional
pores or with an outer mucilagenous coating, now preserved as cement.
They thrive in dense closely-packed thickets in protected lagoons and
mounds. Devonian-Early Carboniferous palaeoberesellids (10 genera
including Exvotarisella, Kamaena, Palaeoberesella) and issinellids (3
genera including Issinella, Pseudoissinella) were succeeded by Late Car-
boniferous-Early Permian beresellids (6 genera including Beresella,
Dvinella, Uraloporella).
7. The most diversified, endemic to a continental block, and often strati-
graphically confined chlorophytes are the Dasycladales sensu stricto.
Seletonellaceae and Diplororaceae are scarce. The presence of Ace-
tabulariaceae is doubtful. The bulk of the Dasycladaceae is observed
in the Tethyan realm. At least 36 genera have been identified (Alber-
taporella, A natolipora, A nthracoporella, A tractyliopsis, Borisovella,
Cabrieropora, Clavaporella, Coelosporella, Columbiapora, Connexia,
Dutroella, Eouraloporella?, Eovelobitella, Epimastopora, Epimastopo-
rella, Gyroporella, Herakella, Japhetella, Koninckopora, Kulikia, Macro-
porella, Masloviporella, Melporella, Nanopora, Paraepimastopora,
Pekiskopora, Pseudogyroporella, Pseudonanopora, Pseudovelebitella,
Queenslandella, Sinarella, Sphinctoporella, Unjaella, Vermiporella.
Windsoporella, Zaporella).
PLATFO RM EDGE O PEN MARIN E OPEN MARINE BARRIER SEMI· RESUICTED RESTRICTED SUPRA TIDAL SEBKAH
(PHOTIC) (WAVE ACTION) (MO UND OR SHOA L) LAGOO N LAGOON
WAVE ACTION
PHOTIC ZONE
1 - CY ANOBACTERIA
2 - SPO NGIOSTROM IDS

--
3 - GARWOODIACEAE
- - --- --
- - ----- - - - - -
4 - UDOTEACEANS
5 - ANCH ICOD IACE AN S

---
~
6 - PALAEOS IPH ONOCLADALES
7 - DASYCLADALES
• •••
B - WETHEREDEllIDS
---
9 - SOlENOPORIDS
10 • UNGDAREllIDS
.ND
\I - A RCHAEOLI THOPH YlllDS -- - -
12 - 13 ST ACHE IINS ."" UNDETE RM INED REO ALGAE
14 _ UMB EllI NIDS +- +- +- +- +- +- +-
+- +- +- +- +- +- +- +- +- +-
15 - CAL CISPHERES
16 _ MIC ROCODIA CE ANS
17 _ TU BIPHYTES
Fig. 1. Idealized reconstruction of a Carbo niferous ca rbo na te p la tform. Distribution and abund a nce of a gl a l microfl ora a nd incertae sed is. Width of
bars proportional to abundance. A rrolVs indicate reworki ng
8. Seven genera of incertae chlorophytes could possibly be related to the

wetheredellids. Asphaltina or Berestovia are observed in high energy
shoals.
The next five groups, from 9 to 13, belong to the red algae.
9. The simple forms of rhodophytes are the long ranging solenoporids (4
genera: Parachaetetes, Pseudochaetetes, Pseudosolenopora, Solenopora).
While there are no records of Carboniferous reef crests built by solen-
oporids, they are often observed as discrete nodules in open marine lagoons
or on open marine ramps.
10. Ungdarellids (5 genera) form important constituents of boundstones and
Komia and Ungdarella are cosmopolitan reef formers.
11. Archaeolithophyllids (2 genera) are crustose or form baffles.
12. Stacheiins are omnipresent in Visean-Namurian carbonates (10 genera
including A oujga lia , Epistacheoides, Mametella, Pseudostacheoides,
Stacheoides).
13. Six other red algae cannot be attributed to well-defined families. The most
widespread is Cuneiphycus and related forms.
14. Charales probably include umbellinids, which thrived in the Devonian.
They peter out at the Devonian-Carboniferous boundary but 6 genera are
still known in the Early Carboniferous (Biumbella, Protoumbella,
Quasiumbella).
15. Calcispheres (Calcisphera, "Radiosphaera" are micritic-walled cysts which
proliferated in lagoons. They floated and were easily dispersed into open
marine facies and spread as far as the platform margin.
16. About twenty incertae sedis (16-17) are tentatively included among
the algae? Microcodiaceans (16) (Microcodium, Palaeomicrocodium)
perforated and dissolved marine grainstones. Tubiphytes (17) erected
Late Carboniferous-Early Permian mounds in association with
Archaeolithoporella.
3 Algal Microfacies
Algal associations permit recognition of marine microfacies belts in a carbonate

ramp or platform.
The deepest algal floras are composed ofstacheiins and solenoporids which
coincide with the deepest part of the photic zone. Higher up in the open marine
facies, rhodophytes are mixed with palaeosiphonocladales. The first dasyclads
and wetheredellids thrived in high energy shallow water shoals. Boundstones
are only formed by early cemented phylloid anchicodiaceans, Tubiphytes or
crustose ungdarellids. Slightly deeper water mud-mounds are formed by
beresellids.
Semi-restricted lagoons have abundant dasyclads, calcispheres and
palaeosiphonocladales, cyanobacteria and nodular codiaceans? Restricted
lagoons have cyanobacteria, spongiostromids, few dasycladales, umbellinids
and calcispheres.
374 B. Mamet
Microcodiaceans developed in lagoons due to fresh water influx, or in

supratidal sabkhas with vadose cementation, and are associated with spon-
giostromids and cyanobacteria.
4 List of Genera
The following are indicated for each genus:
I. Type of the genus with age and location;

2. Diagnosis;
3. Valid species in the Carboniferous, with stratigraphic range and geographic
distribution. Flora is reported from the different basins in the following
conventional order. Western Tethys: North Africa (Libya, Algeria, Mo-
rocco), Western Europe (Spain, France, Belgium, Great Britain, Ireland,
Germany, Carnic Alps, Yugoslavia, Poland), Eastern Europe (Byelorussia,
Russian Platform, Donbass, Middle and Southern Urals).
Eastern Tethys: Turkey, Iran, Afghanistan, Central Asia (Kazakhstan,
Tadzhikhistan, Tian-Shan), Southern China, Indochina, Australia.
Siberian block: Kuzbass, Kolyma, Taimyr, Verkoyansk.
Arctic: Omolon, Arctic Alaska, Arctic Canada.
America: American Cordillera (British Columbia, Alberta, Idaho, Mon-
tana, Utah, Colorado, New Mexico), American Midcontinent (Missouri,
Arkansas, Oklahoma), American Appalachians (Tennessee, Alabama,
Maritime Provinces of Canada). We have no information on South
American microfloras;
4. Suprageneric attributions;
5. Stratigraphic range and distribution for the genus (same convention as
above). When the alga plays an important part in sedimentation, this role
is emphasized.
The list includes taxa from the Devonian-Carboniferous transition

("Strunian") and from the basal Permian (Asselian).
l. A lbertaporella (Johnson 1966) Mamet and Roux 1981
Type of genus. Albertaporella involuta Johnson 1966. Latest Tournaisian,
Alberta.
Diagnosis. Thallus a succession of spherical verticillated units. Pores phloio-
phore, simple, in tufts radiating from a vestibule.
Other Carboniferous species.
A lbertaporella (?) gallatinensis Mamet and Roux 1981. Early Visean, Montana,
Nevada, Utah.
Suprageneric attributions. Albertaporellae (Guven\ 1979) Mamet and Roux
1981 according to Deloffre (1988). Unconvincingly attributed to the Tri-
ploporelleae (Pia 1920) Bassoullet et al. 1979 by Shuysky (1987).
Stratigraphic range and distribution. Reported by Deloffre (1988) as Devon-
ian-Carboniferous. This wide range is based on Giiven~'s assessment that

A lbertaporella included the Siegenian Diplopora constantini Poncet 1965 (=
IvdeUpora Shuysky and Schirchova 1987) and the Late Permian Diplopora
anatoUana Giiven~ 1970. Correct range is restricted to Late Toumaisian-Early
Visean. Endemic to the American Cordillera (Alberta, Montana, Nevada,
Utah).
2. Amorfia Rlicz 1965

Type of genus. Amorfia jaUnki Racz 1965. Lower Moscovian, Spain.
Diagnosis. Thallus crustose, wavy, branching, flattened cylindrical, with hypo-
and perithallic differentiation. Thin polygonal hypothallic tissue and thick
rectangular perithallic cells. Cell threads joined in curved bundles. Concepta-
cles circular.
Suprageneric attribution. Archaeolithophyllaceae Chuvashov 1987.
Stratigraphic range and distribution. Scarce. Same as type of genus.
3. Anatolipora Konishi 1956

Type of genus. Anatolipora carbonica Konishi 1956. Late Visean-Early
Namurian, Japan.
Diagnosis. Thallus club-shaped, small with undivided medulla. Branches
numerous, phloiophore, with strong oblique insertion and slight curvature.
Suprageneric attribution. Salpingoporellinae Bassoullet et al. 1979 according to
Shuysky (1987) and Deloffre (1988).
Stratigraphie range and distribution. Late Toumaisian-Early Namurian, Spain,
France, Belgium, Japan, Alaska, Alberta.
4. Anchicodium Johnson 1946

Type of genus. Anchicodium funile Johnson 1946. Upper Carboniferous,
Kansas.
Diagnosis. Crustose, wavy, phylloid plates. Spongy mass of highly branching
threads, terminated in triangular-shaped "utricles".
Anchicodium gracile Johnson 1946. Late Carboniferous, Kansas, Urals.
Anchicodium magnum Endo 1952. Namurian-Permian, Algeria, Japan.
A nchicodium nodosum Johnson 1946. Late Carboniferous, Kansas.
Anchicodium plumosum Johnson 1946. Late Carboniferous, Kansas.
Suprageneric attributions. Siphonales, Anchicodiaceae according to Shuysky
(1987). Filaments were confused with endolithic algal perforations by Vachard
(1981).
Stratigraphic range and distribution. Locally abundant. Forms bafHestones and
build-ups (early cementation). Middle Carboniferous-Permian. Spain,
Yugoslavia, Russian Platform, Urals, Kazakhstan, Afghanistan, Japan, Alaska,
Kansas.
376 B. Marnet
5. Anthracoporella Pia 1920

Type of genus. Anthracoporella spectabilis Pia 1920. Upper Carboniferous,
Austria.
Diagnosis. Thallus large, branching, vermiporellid-1ike. Medulla prominent.
Branches crowded, numerous, ph10iophore, aspondy1, at right angles from the
medulla.
Anthracoporella girtyi Mamet and Roux 1987. Moscovian-Sakmarian, Can-
adian Arctic.
Anthracoporella vicina Kochansky-Devide and Herak 1960. Upper Carbon-
iferous, Yugoslavia.
Suprageneric attributions. Tribe Dasyporelleae (Pia 1920) Bassoullet et al. 1979
according to Deloffre (1988). Tribe Anthracoporellopsiae Sh uysky 1985 among
the Palaeoberesellaceae.
Stratigraphic range and distribution. Common. Middle Visean? to Permian.
Devonian records are to be disregarded. Forms the "Anthracoporella com-
munity" of Fliige1 (1977) in association with phylloid algae. Spain, Yugoslavia,
Greece, Turkey, Russian Platform, Urals, Tadzhikhistan, Afghanistan,
Kazakhstan, China, Thailand, Sumatra, Japan, Canadian Arctic, Texas.
6. Anthracoporellopsis Maslov 1956

Type of genus. Anthracoporellopsis machaevi Maslov 1956. Middle Carbon-
iferous, Donbass (see Plate 3a).
Diagnosis. Thallus cylindrical, rarely ramified. Medulla divided by irregular
partitions. Pores aspondyl, straight, perforating both cortex and partitions in all
directions.
Suprageneric attributions. Anthracoporellopsiae Shuysky 1985 in Shuysky
(1987). Beresellaceae for Deloffre (1988). Palaeobereselleae? for Mamet and
Roux (1974).
Stratigraphic range and distribution. Rather common. Tournaisian to Early
Permian. Morocco, Algeria, Spain, Belgium, France, Russian Platform, Urals,
Plate I. a-I Idealized reconstructions of thalli. a Columbiapora. Reconstruction showing vestibule

(in Marnet and Roux 1981 b). From Columbiapora johnsoni Marnet 1974. Approximate diameter of
the verticils, up to 1000 mIL. Tournaisian. British Columbia. b Queenslandella (in Mamet and Roux
1983). From Queenslandella jenkinsi Mamet and Roux 1983. Approximate diameter of thallus. up
to 700 mIL. Visean, Australia. c Eovelebitella (Mamet and Roux 1983). From Eovelebitella occitanica
Vachard 1974 emend. Approximate diameter of verticils, 900-1500 mIL. Late Visean, France. d
laphetella (in Mamet and Roux 1983). From laphetella boehmi Mamet and Roux 1983. Approx-
imate diameter of thallus. around 750 mIL. Late Visean. France. e Microcodium. Reconstruction of
the "corn cob" (in Mamet et al. 1987). Outside diameter of the cob. 400-1000 mIL. Plates dissolve and
gouge into surrounding grains. Moscovian. Yukon Territory. f Nansenella. Reconstruction of the
branching habit (in Mamet et al. 1987). Adapted from Nansenella multifurcata Mamet and Roux
1987. Diameter ofmicritized sheaths 25-80 mIL. Bashkirian. Ellesmere. g lssinella Reconstruction
of the branching habit (in Mamet and Roux 198Ia). From lssinella devonica Reitlinger 1954.
~ - I
~j
Diameter of thallus 100 to 600 mIL. Late Visean , Utah . h Kamaenella. Cross-section showing
bifurcation (in Mamet and Roux 1974). From Kamaenella denbighi Mamet and Roux 1974.
Diameter of thallus 80- 140 mIL. Late Visean, Great Britain. i-j Hypothetical views of a phylloid
Anchicodiaceae. i Ivanovia (in Mamet et al. 1987) . Based on Ivanovia lenuissima Khvorova 1946.
Blades reach up to I mrn in thickness. Thallus pluri-rnillirnetric to centirnetric. Moscovian,
Canadian Arctic. j Neoanchicodium (in Marnet et al. 1987). Based on Neoanchicodium calenoides
Endo in Endo and Kanurna 1954. Same order of magnitude as for i. Moscovian , Canadian Arctic.
k Richella (in Marnet et al. 1987). Based on Richella incruslala Mamet and Roux 1987). Diameter
of tubes around 300- 360 mIL. Moscovian, Canadian Arctic. I Epislacheoides (in Marnet and Roux
1977). Hypothallus (E V) with irregular, poorly calcified cells. Peri thallus heavily calcified (EH) , up
to 300 mIL with regular cells (e3). Based on Epislacheoides nephroformis Petryk and Marnet 1972.
Visean , Alberta
378 B. Marne!
0. ! ~II ! ~R
; ~M I III
e f
f".
f~,)
1;"'\', 7 C 9 h
~/
Plate II
Donbass, Kazakhstan, Siberia, (Kuzbass, Omolon), Canadian Arctic, Alaska

(Peratrovich Fm).
7. Aoujga/ia Termier and Termier 1950

Type ofgenus. Aoujga/ia variabilis Termier and Termier 1950. Middle? Visean,
Morocco.
Diagnosis. Thallus encrusting, fusiform. Cells irregular, grossly concentric
around support. Wall strongly calcified, hyaline calcite. No thallus
differentiation.
Aoujgalia elliotti Mamet and Roux 1977. Visean to Sakmarian. Ireland, Siberia
(Kuzbass), Canadian Arctic, Alaska.
Aoujga/ia richi Mamet and Roux 1978. Visean-Baschkirian. Siberia (Kuzbass),
Canadian Arctic, Alaska, Tennessee.
Suprageneric attributions. Originally described as a foraminifer, then trans-
ferred to the "ordre nouveau d'Ischyrosponges Aoujgaliida" by Termier et al.
(1975). Stacheinaceae for Chuvashov (1987) or Stacheiinae for Mamet and
Roux (1978).
Stratigraphic range and distribution. Common from Visean to Namurian.
Reaches the Permian in the Canadian Arctic. Morocco, Algeria, Spain, France,
England, Belgium, Germany, Donbass, Kazakhstan, Siberia, (Kuzbass,
Taimyr, Verkoyansk), Australia, Canadian Arctic, Alaska, American Cordil-
lera, Tennessee .
...
Plate II. a-q Idealized reconstructions of thalli. a-c Clavaporella. a Cross-section; b reconstruction;
c oblique section through pore and vestibule (in Mamet et al. 1987). Based on Clavaporella reinae
Racz 1966. Diameter ofverticils 1000 to 1500 mj.l. Moscovian, Canadian Arctic. d Masloviporidium.
Foliate thallus with pores in horizontal partitions (in Groves and Mamet 1985 after Maslov 1973).
Height of quadratic cells, 20-75 mj.l. Namurian, Donbass. e-h Shapes and disposition of blind pores
among beresellids. All material is Middle Carboniferous. e Beresella (in Mamet et al. 1987). Based
on Beresella erecta Maslov and Kulik 1956. f Dvinella (Dvinella) (in Mamet et al. 1987). Based on
Dvinella comata Khvorova 1949. g Dvinella (Trinodella) (in Mamet et al. 1987). Based on Dvine/la
bi[urcata Maslov and Kulik 1956. h Uraloporella (in Mamet et al. 1987). Based on Uraloporella
variabilis Korde 1950. i Donezella. Reconstruction and longitudinal section (in Mamet et al. 1987).
Based on Donezella lutugini Maslov 1929 emend. j Calci[olium. Reconstruction of thallus by
Skompski (1981). Based on Calci[olium okense Schvetsov and Birina 1935. Late Visean-Early
Namurian, Poland. k Cummingsella. Reconstruction and longitudinal section (in Mamet and Roux
1980). Based on Cummingsella lyoncrossi Mamet and Roux 1980. Diameter of thallus 400-500 mj.l.
Namurian, Scotland. I Orthriosiphonoides. Reconstruction and longitudinal section (in Mamet and
Roux 1980). Based on Orthriosiphonoides salterensis Petryk (in Petryk and Mamet 1972). Diameter
of thallus, 1100-2000 j.lm. Visean, Alberta. m Eovelebitella. Longitudinal section of Eovelebitella
oCcitanica Vachard 1974. Late Visean, Spain. n Epimastopora. Schematic reconstruction of pores (in
Roux 1979). Based on Epimastopora piai Korde 1951. Upper Carboniferous, Urals. 0 Pa-
raepimastopora. Schematic reconstruction of pores (in Roux 1979). Based on Epimastopora kan-
sasensis Johnson 1946. Moscovian, Kansas. p Epimastoporella. Schematic reconstruction of pores
(in Roux 1979). Based on Epimastopora japonica Endo 1950. Early Permian, Japan. q Gyroporella.
Reconstruction showing sha pe of pores (in Mamet et al. 1987). Based on Gyroporella nipponica Endo
and Hashimoto 1955. Upper Carboniferous-Early Permian, Canadian Arctic
380 B. Marne!
=~-
v
Plate III
8. Aphralysia Garwood 1914

Type ofgenus. Aphralysia carbonaria Garwood 1914. Late Tournaisian-Visean,
Great Britain.
Diagnosis. Thallus encrusting, nodular, maze oflayered cystoid cells. In axial
sections, cells are semi-elliptic with concave base attached to convex wall of
preceding row. Wall micritic.
Aphralysia capriorae Mamet and Roux 1975. Visean-Namurian, Algeria, Bel-
gium, France, Great Britain, Alaska, Tennessee.
Aphralysia ferreoli Mamet and Roux 1975. Silurian-Namurian, France, Aus-
tralia, Arkansas, Quebec.
Aphralysia garwoodi Hallett 1970. Late Visean, Great Britain.
Aphralysia matthewsi Mamet and Roux 1975. Tournaisian-Namurian, Great
Britain, Australia, Tennessee, Arkansas.
Suprageneric attributions. Incertae chlorophyte for Mamet and Roux (1975).
Chlorophyte - Wetheredelleae according to Berchenko (1987).
Stratigraphic range and distribution. Known as early as Silurian and quite
abundant in Eifelian of Belgium. Common in Tournaisian-Lower Namurian .
...
Plate OJ. a-v Illustrations of thalli. a Anthracoporellopsis machaevi Maslov 1956 (in Marnet and
Pinard 1985, Peratrovich Island, Alaska, Zone 18, Early Namurian, x28). b Exvotarisella index
(Ehrenberg 1854 emend von Miiller 1879) (in Marnet and Roux 1974, Halecornbe, Great Britain,
Zone 13, Middle Visean, x36). c Nanopora cr. N. fragillissima (Maslov 1939) (in Marnet and Rudloff
1972, Antigonish, Nova Scotia, ZOQe 17, Early Namurian, x29). d Beresella cf. B. polyramosa Kulik
1964 (in Marnet and Rudloff 1972, Ellesmere Island, rnid-Moscovian, x29). e Parachaetetes gar-
woodi (Hinde in Garwood 1912) (in Marnet and Roux 1977, Wath, Great Britain, Zone 9, Upper
Tournaisian, xI2). f Ivanovia tenuissima Khvorova 1946 (in Marnet et al. 1987, van Hauen Pass,
Ellesmere Island, Middle Carboniferous, x8). g Girvanella problematica Nicholson and Etheridge
1878 (in Marnet and Roux 1975b, Cabrieres, France, Zone 16 in" Late Visean, x52). h Fourstonella
sp. (in Marnet et al. 1987, Girty Creek, Ellesmere Island, rnid-Moscovian or younger, xI3).
i Masloviporidium delicata (Berchenko 1982) (in Groves and Marnet 1985, Hale Mountain, Ar-
kansas, Zone 20, Early Bashkirian, xI7). j Windsoporella radiata Marnet and Rudloff 1972 (in
Marnet and Rudloff 1972, Antigonish, Nova Scotia, Zone 17, Early Namurian, x29). kAr-
chaeolithophyllum sp. (in Johnson 1963, Young Co, Texas, Pennsylvanian, x22).IEpistacheoides
connorensis Marnet and Rudloff 1972 encrusting a brachiopod spine (in Marnet and Rudloff 1972,
Connor Lakes, British Columbia. Zone 16sup , Late Visean,xI2).m WetheredellasiluricaWood 1948
(in Marnet and Roux 1983, Jeremiah Hills, Australia, Early Tournaisian. xI7). 0-0 Epimastopora
symetrica (Johnson 1951 )(in Marnet et a1.l987, Little Fish Creek, Yukon Territory, Asselian, x33).
p Eovelebitella robertsi Marnet and Roux 1983 (in Marnet and Roux 1983, Bingleburra, New South
Wales, Australia, Zone 9, Late Tournaisian, x33). q Komia abundans Korde 1951 (in Marnet and
Rudloff 1972, McDarne map area, British Columbia, Zone 18, Early Namurian, x13). r Ungdarella
uralica Maslov 1956 (in Marnet et al. 1987, Girty Creek, Ellesmere Island, Bashkirian, x33).
s Kamaenella tenuis von Moller 1879 (in Marnet and Roux 1980, Denbigh, Great Britain, Zone 15,
Late Visean, x24). t A tractyliopsis weyanti Marnet, Mortelrnans and Roux 1979 (in Marnet and Roux
1983, Old Cannindah, Queensland, Australia, mid-Visean, xI6). u Koninckopora tenuiramosa
Wood 1942 (in Marnet and Roux 1978, Fowler no I Traders, Scott Co, Tennessee, Zone 13, Middle
Visean, x33). v Orthriosiphonoides tenuiramosa Marnet and Rudloff 1972 (in Marnet and Rudloff
1972, Grotto Creek, Alberta, Zone 12, Middle Visean, x12)
382 B. Mamet
Algeria, France, Belgium, Great Britain, Germany, Donbass, Australia, Alaska

(Peratrovich fm), Alberta, British Columbia, Arkansas, Tennessee.
"Aphrostoma" Giirich 1906

Type of genus. Aphrostoma tenerum Giirich 1906. Visean, Belgium.
Diagnosis. Algal-sponge biscuit composed of regular semiplanar layers of,
possibly, dissolved Sphaerocodium, irregularly disposed birdseyes, pelle to ida I
tissue with dissolved Bevocastria and sponges encrusted by Bevocastria.
Suprageneric attributions. Spongiostromidae Giirich 1906. Needs revision.
Stratigraphie range and distribution. Scarce. Visean, Belgium, Poland, Amer-
ican Cordillera.
"A ranea" Hance 1983

Type ofgenus. Aranea eburei Hance 1983, early Visean, Belgium.
Diagnosis. Spider-web of irregular polygonal cells, rounded cystoid cells and
dichotomous tubes.
Suprageneric attributions. Rhodophyte (Solenoporaceae) for Hance (1983).
Problematic nodule formed by different organisms. Bevocastria present in the
holotype.
Stratigraphic range and distribution. Same as type of genus.
9. Archaeolithophyllum Johnson 1956

Type of genus. Archaeolithophyllum missouriense Johnson 1956. Upper Car-
boniferous, Missouri.
Diagnosis. Crustose multifoliate sheets of differentiated cells. Hypothallus
prominent, coaxial, with large polygonal regular cells. Perithallic cells poorly
developed, much smaller, rectangular or square. Conceptacles in the perithallus
(see Shuysky 1987, for illustrations and also Plate 3k).
Archaeolithophyllum delicatum Johnson 1956. Middle and Late Carboniferous,
American midcontinent and Cordillera.
A rchaeolithophyllum donbassicum Kossenko 1972. Visean-Moscovian, Algeria,
Donbass, Germany, Alaska, American Midcontinent (type of the genus Prin-
cipia Brenckle 1982, an Archaeolithophyllum with poorly developed
perithallus).
Archaeolithophyllum johnsoni Racz 1965. Bashkirian-Moscovian, Algeria,
Spain, Canadian Arctic.
Archaeolithophyllum lamellosum Wray 1964. Upper Carboniferous, Kansas.
Archaeolithophyllum va illan i Mamet and Roux 1977. Late Visean-Early
Namurian, France, Spain, Donbass.
Suprageneric attributions. Ancestral coralline for most authors (see Mamet et
a1. (1987) for discussion). Type of new family Archaeolithophyllaceae
Chuvashov 1987.
Stratigraphic range and distribution. Quite common. Oldest reference in
Early Visean of Australia (Mamet and Roux 1983). Forms the "A rchaeolitho-
phyllum community" (Flugel 1977) in assocIatIon with stromatolites and

Tubiphytes from Middle Carboniferous to Early Permian. Frame-builder
in organic build-ups (in association with Neoanchicodium). Also forms "Ar-
chaeolithophyllum oncoid community". Algeria, Morocco, Spain, France,
Germany, Carnic Alps, Donbass, Urals, Russian Platform, Kazakhstan, Sibe-
ria, China, Australia, Arctic Canada, Alaska, American Cordillera, American
midcontinent.
Ardengostella Vachard in Perret and Vachard (1977)

See Dvinella.
"Artophycus" Johnson 1940

Type of genus. Artophycus columnaris Johnson 1940. Upper Carboniferous,
Colorado.
Diagnosis. Described by Johnson as a spongiostromid made of "gently
tapering cones of pillars built of thin, gently arched laminae". Needs
revision.
Suprageneric attribution. Needs revision.
Stratigraphic range and distribution. See type of genus.
10. Asphaltina Mamet in Petryk and Mamet (1972)
Type of genus. Asphaltina cordillerensis Mamet in Petryk and Mamet (1972).
Visean, Alberta.
Diagnosis. Thallus(?) encrusting, composed of large, flexuous, interlaced
cylindrical tubes with oblique partitions. Wall double-layered, a thin external
micritic coating and a very thick pseudo-fibrous inner layer.
Suprageneric attributions. An incertae sedis reported to the tribe Wetheredellae
by Berchenko in Shuysky (1987).
Stratigraphic range and distribution. Recorded from Devonian to Middle
Carboniferous. Very common in North American Cordillera from which it was
originally described in high energy carbonates. Great Britain, France, Siberia
(Kusbass, Kolyma), Canadian Arctic,Alaska, including Peratrovich fm, Yukon
Territory, Alberta, British Columbia, Idaho, Montana, Nevada, Utah, Ok-
lahoma, Kansas, Iowa, Arkansas, Tennessee, Alabama.
11. Asphaltinella Mamet and Roux 1978

Type ofgenus. Asphaltinella horowitzi Mamet and Roux 1978. Tennessee, Early
Namurian.
Diagnosis. Thallus (?) encrusting, composed of interwoven layers of cylindrical
tubes as in Asphaltina (hence the name). Wall single layered.
Asphaltinella? bangorensis Mamet and Roux 1978. Frasnian-Namurian. Mo-
rocco, American Midcontinent, Tennessee.
Asphaltinella peratrovichensis Mamet and Pinard 1983. Early Bashkirian, Pe-
ratrovich fm, Alaska.
384 B. Marnel
Suprageneric attribution. Algae(?) incertae sedis.

Stratigraphic range and distribution. Common in North America in high energy
carbonates. Scarce elsewhere.
Frasnian-Bashkirian. Morocco, Spain, Belgium, Afghanistan, Australia, Can-
adian Arctic, Alaska, Yukon Territory, Alberta, Wyoming, Nevada, Utah,
Arizona, New Mexico, Arkansas, Kansas, Oklahoma, Tennessee, Alabama.
12. Atractyliopsis (Pia 1937) Accordi 1956

Type ofgenus. Atractyliopsis lastensis Accordi 1956. Permian, Italy.
Diagnosis. Thallus acicular or subcylindrical, undivided. Stem undivided,
prominent. Incompletely calcified cortex containing a single row of spherical
cysts? (endospores?).
Atractyliopsis cumberlandensis Rich 1974. Upper Tournaisian-Namurian,
Western Tethys (Morocco, Belgium, France), North America (Tennessee,
Alabama, Alberta).
A tractyliopsis forezi Mametand Roux 1975. Visean, France, Belgium, Australia.
A tractyliopsis minima Mamet and Roux 1978. Visean, Tennessee, Alaska.
Atractyliopsis weyanti Mamet, Mortelmans and Roux 1979. Visean, France,
Belgium, Australia (see Plate 3t).
Suprageneric attributions. Aciculelleae Bassoullet et al. 1979 sensu Berchenko
and Saltovskaja (1987) (see Elliott 1971, for hypothetical ontogeny). Tribe
emended by Deloffre (1988).
Stratigraphic range and distribution. Widespread, but fragile and often broken.
Usually associated with red algae (Archaeolithophyllum) or phylloid algae
(Neoanchicodium). Upper Tournaisian - Upper Permian. Eurasia (Belgium,
France, Hungary, Italy, Yugoslavia, Afghanistan, China, Japan, Sumatra),
North Africa (Libya), Australia, North America (Tennessee, Alabama).
"Baculella" Conil and Dreesen 1985

Type of genus. Baculella gem ina Conil and Dreesen 1985. Devonian-Carbon-
iferous transition, Belgium.
Diagnosis. Chain of pyriform chambers ending by narrow neck. Wall fibrous
radial on a micritic layer, perforated by thin undivided pores.
Suprageneric attribution. Microproblematic. Morphologically similar to the
incertae sed is Saccamminopsis, but with a wall structure similar to the alga
Issinella.
Stratigraphic range and distribution. See type of genus. Reported to form "algal
binds tones" .
13. Beresella (Makhaev? 1939) Maslov and Kulik 1956

Type of genus. Beresella erecta Maslov and Kulik 1956. Moscovian, Russian
Platform (see Plate 2e).
Diagnosis. Calcareous cylindrical tube. rarely dichotomous. Cortex perforated
at regular intervals by short, thin. blind cortical branches. micritized and
forming "dark bands". They are separated by "clear bands" of non-perforated

cortex. Thallus possibly formerly surrounded by mucilaginous coating, now
preserved as a continuous cement layer.
Beresella bilgutayae Giiven~ 1965 (probably a Beresella polyramosa). Bashki-
rian, Turkey.
Beresella herminae Racz 1965. Bashkirian, Spain, Algeria.
Beresella ishimica Kulik 1964. Middle Carboniferous, Russian Platform,
Canadian Arctic.
Beresella polyramosa Kulik 1964. Bashkirian-Sakmarian, Eurasia, Canadian
Arctic, North America (see Plate 3d).
Beresella translucea Kulik 1964. Middle Carboniferous. Russian Platform,
Central Asia.
Suprageneric attributions. Bereselleae Maslov and Kulik 1956.
Beresellaceae (Maslov and Kulik 1956) trans. Shuysky 1985 (and Deloffre 1988)
Concept of Beresellidae as a sponge (Termier, Termier and Vachard 1977) is
untenable.
Stratigraphic range and distribution. One of the most important "reef' builders
in Carboniferous time (baffiestone). First report in Serpukhovian Zone 18.
Common to very abundant in Bashkirian-Moscovian and still widespread up to
the Early Permian. Cosmopolitan: Eurasia, North Africa, Arctic, Nevada, Utah,
Idaho.
14. Berestovia Berchenko 1982

Type of genus. Berestovia filaris Berchenko 1982. Serpukhovian, Donbass.
Diagnosis. Thallus crustose composed of continuous rows of highly calcified
hemispherical to spherical cells. Rows asymmetrical, flattened at base and wavy
at top. Wall yellowish. Morphology of pores, if any, doubtful.
Suprageneric attribution. Linked to the Donezelleae by Shuysky (1987), but
presence of pores is debatable and attribution is unclear.
Stratigraphic range and distribution. Reported only once by Berchenko, but a
common producer of oncoids in the Wahoo Limestone of Alaska (Zone 21). Also
common in Oklahoma, Arkansas, Alabama.
15. Bevocastria Garwood 1931

Type ofgenus. Bevocastria conglobata Garwood 1931. Late Tournaisian, Great
Britain.
Diagnosis. Thallus nodular, formed by meshwork of numerous anastomosed,
branching, micritized tubes. Tubes have constrictions at regular intervals.
Bevocastria amplefurcata Pia 1932. Devonian-Upper Carboniferous. Poland,
Russian Platform, Iran.
Bevocastria fragilis Berchenko 1981 is synonymous with Bevocastria conglobata
Garwood 1931.
Bevocastria kuzbassi Maslov 1956. Famennian-Visean, Kuzbass.
386 B. Marnel
Bevocastria hubbardi Mamet and Roux 1975. Lower Carboniferous, Western

Europe.
Bevocastria uralica Kulik 1973. Serpukhovian, Urals.
Suprageneric attributions. Controversial alga. Codiaceae for Mamet and Roux
1975, Stignomenales for Ischenko 1985, Garwoodiaceae for Luchinina 1987.
Stratigraphie range and distribution. Cambrian (as "Dictyophycus") to Sak-
marian. Although rarely recorded, omnipresent in Devonian-Carboniferous
lagoons. Forms nodules ("Glomus bevocastriosus" Radionova 1976), spon-
giostromids, oncoids and composite nodules with ctenostomates and sponges.
Eurasia, Canadian Arctic, North America.
16. Biumbella Mamet 1970
Type ofgenus. Biumbella braznikhovae Mamet 1970. Devonian-Carboniferous
transition, Donbass.
Diagnosis. Utricular (1) cavity irregularly spherical. Apical opening closed by
conical lid on a neck. Wall two-layered, with a prominent pseudo-fibrous
yellowish layer.
Synonyms. Pseudoumbella Berchenko 1971. Quasiumbelloides Berchenko
1971.
Biumbella donbassica (Berchenko 1971). Tournaisian, Donbass.
Biumbella tenuis (Berchenko 1971). Tournaisian, Donbass.
Biumbella venusta (Berchenko 1971). Tournaisian, Donbass.
Suprageneric attributions. Biumbellidae Berchenko 1974. Umbellaceae Fur-
senko 1959 for Berchenko in Shaikin (1987).
Stratigraphic range and distribution. Scarce. Famennian-Tournaisian. En-
demic to the Donbass.
17. Borisovella Ivanova 1988

Type ofgenus. Borisovella turbinata Ivanova 1988. Serpukhovian, Urals.
DiagnosiS. Thallus cylindrical, with continuous (1) stem. Cortex perforated by
simple branches with very oblique insertion on medulla. Branches numerous,
pedonculated, rapidly expanding and ampuliform.
Suprageneric attribution. Cyclocrinae Pia 1927.
18. Brazhnikovia Berchenko 1981

Type ofgenus. Brazhnikovia undata Berchenko 1981. Devonian-Carboniferous
boundary, Donbass.
Diagnosis. A slightly tapering kamaenid with short irregular partitions. Mor-
phology of branches is unclear.
Suprageneric attributions. Originally linked to the Palaeobereselleae by Ber-
chenko (1981), then attributed to the Anthracoporellopsiae Shuysky 1985 by
Shuysky (1987). Genus probably valid, but restudy necessary as nature of the
branches is poorly known.
"Buzgulella" Korde 1951

Type ofgenus. Buzgulella serrata Korde 1951. Middle Carboniferous, Urals
Diagnosis, attribution and stratigraphy. Need revision.
19. Cabrieropora Mamet and Roux 1975

Type of genus. Cabrieropora pokornyi Mamet and Roux 1975. Late Visean
(Zone 16 j ), France.
Diagnosis. Thallus long, cylindrical, composed or regular barrel-shaped arti-
cles. Expansion of medulla forms important spherical vestibule. Tufts of six
divergent branches, each thick and phloiophore.
Suprageneric attributions. Velebitelleae (Vachard 1977) Mamct and Roux
1983. Incorrectly attributed to Linoporallinae Pia 1927 by Shuysky (1987).
Stratigraphic range and distribution. Scarce. Upper Visean (Zones 14 to 16)
France, Spain.
20. Calcifolium Schvetsov and Birina 1935

Type of genus. Calcifolium okense Schvetsov and Birina 1935. Russian
Platform.
Diagnosis. Thallus composed of a succession of tubular siphons, rapidly ex-
panding into cups or petals. Filaments occupy whole thickness of petals and
branch dichotomously (see Plate 2j).
Calcifolium bruntonense Johnson 1958 (identical to Calcifolium okense).
Calcifolium punctatum Maslov 1956. Latest Visean (Zones 16 j and 16.), Great
Britain, Poland, Russian Platform, Donbass, Urals.
Supragenericattributions. Codiaceae (Mametand Roux 1977; Skompski 1981).
Type of new tribe Calcifoliae Shuysky 1987, placed among Anchicodiaceae
Shuysky 1987. Attribution to sponges by Termier et al. (1977) (as Calcifoliida)
is unwarranted.
Stratigraphic range and distribution. Calcifolium is often referred to
"phylloids", although there is no proof of a leafiike habit. A good stratigraphic
marker for the Visean-Namurian boundary. Occidental part of the Tethys:
Algeria, Morocco, Spain, Great Britain, France, Poland, Russian Platform,
Donbass, Urals, Tadzhikhistan.
21. Calcisphaera Williamson 1881

Type ofgenus. Calcisphaera laevis Williamson 1881. Late Visean, Great Britain.
(Type designation by Andrews 1955). Subsequent designation of Calcisphaera
cancellata Williamson 1881 by Va chard (1977) is questionable.
Diagnosis. Microcrystalline, hollow, calcite spherule. Wall one-layered. Pores
(?) scarcely noticeable. No known functional aperture.
Synonymy. Granulosphaera Derville 1931 OBJ, Pachysphaera Rauzer-Cher-
noussova 1948, OBJ, Pachysphaera Conil and Lys 1964 OBJ, Pachysphaerina
Conil and Lys 1967 OBJ.
388 B. Mamet
Calcisphaera pachysphaerica (Pronina 1963). Visean-Baschkirian, cosmo-

politan.
Suprageneric attributions. Controversial since its origin. Kazmierczak (1976)
and Vachard and Tellez-Giron (1986) have proposed volvocale affinity for the
Calcisphaeridae. Most Russian authors still attribute genus to foraminiferal
family Calcisphaeridae Reitlinger 1960 OBJ. The debate exists since Wil-
liamson who thought (1881, p. 525) that they were either "reproductive capsules
of some marine form of vegetation" or "some type of extinct Protozoa". Recent
authors have shown their resemblance with dasycladacean reproductive cysts
(Wray 1977a). Mamet 1973 has illustrated an in situ crown of Calcisphaera
refuting a protozoan origin. Loeblich and Tappan (1988) list them among the
taxa erroneously regarded as foraminifers.
Stratigraphic range and distribution. Middle Devonian-Middle Carboniferous.
Extremely abundant in Tournaisian-Visean lagoons. Easily dispersed
throughout entire carbonate platforms. "Calcispherites" are quite similar
to the Late Carboniferous-Permian "algen sporen assemblages" of Flugel
(1977). Cosmopolitan. Eurasia, North Africa, Australia, Arctic, North
America.
Chabakovia Vologdin 1939

See Renalcis.
"Chantonia" Termier, Termier and Vachard 1977

Type of genus. Ungdarella maslovi Chanton-Guven~ 1965. Late Visean,
Algeria.
Diagnosis. Needs revision as original material is poorly preserved. Seems to be
a mixture of Ungdarella, Chuvashovia and Epistacheoides.
Suprageneric attributions. The type described as a red alga, was transferred to
the "sponges" (Aoujgaliida, Termier et a1. 1977).
Stratigraphic range and distribution. Needs revision.
"Chondrostoma" Gurich 1906

Type ofgenus. Chondrostoma problematicum Gurich 1906. Visean, Belgium.
DiagnOSiS. Needs revision. The four species described by Gurich (the type
Chondrostoma problematicum, Chondrostoma globuliferum, Chondrostoma in-
termixtum and Chondrostoma vermiculiferum) show various cyanophyte mats,
more or less bioturbated, some in situ, some reworked, some with mud cracks
and algal chips. Faecal pellets, hard lumps, algal pellets and in situ sponge
spicules are present.
Suprageneric attribution. Spongiostromidae Gurich 1906.
Stratigraphie range and distribution. Needs revision.
"C/aracrusta" Vachard in Va chard and Montenat (1981)

Type of genus. Girvanella catenoides Homann 1972. Permian, Carnic Alps.
Diagnosis, suprageneric attribution, stratigraphic range and distribution. Needs

revision. Most Middle Carboniferous forms are probably Berestovia.
22. Clavaporella Kochansky-Devide and Herak 1960

Type of genus. Clavaporella caliciformis Kochansky-Devide and Herak 1960.
Permian, Yugoslavia.
Diagnosis. Thallus cylindrical, unramified, a succession of calyx-shaped
asymmetrical euspondyl units. Composite vestibule followed by tufts of
phloiophore branches. Stem isodiametrical, cylindrical, unsegmented. Three to
four rows of simple pores form verticils.
Clavaporella reinae Racz 1966. Moscovian-Upper Carboniferous, Spain, Urals,
Canadian Arctic, (see Plate 2a,b,c).
Suprageneric attributions. Albertaporelleae (Giiven<s 1979) Mamet and Roux
1981 for Deloffre 1988. Triploporellinae (Pia 1920) Bassoullet et al. 1979
according to Shuysky (1987).
Stratigraphic range and distribution. Moscovian (scarce) to Permian
(abundant). Spain, Yugoslavia, Turkey, Caucasus, Urals, Afghanistan, China,
Canadian Arctic.
Coactilum Maslov 1956

See Sphaerocodium.
"Codonophycus" Fenton and Fenton 1939

Type of genus. Codonophyc1,ls austinii Fenton and Fenton 1939. Tournaisian,
Wyoming.
Diagnosis, suprageneric attribution and stratigraphic range. Needs revision
(algal biscuit).
23. Coelosporella Wood 1940

Type ofgenus. Coelosporella wetheredii Wood 1940. Late Visean, Great Britain.
DiagnOSiS. Thallus cylindrical, with prominent unsegmented medulla. Poorly
calcified cortex with one or two rows of lemon-shaped cavities ("sporangial"
cavities) open to the outside.
Coelosporella jonesii Wood 1940. Upper Visean, Great Britain, Belgium,
Donbass, Urals, Tennessee.
Suprageneric attributions. Aciculelleae Bassoullet et al. 1979 for Deloffre (1988)
and Berchenko and Saltovskaja (1987). See Elliott (1971) for hypothetical
ontogeny and function of rod.
Stratigraphic range and distribution. Quite scarce and fragile. Late Tournaisian
to Serpukovian. Spain, Great Britain, Belgium, Germany, France, Donbass,
Ural, Nepal, American Cordillera and midcontinent, Tennessee, Alabama,
Newfoundland.
390 B. Mamet
24. Columbiapora (Mamet 1974) Mamet and Roux 1981b

Type of genus. Columbiapora johnsoni Mamet 1974. Middle Tournaisian,
British Columbia (see Plate la).
Diagnosis. Thallus, a succession of subspherical, verticillated units. Medulla
sub cylindrical, with some expansion at junction ofverticils. Tufts of six fusiform
branches diverge from a short composite vestibule, a cylindrical expansion of
the medulla.
Suprageneric attributions. Incorrectly attributed to the Gyroporelleae Pal 1976
emendBassoulletetal. 1979 by Shuysky (1987). Albertaporelleae (Giiven<s 1979)
Mamet and Roux 1981 b for Deloffre (1988).
Stratigraphic range and distribution. Middle to Late Tournaisian. Endemic to
the American Cordillera (Columbiapora mameti Vachard 1977 from the Ser-
pukovian of France does not belong to the genus).
Coni/alia Vachard in Massa and Vachard (1979)

See Cuneiphycus.
25. Connexia Kochansky-Devide 1970

Type of genus. Connexia fragilis Kochansky-Devide 1970. Moscovian,
Yugoslavia.
Diagnosis. Thallus a succession of claviform units. Central medulla cylindrical,
unsegmented. Each unit is stellate with five to six bifid projections ("horns").
Branches with second and third order dichotomy.
Suprageneric attribution. Triploporelleae (Pia 1920) Bassoullet et al. 1979.
Stratigraphic range and distribution. Both Deloffre (1988) and Shuysky (1987)
restrict the genus to the Permian. However, the type is Moscovian and the correct
range is Moscovian-Early Permian as reported by Milanovic (1982). Yugos-
lavia, Central Asia.
Contortoporidium Maslov 1973 NOM. NUD.

See Masloviporidium.
"Corydopodium" Derville 1931

Type of genus. Corydopodium pruvosti DerviJle 1931. Visean, France.
DiagnOSiS. Flattened algal biscuit composed of birdseyes (sponge canals?)
interspersed with Bevocastria tubes and algal peloids of unknown affinity.
Suprageneric attribution. Spongiostromidae.
Stratigraphic range. Scarce. Lower Carboniferous, Great Britain, France.
Belgium, Germany.
C ostatumbella Berchenko 1974

See Eoumbella Platonov 1974
26. Cribrokamaena Brenckle 1985

Type ofgenus. Cribrokamaena citrosa. Brenckle 1985. Visean, Nevada.
Diagnosis. Thallus (?) slender, cylindrical, branching. Medulla divided by
regularcribrated pseudo-septa. Wall hyaline pseudo-fibrous, perforated by thin
pores.
Note. Part of the original material may belong to Issinella. Cribrated partitions
are unreported among Palaeosiphonocladales.
Cribrokamaena furcillata Brenckle 1985. Visean, Nevada, Missouri.
Suprageneric attribution. At origin, Palaeoberesellaceae? Mamet and Roux
1974.
Stratigraphic range and distribution. Endemic to North America. Common in
American Cordillera (Alberta, British Columbia, Wyoming, Montana, Utah,
Nevada, Colorado), American midcontinent (Iowa, Missouri, Illinois) and
Tennessee-Ala bama.
Cribroporidium Maslov 1973 NOM. NUD.

See Masloviporidium Groves and Mamet 1985.
27. Cribrosphaeroides Reitlinger 1959

Type of genus. Cribrosphaera simplex Reitlinger 1954. Frasnian, Russian
Platform.
Diagnosis. Thallus? ovoid, irregular. Wall micritized, perforated by coarse,
regular straight "pores". No aperture.
Carboniferous species.
Cribrosphaeroides ovalis Poyarkov 1961. Frasnian to Devonian-Carboniferous
transition. Donbass, Urals, Siberian Platform, Tian-Shan, Turkey.
Cribrosphaeroides robusta (Mikluko-Maklai 1965). Givetian to Devonian-
Carboniferous transition. Urals, Siberian Platform, Tian-Shan, Turkmenistan,
Omolon Massif.
Cribrosphaeroides simplex donica Brazhnikova and Rostovceva in Aizenberg
and Brazhnikova 1966. Devonian-Carboniferous transition, Donbass.
Suprageneric attributions. Controversial. Cribrosphaeroides (and Auroria)
could be foraminifers? (Cribrosphaerinae Zadorozhnyi and Yuferev 1984), but
they lack aperture and their "pores" normally indicate dasyclad branches.
Stratigraphie range and distribution. Eifelian to Tournaisian. Belgium, Poland,
France, Germany, Donbass, Russian Platform, Urals, Kazakhstan, Tadhzik-
histan, Turkmenia, Tian-Sian, Siberian Platform, Kuzbass, China, Omolon
Massif.
28. Cummingsella Mamet and Roux 1980

Type of genus. Cummingsella lyoncrossi Mamet and Roux 1980. Lower
Namurian, Scotland (see Plate 2b).
392 B. Marne!
Diagnosis. Thallus, succession of barrel-shaped articles. Medulla cylindri-

cal, unsegmented. Branches (?) or cortical tubes (?) numerous, as random
"verticils" (?)
Dichotomy of first to second order depending on position of branch.
Cummingsella bingleburrae Mamet and Roux 1983. Late Tournaisian,
Australia.
Suprageneric attributions. Undetermined tribe among the Dasycladales for
Deloffre (1988). Dasycladacean? or codiacean? for Mamet and Roux (1980).
Could be a Udoteacean?
Stratigraphic range and distribution. Very scarce. Only two Lower Carbonife-
rous occurrences.
29. Cuneiphycus Johnson 1960

Type of genus. Cuneiphycus texana Johnson 1960. Middle Carboniferous,
Texas.
Diagnosis. Thallus tapering, more or less encrusting, without hypo- and pe-
rithallic differentiation. Rows of regular, thin, wedge-shaped cells. Thallus often
difficult to separate from Fourstonella. Concept needs revision.
Synonym. Eflugelia, Coni/alia.
Cuneiphycus aliquantulus Johnson 1960. Visean-Moscovian, Algeria, Spain,
Belgium, Donbass, Siberia, Canadian Arctic, New Mexico, Texas.
Cuneiphycus johnsoni Flugel 1966. Visean-Permian, Carnic Alps, Urals, Can-
adian Arctic, Idaho, Kansas.
Cuneiphycus texana Johnson 1960. Visean-Upper Carboniferous, Western
Tethys, (Morocco, Algeria, Great Britain, Russian Platform, Donbass), North
America (Alaska, Texas, Oklahoma).
Suprageneric attributions. Undetermined rhodophycophyte family for Mamet
et al. (1987). New tribe Mametelleae Ch uvashov 1987. Attributed to the sponges
by Termier et al. (1977) (as Ptychocladiidae Ellias 1950).
Stratigraphic range and distribution. Abundant. Long ranging from Visean to
Permian. Libya, Algeria, Morocco, Spain, Great Britain, France, Donbass,
Siberia, Urals, Iran, Afghanistan, China, Alaska, Idaho, American midcon-
tinent, Texas, New Mexico.
30. Cylindrofolia Brenckle and Groves 1987.

Type ofgenus. Cylindrofolia glenisteri Brenckle and Groves 1987. Tournaisian,
Iowa.
Diagnosis. Thallus unsegmented, with continuous cylindrical medulla.
Cortex poorly calcified, perforated by horizontal, wavy, irregular, frond-like
appendages.
Suprageneric attribution. Incertae sedis. As noted by Brenckle and Groves,
there are no known horizontal processes among bona fide dasyc1adales. This
puzzling genus has some similarities with Menselina, another Carboniferous

microproblematicum.
31. Dasyc/adophycus Elliott 1982

Type of genus. Dasyc/adophycus ensomi Elliott 1982. Mid-Visean, Great
Britain.
Diagnosis. Non-calcified central stem with verticils of four branches. Each
branch divides into four short branch lets.
Suprageneric attribution. Uncalcified Dasycladales for Elliott (1982).
Dasyporella Stolley 1893

Genus restricted to the Lower Palaeozoic, but often mentioned in the Visean-
Serpukhovian by Russian authors (e.g. Dasyporella kulikae Muromtseva 1979,
Dasyporella maslovi Kulik 1973, Dasyporella popovae Kulik 1973, etc.). These
taxa ought to be transferred to genuine Carboniferous Dasyporelleae and
Uraloporelleae (Anthracoporella, /ssinella, Nanopora, etc.).
32. Dokutchaevskella Berchenko 1981

Type of genus. Dokutchaevskella inaequalis Berchenko, Devonian-Carbon-
iferous transition, Donbass.
Diagnosis. Thallus long, slender, cylindrical, unsegmented. Medulla div-
ided by uneven projections of the cortex, forming irregularly-shaped cells.
Branches thin, straight, perpendicular or slightly oblique to the medulla.
Needs further study. Seems quite similar to Pseudonanopora Mamet and
Roux 1975.
Suprageneric attributions. Dasyporelleae Pia 1927 for Berchenko (1981). Ex-
votariselleae Shuysky 1985 for Shuysky (1987).
Stratigraphic range and distribution. Devonian-Carboniferous transition,
Donbass and Southern Urals. Needs revision.
33. Donezella Maslov 1929 non sensu Vachard in Meissami et al. 1978
Type of genus. Donezella lutugini Maslov 1929. Middle Carboniferous,
Donbass.
Diagnosis. Thallus cylindrical, slightly tapering, tubular, sinuous, constricted
and ramified in all directions. Medulla divided by thin irregular projections of the
cortex. Cells grossly rectangular; height greater than width. Cortex perforated
by blind pores, thin, unramified, forming dark micritic bands from one partition
to the other. During life, thallus may have been covered by thin mucilagenous
coating, now preserved as clear cement (see Plate 2i).
Synonyms. Goksuella Giiven<s 1965 and probably Praedonezella Kulik 1973.
394 8. Mamet
Donezella cespaeformis (Kulik 1973). Latest Visean, basal Bashkirian. Spain,

France, Donbass, Urals.
Donezella lunaensis Racz 1964 is probably synonymous with Donezella lutugini.
Suprageneric attributions. Palaeosiphonocladales of the tribe Donezelleae as
emended by Shuysky (1985). Maslov's original description was very sketchy and
has caused much taxonomic controversy. As with many Palaeozoic siphono-
cladales, this genus has been attributed to foraminifers, sponges, and
pseudosponges.
Stratigraphic range and distribution. Very abundant in the Middle Carbon-
iferous. Forms Donezella baftlestones (Mamet et al. 1979). Latest Visean-
Namurian (scarce), Bashkirian-Moscovian (very abundant). Late Carbonife-
rous-Early Permian (scarce). Cosmopolitan in the Northern Hemisphere.
Forms baftlestones and mud-mounds in Eurasia, North Africa and North
America.
Dromastacheoides Vachard in Perret and Vachard (1977)

See Epistacheoides.
34. Dutroella Mamet and Roux 1978

Type ofgenus. Dutroella scotti Mamet and Roux 1978. Late Visean (Zone 16d,
Tennessee.
Diagnosis. Thallus, a succession of thin barrel-shaped units. Medulla thick.
Cortex perforated by a single row of perpendicular, simple, straight, phloio-
phore branches. Needs further study.
Suprageneric attribution. Unconvincingly placed with the Mizziinae Bas-
soullet et al. 1979 by Deloffre (1988). Should be placed among the Diplopo-
raceae.
35. Dvinella Khvorova 1949

Type of genus. Dvinella comata Khvorova 1949. Middle Carboniferous, Rus-
sian Platform (see Plate 2f).
Diagnosis. Thallus, a long, cylindrical calcareous tube. Cortex perforated by
blind pores, oblique and dichotomous, forming a triangular "dark ring",
separated by cement filled "clear rings". During life, thallus may have been
covered by a thin mucilagenous coating, now preserved as clear cement.
Three subgenera have been proposed, but they are often difficult to separate:
Dvinella (Dvinella) Khvorova 1949 has branches with single dichotomy.
Dvinella (Trinodella) Maslov and Kulik 1956 has pores with double dichotomy.
The subgenus Dvinella (Ardengostella) Vachard in Perret and Va chard 1977 is
a Dvinella in Khvorova's sense.
Dvinella (Trinodella) aequalis Kulik 1964. Moscovian, Russian Platform.
Dvinella (Trinodella) bi{urcata Maslov and Kulik 1956. Bashkirian-Lower
Permian, Spain, Russian Platform, Canadian Arctic (Plate 2g).

Dvinella (Dvinella) crassitheca Kulik 1964. Moscovian, Hungary, Yugoslavia,
Russian Platform, Canadian Arctic.
Dvinella cuvillieri GUven<s 1965. Moscovian, Turkey.
Dvinella (Dvinella) distorta Kulik 1964. Moscovian, Russian Platform.
Dvinella (Dvinella) gracilis Kulik 1964. Moscovian, Russian Platform, Can-
adian Arctic.
Dvinella (Trinodella) varialonga Kulik 1964. Moscovian, Russian Platform.
Suprageneric attributions. Bereselleae Maslov and Kulik 1956.
Beresellaceae (Maslov and Kulik 1956) trans. Shuysky 1985 (and by Deloffre
1987).
Stratigraphic range and distribution. Abundant and very important for car-
bonate production. Forms "baffiestones" in mud-mounds and in "reefs".
Namurian-Bashkirian up to Early Permian. Cosmopolitan in the Northern
Hemisphere. Eurasia, North Africa, Canadian Arctic, North America.
Eflugelia Vachard in Massa and Vachard (1979)

See Cuneiphycus. See also Fourstonella.
36. Einorella Saltovskaja 1984

Type of genus. Einorella globosa Saltovskaja 1984. Middle Carboniferous,
Tadhzikistan.
Diagnosis. Thallus, a slender calcareous tube, rarely dichotomous, with regular
constrictions at each internal pseudoseptation. Cortex perforated by thin blind
pores forming micritized "black rings". Unperforated parts form thin "clear
bands". Thallus may originally have been covered by a mucilaginous coating,
now preserved as a continuous cement layer. Needs further study.
Suprageneric attributions. Bereselleae Maslov and Kulik 1956. Beresellaceae
(Maslov and Kulik 1956) trans. Shuysky 1985 (and Deloffre 1987).
37a. Elenia Poyarkov 1965

Type of genus. Umbella famena Bykova in Bykova and Polen ova (1955).
Famennian, Russian platform.
Diagnosis. Utricular (?) spherical cavity. Parallel longitudinal ribs. No basal
projection. Aperturallid unknown. Wall two-layered.
Synonym. Spinumbella Platonov 1974
Elenia costulata Berchenko 1974. Devonian-Carboniferous transition,
Donbass.
Elenia formosa Berchenko 1974. Devonian-Carboniferous transition, Donbass.
Elenia spinosa (Conil and Lys 1964) Devonian-Carboniferous transition,
Belgium.
Suprageneric attributions. Umbellaceae Fursenko 1959.
396 B. Mamet
Stratigraphic range and distribution. Mainly Upper Devonian (Frasnian-

Famennian) up to the Devonian-Carboniferous transition. Restricted to the
western Tethys. Belgium, France, Poland, Russian Platform, Urals, Donbass.
37b. Eoc/ypeina sensu Poncet 1987 non Vachard 1985

Type ofgenus. Eoc/ypeina crassa Poncet 1987. Moscovian, Algeria.
Diagnosis. Thallus, a succession of glass- or basket-shaped segments. Primary
branches in verticils, of even diameter, first oblique to the axial cavity, then
horizontal at their distal part.
Suprageneric attributions. Originally described as Dasycladaceae. Clypeineae
(Elliott 1968) Bassoullet et al. 1979 for Deloffre (1988).
38. Eokoninckopora Saltovskaja 1984

Type of genus. Eokoninckopora einori Saltovskaja 1984. Tournaisian,
Tadzhikistan.
Diagnosis. Thallus (?) very irregular, vermiporellid-shaped. Cortex perforated
by very thick polygonal to sub polygonal pores. Wall structure unclear.
Suprageneric attribution. Linked to the Cyclocrinae Pia 1927 by Shuysky (1987)
although there appears to be no relation with Koninckopora. Needs further
study. Could be a partially dissolved bryozoan.
39. Eolithoporella Johnson 1966

Type of genus. Eolithoporella dawsoni Johnson 1966, Lower Carboniferous,
Alberta.
Diagnosis. Reported by Johnson as a thin crust of vertically elongated cells
resembling Litostroma.
Eolithoporella europaea (Kochansky-Devide 1970) (as Litostroma), Moscovian,
Yugoslavia.
Suprageneric attribution. Johnson thought it belonged to an undetermined
family among the red algae. It could very well be a megafaunal fragment.
Stratigraphie range and distribution. Scarce. Lower to Middle Carboniferous.
Alberta, Yugoslavia.
40. Eoumbella Platonov 1974

Type ofgenus. Umbella ollaria Bykova in Bykova and Polenova (1955). Upper
Devonian, Russian Platform.
Diagnosis. Utricular(?) cavity spherical. Coarse external irregular ribs. Ex-
tensive basal projection as in Umbellina. Apertural lid unknown. Wall
two-layered.
Eoumbella slriatella (Berchenko 1974). Devonian-Carboniferous transition,
Donbass.
Suprageneric attribution. Umbellaceae Fursenko 1959.

Stratigraphic range and distribution. Mainly Upper Devonian, but ranges into
the Devonian-Carboniferous transition. Russian Platform, Urals, Donbass,
Siberian Platform.
41. Eouraloporella Berchenko 1981

Type of genus. Eouraloporella kordeae Berchenko 1981. Devonian-Carbon-
iferous transition, Donbass.
Diagnosis. Thallus long, slender, cylindrical, unsegmented with central stem
conspicuous and occasional pseudoseptations. Pores functional, thin, straight,
aspondyl, at right angle to the medulla. Needs revision.
Suprageneric attribution. In spite of its name, and attribution to the Uralopo-
rellaeae by Shuysky (1985), the taxon is not a beresellid as the pores are
functional.
42. Eovelebitella (Vachard 1974) emend. Mamet and Roux 1983

Type of genus. Eovelebitella occitanica (Vachard 1974) emend. Mamet and
Roux 1983. Late Visean, Zone 16 j , France. (Note: The original material
comprises two genera, Eovelobitella sensu stricto as redefined and laphetella
Mamet and Roux 1983) (see Plates Ic and 2m).
Diagnosis. Thallus metaspondyl, a succession of subspherical articles.
Medullar segments biconvex, with medullar projections forming a thick ves-
tibule. Branches phloiophore, in thick dense tufts (about 6-8 branches in a
random thin section, probabJe around thirty in three dimensions).
Eovelebitalla robertsi Mamet and Roux 1983, Tournaisian-Visean boundary,
Australia (Plate 3p).
Suprageneric attribution. Diploporeae Pia 1920, emend. Giiven~ 1979, non
sensu Bassoullet et al. (1979).
Stratigraphic range and distribution. Locally abundant. Tournaisian and Vi-
sean. France, Spain, Australia.
43. Epimastopora ("Pia 1922") (emend. Korde 1951) sensu Roux 1979.
Type of genus. Epimastopora piai Korde 1951 (non piae Bilgiitay 1960). Late
Carboniferous, Urals (see Plate 2n).
Diagnosis. Thallus cylindrical, with very large stem. Cortex perforated by
dumb-bell shaped branches. In equatorial sections, branches are subpolygonal
to spherical.
See Roux (1979) for discussion of type and elucidation of the taxonomy.
Synonym. Globuliferoporella Chuvashov 1974.
Epimastopora primae va (Chuvashov and Anfimov 1988). Moscovian, Urals.
Epimastopora symetrica (Johnson 1951) (as Gyroporella). Upper Carbonifer-
ous-Permian. Carnic Alps, Urals, Canadian Arctic, Texas, see Plate 3n-o.
398 B. Marnet
Suprageneric attributions. Mastoporeae (Pia 1920) emend. Deloffre 1988.

Cyclocrineae Pia 1927, Mastoporinae Pia 1927 for Shuysky (1987).
Stratigraphic range and distribution. Important and abundant. Forms the
"Epimastopora community" in association with Gyroporella and M izzia in outer
shelf facies. Upper Carboniferous-Permian. Spain, Carnic Alps, Yugoslavia,
Turkey, Urals, Japan, Canadian Arctic, Yukon Territory, Texas.
44. Epimastoporella Roux 1979

Type of genus. Epimastopora japonica Endo 1951. Early Permian, Japan,
Plate 2p.
Diagnosis. Thallus cylindrical, with large, important medulla. Cortex perfo-
rated by ovoid branches, perpendicular to the cavity. In equatorial section,
branches are sub polygonal to spherical.
See Roux (1979) for discussion. (Partially Epimastopora auctores and
Pseudoepimastopora auctores).
Epismastoporella bodoniensis (Racz 1965) (as Epimastopora). Moscovian, Spain.
Epismastoporella camasobresensis (Racz 1966) (as Epimastopora). Moscovian,
Spain.
Epismastoporella hunzaensis (Zan in Buri 1965) (as Epimastopora). Upper
Carboniferous-Permian, Carnic Alps, Pakistan, Canadian Arctic.
Epismastoporella iwaizakensis (Endo 1953) (as Epimastopora). Upper Car-
boniferous-Permian, Carnic Alps, Turkey, Japan, Canadian Arctic.
Epimastoporella japonica (Endo 1951) (as Epimastopora). Upper Carbonifer-
ous-Permian, Carnic Alps, Yugoslavia, Turkey, Japan, Canadian Arctic.
Epimastoporella macropora (Maslov 1956) (as "Koninckopora"). Serpuk-
hovian-Bashkirian, France, Donbass, Urals, Tian-Shan.
Epimastoporella micropora (Maslov 1956) (as "Koninckopora'). Serpuk-
hovian-Bashkirian, Donbass, Urals.
Epimastoporella rolloensis (Racz 1965). (as Epimastopora). Moscovian-Per-
mian, Spain, Urals.
Suprageneric attribution. Mastoporeae (Pia 1920) emend. Deloffre 1988.
Stratigraphic range and distribution. First occurrence, Middle Carboniferous.
Abundant, Upper Carboniferous and Permian. Tunisia, Spain, Carnic Alps,
Turkey, Russian Platform, Urals, Pakistan, Malaya, Japan, Canadian Arctic.
45. Epiphyton Bornemann 1886

Type of genus. Epiphyton flabella tum Bornemann 1886. Cambrian, Sardinia.
DiagnOSiS. Thallus branching, composed of highly dichotomous tufts of stems.
Original cellular microstructures controversial.
Epiphyton buldyricum Antropov 1955 (synonym, Epiphyton aegeensis Guven«
1981) Frasnian-Tournaisian. Turkey, Russian Platform, Urals. China.
Suprageneric aflribulions. Controversial. Attributed to red, green and blue-
green algae (see Saltovskaja 1984, for discussion). Placed by Luchinina (1987)
as Epiphytaceae Korde 1959 among the Hormogonophyceae (Geitler) Elenkin

1934.
Stratigraphic range and distribution. Reported from the Proterozoic to the
Carboniferous. Important in Cambrian archaeocyathid mounds (nearly one
hundred "species" in the literature). Still abundant in Devonian reefs. Scattered
at the Devonian-Carboniferous transition and in the Tournaisian where it
becomes extinct.
46. Epistacheoides Petryk and Mamet 1972

Type of genus. Epistacheoides nephroformis Petryk and Mamet 1972. Visean,
Alberta. (Synonym: Dromastacheoides wilsoni Vachard in Perret and Vachard
1977) (see Plate II).
Diagnosis. Thallus encrusting, crustose, irregular. Hypothallic tissue poorly
calcified, vesicular, surrounded by two or three rows of strongly calcified
perithallic rectangular cells. Wall hyaline, yellowish calcite.
Synonyms. Dromastacheoides Vachard in Perret and Vachard 1977 and
Roquesselsia Termier et al. 1977.
Epistacheoides chan toni Mamet and Roux 1977 (synonym: Roquesselsia radians
Termier et al. 1977). Upper Visean, Algeria, France, Ireland.
Epistacheoides connorensis Mamet and Rudloff 1972 (synonym; Dromasta-
cheoides topi Vachard 1981). Visean-Namurian (Asselian-Sakmarian in the
Arctic). Algeria, Morocco, Great Britain, Belgium, France, Urals, Afghanistan,
China, Alaska, Canadian Arctic, British Columbia, Tennessee, Newfoundland
(see Plate 31).
Suprageneric attributions. Stacheiinae Loeblich and Tappan 1961. New tribe
Pseudostacheoidaea Chuvashov 1987. Erroneously attributed to the sponges by
Vachard (1977) (as Aoujgaliida).
Stratigraphie range and distribution. Very common in the Visean-Namurian-
Baschkirian. Algeria, Morocco, Spain, France, Belgium, Great Britain, Ger-
many, Urals, Afghanistan, China, Canadian Arctic (where is reaches Early
Permian), Yukon Territory, Alaska, British Columbia, Alberta, Tennessee,
Newfoundland.
47. Eugonophyllum Konishi and Wray 1961

Type of genus. Eugonophyllum johnsonii Konishi and Wray 1961. Late Car-
boniferous, New Mexico.
Diagnosis. Thallus composed of large blades, standing erect on substratum.
Medulla with dichotomous filaments (?) often obliterated by early cemen-
tation. Subcortex contains anastomosed crescent-shaped utricles (?) com-
municating through the cortex by fine perpendicular branchlets. (Synonym.
Could be similar to Paradella Maslov 1956 which has priority. However
Maslov's taxon is ill defined and Eugonophyllum has been extensively used
since its erection).
400 B. Marnet
Eugonophyllum johnsonii uralicum Rauser-Chernoussova and Korolyuk 1981.

Moscovian, Urals.
Eugonophyllum magnum (Endo 1951). Middle Carboniferous- Lower Permian,
Yugoslavia, Turkey, Japan.
Eugonophyllum mulderi Racz 1965, Moscovian, Spain, Yugoslavia.
Suprageneric attributions. "Phylloid algae" (vernacular). Anchicodiaceae
Shuysky 1987, new tribe Ivanoviae Shuysky 1987.
Stratigraphic range and distribution. Important and widespread. For a recon-
struction of a phylloid algal community, see Toomey and Babcock (1983).
Forms mounds in Moscovian-Sakmarian. Spain, Carnic Alps, Turkey, Urals,
Kazakhstan, Japan, Canadian Arctic, Kansas, Texas, New Mexico.
48. Evlania Bykova 1952

Type of genus. Evlania transversa Bykova 1952. Upper Devonian, Russian
Platform.
Diagnosis. Thallus(?) attached, irregularly shaped. Internal cavity divided by
pseudo-septa. Wall hyaline, yellowish, perforated by numerous dichotomously
divided pores. Needs revision.
Suprageneric attributions. Controversial. Variously interpreted as foraminifer,
sponge or alga.
Stratigraphie range and distribution. Givetian-Visean. Morocco, France, Bel-
gium, Czechoslovakia, Urals, Donbass, Russian Platform, Alberta, Iowa.
49. Exvotarisella Elliott 1970

Type of genus. Nodosinella index Ehrenberg sensu von Moller 1879 (synonym:
Exvotarisella maponi Elliott 1970). Late Visean, Russian Platform (see Plate 3b).
DiagnOSiS. Thallus, a long cylindrical tube. Medulla divided by irregular
cavities formed by thick pseudosepta. Branchlets thin, numerous, aspondyl.
Second to third order ramifications.
Exvotarisella dili Vachard in Oil et al. 1977. Visean. Turkey.
Suprageneric attributions. Tribe Exvotariselleae Shuysky 1985 according to
Shuysky (1987). Palaeobereselleae Mamet and Roux 1974 for Deloffre (1988).
Stratigraphic range and distribution. Scarce in Devonian-Tournaisian. Abun-
dant in Late Visean-Namurian. In association with other palaeoberesellids,
forms baffiestones in protected lagoons. Algeria, Morocco, Spain, Great Britain,
France, Belgium, Germany, Poland, Turkey, Russian Platform, Urals, Donbass,
Kuzbass, Arctic Siberia, Omolon, Australia. Alberta, British Columbia.
Newfoundland.
50. Fasciella Ivanova 1973

Type o[genus. Fasciella kizilia Ivanova 1973 (synonyms: Shartymophycus [usus
Kulik 1973. Fasciella ivanovae Saltovskaja 1984. and Fasciella ramosa Sal-
tovskaja 1984). Late Visean-Serpukhovian. Urals.
Diagnosis. Thallus encrusting, composed of irregular concentric layers of

elongated, undulating cells. Recrystallization often obscures shape of cells.
Walls calcareous, yellowish, hyaline.
Suprageneric attributions. Incertae sedis. Could be a chlorophyte or a
rhodophyte.
Stratigraphic range and distribution. Very common in high energy open marine
grainstones. Visean-Bashkirian. Algeria, Morocco, Spain, Ireland, Great Brit-
ain, France, Belgium, Germany, Poland, Russian Platform, Urals, Donbass,
Tadzhikhistan, China, Alaska, Idaho, Quebec.
51. Foliophycus Johnson 1960

Type of genus. Foliophycus llanoensis Johnson 1960. Middle Carboniferous,
Texas. Type extensively recrystallized.
Diagnosis. Thallus long with slender leafy fronds. They are composed of long
threads of rectangular or barrel-shaped cells emerging from central axis and
branching dichotomously. No perithallic-hypothallic differentiation. Needs
revision.
Suprageneric attribution. Undetermined family among the rhodophytes. The
closest known taxon is Masloviporidium Groves and Mamet 1985.
Stratigraphic range and distribution. Scarce. Namurian-Moscovian. Algeria,
Canadian Arctic, American midcontinent and Texas.
52. Fourstonella Cummings 1955

Type of genus. Stacheia fusiformis Brady 1876. Visean, Great Britain.
Diagnosis. Thallus fusiform, symmetrical, encrusting around a central support.
Thin rows of minute, subparallel laminae, slightly overlapping. Insertion of
rows oblique on the support. Cells quadratic, regular. Wall calcareous, yel-
lowish, hyaline (see Plate 3g).
Fourstonella irregularis Mamet and Roux 1977. Late Visean. Algeria, Morocco,
France, Siberia.
Suprageneric attribution. Originally considered a foraminifer, it has been
transferred to the ungdarellids by Mamet and Rudloff (1972), a transfer based
on the nature of the wall. Stacheinaceae Loeblich and Tappan 1961. New tribe
Mametellae Chuvashov 1987.
Stratigraphic range and distribution. Visean-Namurian. Reaches the Mos-
covian in the Canadian Arctic. Algeria, Morocco, Great Britain, France, Ger-
many, Poland, Russian Platform, Siberia (Verkhoyansk), Canadian Arctic,
American Midcontinent, Nova Scotia, Newfoundland.
53. Frustulata Saltovskaja 1984

Type of genus. Frustulata asiatica Saltovskaja 1984. Visean, Tadzhikhistan.
Diagnosis. Thallus cylindrical, regularly branching. Medulla divided by ir-
regular pseudosepta. Cortex perforated by thin, simple pores. Further in for-
402 B. Marne!
mation is needed to differentiate it from Kulikaella Berchenko 1981.

Suprageneric attribution. Systematic position debatable (see Shuysky 1987).
54. Garwoodella Paul 1938

Type of genus. Garwoodella thallus Paul 1938. Visean, Germany.
Diagnosis. Thallus nodular composed of closely-packed layers of micritized
tubes. Two different types of tube: a) regular, with simple, low-angle dicho-
tomy; b) flexuous, irregular, pinched, with erratic dichotomy. Needs revision.
Could be an association of Ortonella and Mitcheldeania. Needs revision.
Suprageneric attribution. Originally Schizophyceae. Probably a nodular
codiacean.
Stratigraphie range and distribution. Visean. Germany, Belgium, American
midcontinent.
55. Garwoodia Wood 1941

Type of genus. Mitcheldeania gregaria Nicholson 1888 emend. Wood 1941.
Tournaisian, Great Britain.
Diagnosis. Thallus nodular, composed of radiating, dichotomous, micritized
tubes. These tubes, of even diameter, bifurcate at right angles.
Garwoodia zonata (Derville 1931). Visean, France.
Suprageneric attributions. Nodular codiacean for Mamet and Roux (l975b)
Hormogonophyceaea (Geitler) Elenkin 1934 for Luchinina (1987).
Stratigraphic range and distribution. Common in lagoons and semi-restricted
waters. Ordovician to Carboniferous. Scarce in Permian. Great Britain, Bel-
gium, France, Carnic Alps, Turkey, Podolia, Urals, Siberia, Japan, Canadian
Arctic, American Cordillera and midcontinent.
56. Girvanella Nicholson and Etheridge 1878

Type of genus. Girvanella problematica Nicholson and Etheridge 1878 emend.
Wood 1957. Ordovician, Great Britain, see Plate 3g.
Diagnosis. Micritized sheaths of random filaments. Filaments do not branch
and have even diameter. They are continuous and without partitions.
Synonyms: Nicholsonia Korde 1973, Batinevia Korde 1973.
Danielli (1981) cites many Paleozoic Girvanella which should be transferred to
Ellesmerella, Sphaerocodium, Mitcheldeania, Obruchevella, Razumovskia,
Malakhovella, etc.
Mamet and Roux (1975b) recognize four species based on the diameter of the
filament. The first three species are cosmopolitan and very long ranging:
Girvanella staminea Garwood 1931.
Girvanella werheredii Chapman 1908 (Girvanella incruslans Wethered 1890. non
Borneman 1886).
Girvanella problematica Nicholson and Etheridge 1878 emend. Wood 1957

(= Girvanella ducii Wethered 1890).
Girvanella kasakiensis Maslov 1949 is known from France, Great Britain,
Poland, Russian Platform, Urals, Kazakhstan, China, Australia.
Suprageneric attributions. Controversial, but consensus is that girvanellids are
calcified cyanobacteria. Otherwise attributed to Chlorophycophyta, Poros-
tromata, Oscillatoriales, Nostocales, Hormogoneae, etc. Type of the family
Girvanellacea Luchinina 1975.
Stratigraphic range and distribution. Very common, not only in restricted
lagoons, but through the whole range ofthe photic zone in carbonate shelves or
ramps. Many mudstones-wackestones are formed by intertwined girvanellid
tubes. Forms "Girvanella community" of FlUgel (1977). Proterozoic to Cre-
taceous. Eurasia including Siberia, North Africa, Australia, Canadian Arctic,
North American Cordillera and midcontinent, Maritime Provinces of Canada.
57. Gissarella Saltovskaja 1979

Type of genus. Gissarella elegantula Saltovskaja 1979. Middle Carboniferous,
Tadzhikhistan.
Diagnosis. Needs revision. Hypothetical reconstruction of Saltovskaja in-
dicates a euspondyl thallus with cylindrical axial stem and unramified branches.
Suprageneric attribution. Doubtfully reported to the Luliporeae Shuysky 1986
among the Acetabulariaceae.
Globuliferoporella Chuvashov 1974

See Epimastopora.
Goksuella GUven<s 1965

See Donezella.
58. Gouldina Johnson 1940

Type of genus. Gouldina carbonaria Johnson 1940. Middle Carboniferous,
Colorado.
DiagnOSiS. Needs revision. A mammillated algal biscuit composed of un-
dulating laminae (see also Shermanophycus).
Suprageneric attribution. Spongiostromidae.
Stratigraphic age and distribution. See type of genus.
Grozdilovella Chermnykh 1972

See Protoumbella.
59. Gyroporella (GUmbel 1872) Benecke 1876

Type of genus. Gyroporella vesiculifera GUmbel 1872. Trias, Alps.
404 B. Marne!
Diagnosis. Thallus aspondyl, club-shaped. Medulla of similar shape. Cortex

perforated by important phloiophore? branches.
? Gyroporella constricta Kochansky-Devide 1970. Moscovian, Yugoslavia.
Gyroporella intraseptata Kochansky-Devide 1970. Moscovian, Yugoslavia.
? Gyroporella likana Kochansky-Devide 1970. Moscovian, Yugoslavia.
Gyroporella nipponica Endo and Hashimoto 1955. Late Carboniferous - Late
Permian. Spain, Yugoslavia, Urals, Kazakhstan, China, Japan, British
Columbia, Canadian Arctic (see Plate 2q).
? Gyroporella primitiva Ra user-Chernoussova and Korolyuk 1981. Moscovian,
Urals.
? Gyroporella prisca Kochansky-Devide 1970. Moscovian, Yugoslavia.
? Gyroporella shimanensis Chuvashov and Anfimov 1988. Moscovian, Urals.
Suprageneric attributions. Tribe Gyroporelleae (Pal 1976) Bassoullet et al. 1979
for Deloffre (1988). Subtribe Gyroporellinae Berchenko 1987 for Shuysky
(1987).
Stratigraphic range and distribution. Moscovian (scarce) to Jurassic. Most
abundant in Permian-Triassic where it is cosmopolitan in the Northern
Hemisphere.
60. Halenopora Hance 1983

Type ofgenus. Hale nopora radia ta Hance 1983. Tournaisian -Visean boundary,
Belgium.
Diagnosis. Thallus(?) cylindrical, composed of three to five poorly calcified
central canals of even diameter from which emerge thin, oblique lateral canals
which become progressively thicker and more perpendicular to the thallus.
Suprageneric attribution. The idealized reconstruction of Hance shows two
series of fertile and non-fertile branches derived from the stem and suggests an
euspondyl dasycladacean. Illustrations show a udoteacean, similar to
Dimorphosiphonoides.
Stratigraphic range and distribution. See type of the genus.
61. Hedstroemia Rothpletz 1913

Type of genus. Hedstroemia halimedoidea Rothpletz 1913. Silurian, Gotland.
Diagnosis. Nodule composed of radially disposed cylindrical tubes which have
multiple (triple to quadruple and higher) dichotomies.
Hedstroemia corymbosa Pia 1937. Visean, France, Great Britain.
Hedstroemia koninckoporoides Vachard 1988 is not an alga.
Suprageneric attributions. Nodular codiacean for Mamet and Roux (l975b).
Porostromata for Poncet (1986), Garwoodiaceae for Luchinina (1987).
Stratigraphic range and distribution. A widespread Ordovician-Silurian-
Devonian taxon that peters out in the Lower Carboniferous. Highest recorded
occurrence is Namurian. Eurasia. Australia. North America.
62. Herakella Kochansky-Devide 1970

Type of genus. Herakella paradoxa Kochansky-Devide 1970. Moscovian,
Yugoslavia.
Diagnosis. Thallus cylindrical, segmented, asymmetrically annulated, meta-
spondyl. Medulla cylindrical. Vestibule followed by tufts of at least ten
phloiophore branches, with first to second order dichotomy.
Suprageneric attributions. Diploporeae? (Pia 1920) Giiven<; 1979 according to
Deloffre (1988). Linoporalinae (Pia 1927) Bassoullet et al. 1979 for Shuysky
(1987).
Stratigraphie range and distribution. Middle to Late Carboniferous. Yugos-
lavia, Central Asia.
Hikorocodium Endo 1951

Type ofgenus. Hikorocodium elegantula Endo 1951. Permian, Japan. Although
reported twice by Endo from the Carboniferous of Japan, this incertae sedis is
restricted to the Permo-Triassic (Endo 1951, 1957).
63. Iberiaella Racz 1984

Type of genus. Iberiaella carbonica Racz 1984, Middle Carboniferous, Spain.
Diagnosis. Thallus encrusting or crustose, composed of colony-forming,
thread-like wavy thin tubes with constrictions. They branch randomly, forming
an anastomosing pattern. Wall calcareous, yellowish, hyaline.
Suprageneric attribution. Undetermined algal family for Racz (1984). Seems
akin to Berestovia Berchenko 1983.
Stratigraphie range and distribution. See type of genus.
64. Intextulella Petryk in Petryk and Mamet 1972

Type of genus. Intextulella agglomerata Petryk in Petryk and Mamet 1972.
Visean, Alberta.
Description. Thallus encrusting, composed of regular strands of crustose,
layered, rectangular cells weaving uncalcified intercellular tissue. Wall yel-
lowish, clear. hyaline. Needs revision.
Suprageneric attribution. Undetermined family among the rhodophytes.
65. Issinella (Reitlinger 1954) Mamet and Roux 1981a

Type of genus. Issinella devonica Reitlinger 1954 as a Devonian siphonales,
Russian Platform (see Plate 19).
Description. Thallus cylindrical, aspondyl, erect. Medulla important, divided
by septiform processes. Cortex fibro-radial, perforated by thin, randomly
disposed, unramified branches. Conceptacles subcortical in regular rows.
Synonym. Luteotubulus Vachard in Termier et al. (1977).
406 B. Mamet
Issinella grandis Chuvashov 1965. Upper Famennian - Upper Tournaisian.

France, Urals, Donbass, Siberia (Kolyma).
Issinella ilychensis Ivanova 1988. Visean, Urals.
Issinella sainsii Mamet and Roux 1975. Upper Famennian-Visean. France,
Belgium, Siberia (Kolyma, Omolon), Alaska, Utah.
Suprageneric attributions. A green alga for Reitlinger (1954) and an
"algosponge" for Termier, Termier and Vachard 1977. Placed among the
Issinelleae Deloffre 1987 by Deloffre (1988). Erroneously considered a Da-
syporelleae Pia 1920 by Shuysky (1987).
Stratigraphic range and distribution. Middle Devonian-Late Visean. Very
common. Forms bafftestones associated with other tubular algae, palaeobere-
sellids and calcispheres. Morocco, Spain, Belgium, France, Ireland, Turkey,
Russian Platform, Urals, Donbass, Siberia (Kuzbass, Kolyma, Omolon), China,
Australia, Alaska, Bri tish Col urn bia, Al berta, Utah, M on tana , Tennessee, Nova
Scotia.
66. Ivanovia Khvorova 1946

Type of genus. Ivanovia tenuissima Khvorova 1946. Late Carboniferous, Rus-
sian Platform (see Plates Ii and 3f).
Diagnosis. Thallus composed of large blades, standing erect on substratum.
Central zone with dichotomous poorly calcified filaments(?), usually obliterated
by cementation. Cortex contains rows of cylindrical utricles(?) with branches
perpendicular to the exterior.
Suprageneric attribution. "Phylloid algae" (vernacular). New tribe Ivanoviae
Shuysky 1987 among the Anchicodiaceae Shuysky (1987).
Stratigraphic range and distribution. First occurrence in Middle Carboniferous.
Abundant from Moscovian to Early Permian. Often associated with Eugono-
phyllum and Neoanchicodium in phylloid mounds characterized by Tubiphytes.
Algeria, Spain, Carnic Alps, Russian Platform, Urals, China, Canadian Arctic,
Yukon, Colorado, New Mexico, Texas.
Izhella Antropov 1955

See Renalcis.
67. laphetella Mamet and Roux 1983

Type ofgenus. laphetella boehmi Mamet and Roux 1983. Late Visean, France
(see Plate I d).
DiagnOSiS. Thallus cylindrical, annulated. Verticils separated by slight con-
strictions. Medulla cylindrical with poorly defined expansions forming ves-
tibule. Tufts of phloiophore branches, slightly arched.
Suprageneric attribution. Diploporeae (Pia 1920) Guven<; 1979 according to
Deloffre (1988).
Stratigraphic range and distribution. Late Visean Zone 16 j and 16 s ' France,
Spain.
68. Kamaena Antropov 1967

Type ofgenus. Kamaena delicata Antropov 1967 (= Kamaena simplex Eickhoff
1968). Devonian-Carboniferous transition, Russian Platform.
Diagnosis. Thallus long, slender, cylindrical, with dichotomic ramifications.
Medulla divided by regular pseudosepta, of even thickness and perpendicular
to the cortex. Inner cells rectangular to subrectangular. Cortex perforated by
thin, straight, undivided, randomly disposed branches.
Synonyms. Carboniferous "Nodosinella" of the literature, Devonoscalae
Langer 1979, Subkamaena Berchenko 1981.
Kamaena awirsi Mamet and Roux 1974. Devonian-Carboniferous transition to
Visean. Ireland, Great Britain, Belgium, Donbass, Siberian Platform, Kolyma.
Kamaena concaviuscula (Berchenko 1981). Devonian-Carboniferous transi-
tion, Donbass.
Kamaena itkillikensis Mamet in Petryk and Mamet (1972). Lower Carbonifer-
ous, Great Britain, Belgium, Donbass, Urals, Siberia, Alaska, Alberta.
Kameana lata Ivanova 1988. Visean, Urals, Siberian Platform (Kolyma,
Omolon).
Kameana maclareni Mamet and Rudloff 1972. Lower Carboniferous, Alberta,
British Columbia, Alaska.
Kamaena magna Ivanova 1988. Famennian-Visean. Morocco, Ireland, France,
Urals
Kameana pirleti Mamet and Roux 1974. Lower Carboniferous, Great Britain,
France, Belgium, Donbass, Urals.
Kameana razdolnica (Berchenko 1981). Devonian-Carboniferous transition,
Donbass, Urals
Kameana sibirica Ivanova 1988. Tournaisian, Siberian Platform.
Suprageneric attributions. Palaeobereselleae Mamet and Roux 1974. Tribe
Kamaeneae Shuysky 1985 among the Palaeosiphonocladales Shuysky 1985.
Stratigraphic range and distribution. Abundant in Carboniferous lagoons as-
sociated with Palaeoberesella. Middle Devonian - Lower Carboniferous.
Morocco, Algeria, Spain, Ireland, Great Britain, France, Belgium, Germany,
Poland, Russian Platform, Donbass, Urals, Central Asia, Afghanistan, Siberia,
(Taimyr, Kolyma, Omolon), Australia, Alaska, Alberta, American Cordillera,
Iowa, Newfoundland.
69. Kamaenella Mamet and Roux 1974

Type of genus. Kamaenella denbighi Mamet and Roux 1974. Late Visean,
Great Britain (see Plate 1h).
Diagnosis. Thallus cylindrical, with extensive twisted bifurcations. Medulla
divided by regular, even pseudoseptations, perpendicular to the cortex.
Branches straight, very thin. Medullar cavities have height greater than width.
Kamaenella tenuis. (von Moller 1879). Visean, Morocco, Spain, France, Great
Britain, Russian Platform (see Plate 3s).
408 B. Marne!
Suprageneric attributions. Palaeobereselleae Mamet and Roux 1974. Tribe

Kamaeneae Shuysky 1985 (as a Palaeosiphonocladales Shuysky 1985).
Stratigraphic range and distribution. Lower Carboniferous. Morocco, Spain,
Great Britain, France, Russian Platform, Donbass, Urals, Tadhzikhistan,
Newfoundland.
70. Komia Korde 1951

Type ofgenus. Komia abundans Korde 1951, Middle Carboniferous, Urals. (=
Komia eganensis Wilson and Waines 1963) (see Plate 3q).
Diagnosis. Thallus ramose, cylindrical, robust. Hypothallic tissue thin, a flaring
bundle of regular, elongated, rectangular cells. Perithallic tissue much more
conspicuous, composed of quadrangular cells perpendicular to the central axis.
Wall yellowish, hyaline. Conceptacles in cortical cells. Close to Ungdarella and
hard to differentiate unless the section shows the hypothallic tissue.
Suprageneric attributions. Controversial with unfounded attributions to
echinoderms, stromatoporoids, hydrozoans and calcisponges. Korde's original
description is correct and with its companion Ungdarella, the genus belongs to
the Ungdarellaceae Maslov 1962.
Stratigraphic range and distribution. Very abundant. Forms the "Komia com-
munity" of Fltigel (1977) (shelf margin community of Wilson 1975). First
probable occurrence in Namurian (Zone 18 of the Nizi Formation, British
Columbia). Last occurrence, Early Permian. Cosmopolitan in the Northern
Hemisphere. Morocco, Spain, Great Britain, Urals, Donbass, Kazakhstan, Iran,
Japan, Alaska, Canadian Arctic, American Cordillera and midcontinent.
7l. Koninckopora Lee 1912

Type of genus. Calamopora infiata de Koninck 1842. Visean, Belgium.
Diagnosis. Thallus large, arched to cylindrical with an conspicuous medulla.
Cortex, by comparison, thin, coarsely perforated by subpolygonal straight
branches. Wall two-layered: an inner micritic layer and a yellowish pseudo-
fibrous layer.
Synonym: Coeloceratoides Derville 1931, Uragiella Maslov 1935, Koninck-
oporoides Rich 1974.
Koninckopora minuta Weyer 1968 (= Koninckopora sahariensis Chanton 1964
pars).
Koninckopora mortelmansi Mamet 1973 (= Koninckoporoides monteaglensis
Rich 1974).
Koninckopora pruvosti Gi.iven~ 1966 (= Koninckopora sahariensis Chanton
1964, pars).
Koninckopora tenuiramosa Wood 1942 (see Plate 3u).
All these species are Visean and, except for pruvosti, are cosmopolitan in the
Northern Hemisphere and Australia. Koninckopora pruvosti is known from
Morocco, Spain, France, Belgium, Germany. Turkey. Iran and Australia.
Suprageneric attributions. Controversial with attributions to the tabulates,
bryozoans, cephalopods. Belongs to the cyclocrinitid Seletonellaceae either as

Mastoporeae (Pia 1920) (Deloffre 1988) or as Cyclocrineae Pia 1929 (Shuysky
1987).
Stratigraphic range and distribution. Very common in high-energy shoals and
banks. Fragile and usually observed as fragments. Cosmopolitan from Zone II
to 16 (Early to Late Visean) in Eurasia, including Siberian Arctic (Taymyr,
Omolon), North Africa, Australia, Canadian Arctic and elsewhere in North
America. Extends up to Zone 18 in Southern China.
Koninckoporoides Rich 1974

See Koninckopora.
72. Kulikia Golubtsov 1961

Type of genus. Kulikia sphaerica Golubtsov 1961. Upper Visean, Byelorussia.
Diagnosis. Thallus metaspondyl, a succession of spherical units. Central
medulla conspicuous, cylindrical, continuous but with constrictions. Verticils
with first order ramifications. Branches(?) nearly spherical, communicating by
round pores. Needs to be partially revised. Original description is confusing and
the emendation of Skompski (1984) combines two taxa. The best published
photograph of the genus is in Shuysky (1987, PI. XIII, Fig. 5). Similar to, but
distinct from, the aciculellid Sphinctoporella.
Suprageneric attribution. Aciculelleae Bassoullet et al. 1979 emend. Deloffre
1988.
Stratigraphic range and distribution. Scarce. Late Visean. Belgium, Poland,
Russian Platform, Ukraine.
73. Litostroma Mamay 1959

Type of gen us. Litostroma oklahomense Mamay 1959, Middle Carboniferous,
Oklahoma.
Diagnosis. Thallus, encrusting, flat platelets, one cell thick. Cells rather ir-
regular, subpolygonal in axial section.
Suprageneric attribution. Undetermined family among the rhodophytes?
Stratigraphic range and distribution. Very scarce. Lower and Middle Carbon-
iferous. Oklahoma, Newfoundland.
Luteotubulus Vachard in Vachard et al. (1977)

See Issinella.
74. Macroporella Pia 1912

Type ofgenus. Macroporella dinarica Pia 1912. Trias, Yugoslavia.
Diagnosis. Thallus cylindrical, unsegmented, with conspicuous medulla.
Phloiophore primary branches in irregular oblique whorls.
410 B. Marne!
Macroporella ginkeli Racz 1965. Bashkirian-Moscovian, Spain, Urals.

Macroporella ortashensis Rauser-Chernoussova and Korolyuk 1981. Mos-
covian, Urals.
Suprageneric attributions. Macroporellinae (Pia 1920) emend. Bassoullet et a1.
1979 trans. Berchenko and Saltovskaja 1987.
Stratigraphic range and distribution. Bashkirian to Jurassic. Abundant from
Permian to Triassic. Eurasia. North America.
"Malacostroma" Giirich 1906

Type of genus. Malacostroma concentricum Giirich 1906, Visean, Belgium,
France.
Diagnosis. Regularly layered algal biscuit, composed of loose peloidal mesh-
work of Bevocastria filaments and darker, heavily micritized layers with Gir-
vanella. The two other species, "Malacostroma" plumosum Giirich 1906 and
"M". undulosum Giirich 1906 have similar, but more irregular layering.
Suprageneric attributions. Spongiostromidae.
Stratigraphic range and distribution. Visean of Belgium. Questionable in the
Visean of the American midcontinent.
75. Malakhovella Mamet and Roux 1977

Type ofgenus. Malakhovella malakhovae Mametand Roux 1977, Visean, Urals.
Diagnosis. Bundles of intertwined micritized sheaths of filaments forming
pseudo-segments.
Malakhovella sparsa Saltovskaja 1984. Visean. Tadzhikhistan.
Suprageneric attributions. Incertae sedis. Cyanophyte? Girvanellaceae Lu-
chinina 1975 according to Luchinina (1987).
Stratigraphic range and distribution. Lower Carboniferous, Belgium, Urals,
Tadzhikhistan.
76. Mametella Brenckle 1977

Type of genus. Mametella chautauquae Brenckle 1977. Visean, midcontinent
(= Stacheia? skimoensis Mamet and Rudloff 1972).
Diagnosis. Thallus encrusting. fusiform or lemon-shaped. Cells quadratic. in
regular concentric rows. Wall. a yellowish hyaline calcite.
Suprageneric attribution. New tribe Mametellae Chuvashov 1987. A
Stacheiinae.
Stratigraphic range and distribution. Quite common in North America (Alaska.
Cordillera. midcontinent) and scarce in Europe (Ireland. Great Britain and
Siberia (Kuzbass. Verkoyansk). Tournaisian - Lower Namurian.
77. Masfoviporclla Kulik 1973

Tvpe olgenus. Masfoviporella cafixoidca Kulik 1973. Serpukhovian. Urals.
Diagnosis. Thallus cylindrical. a succession of vertic ilia ted. protruding. regular
crowns. Branches euspondyl. simple. Insertion on stem is unclear.

Masloviporella conili Vachard III Perret and Vachard 1977. Serpukhovian,
France.
Suprageneric attribution. Clypeineae (Elliott 1968) Bassoullet et al. 1979, ac-
cording to Deloffre (1988) and Shuysky (1987).
Stratigraphic range and distribution. Serpukhovian. France, Urals.
78. Masloviporidium Groves and Mamet 1985
Type of genus. Donezella delicata Berchenko 1982. Serpukovian-Bashkirian

boundary, Donbass (see Plate 3i).
Diagnosis. Thallus sheet-like, foliate, branching, composed of arcuate rows of
wedge-shaped cells. Horizontal partitions heavily calcified. Vertical partitions
thinner, irregular. Pores connect continuous cells of adjacent rows. Wall cal-
careous, hyaline (see Plate 2d).
Suprageneric attribution. Incertae familia among the rhodophytes.
Stratigraphic range and distribution. The only known Carboniferous red alga
which has precise stratigraphic connotations. Morrowan-Atokan (Zones 20-21)
in North America, uppermost Serpukhovian-basal Bashkirian (Zones 20-21) in
Eurasia. Algeria, Donetz, Central Asia, China, Alaska, Idaho, Utah, Arkansas,
Oklahoma, Kansas, Texas.
79. Mellporella Racz 1965
Type ofgenus. Mellporella anthracoporellaformis Racz 1965. Moscovian, Spain.

Diagnosis. Thallus cylindrical, unbranched. Medulla very important. Cortex
thin, perforated by numerous random, short, phloiophore pores which taper out
before bifurcation.
Suprageneric attributions. Dasyporelleae (Pia 1920) Bassoullet et al. 1979,
according to Deloffre (1988).
Stratigraphic range and distribution. Scarce. Late Bashkirian-Moscovian,
Spain, Algeria.
80. Menselina Antropov 1967

Type of genus. Menselina clathrata Antropov 1967. Devonian-Carboniferous
transition, Russian Platform.
Diagnosis. Thallus(?) conical. Wavy horizontal calcified partitions which could
correspond to fronds? Vertical partitions poorly developed. No central stem.
Menselina rotunda Berchenko 1981. Devonian-Carboniferous transition,
Donbass, Siberia (Altai, Omolon).
Menselina triangula Berchenko 1981. Devonian-Carboniferous transition,
Donbass.
Suprageneric attributions. Microprob1ematic. "Problematic clear cones" of
Brazhnikova and Rostovceva (1966). "Problematic algae" of Bogush et al. (1970).
Stratigraphic range and distribution. Devonian-Carboniferous transition.
412 B. Marnet
Belgium, France, Russian Platform, Donbass, Urals, Siberia (Altai, Omolon),

Afghanistan.
81. Microcodium GlUck 1912

Type of genus. Microcodium elegans GlUck 1912. Miocene, Germany.
Diagnosis. A cob-shaped thallus(?) with a central micritized cavity surrounded
by peripheral radial plates. Each plate contains a central, thin, cylindrical tube.
The "corn-grain" plates often corrode the surrounding carbonates (see Plate 1e).
Suprageneric attributions. Highly controversial. Originally placed among the
codiaceans. Maslov erected the family Microcodiaceae (1960). See Klappa
(1978, table 1) for attributions ranging from fungi to vascular plant rootlets.
Often considered as indicative of pedogenesis. Erroneously considered as
Cenozoic infiltrations (Bodergat 1975). Refer to Mamet and Roux (1982) for
discussion of Palaeozoic occurrences. See also the thesis of Plaziat (1984).
Stratigraphic range and distribution. Oldest documented occurrence in the
Emma Fjord Formation of the Canadian Arctic (Visean). Very common in
Middle-Carboniferous and Permian shallowing-up sequences of the Canadian
Arctic. Cosmopolitan in the Mesozoic and Cenozoic of the Northern
Hemisphere.
82. Mitcheldeania (Wethered 1886) Mamet and Roux 1975b.

Type of genus. Mitcheldeania nicholsoni Wethered 1886 pars. Lower Carbo-
niferous, Great Britain.
Diagnosis. Thallus nodular, an aggregate of micritized, highly dichotomous
sheaths. Tubes are flexuous and pinched at point of bifurcation.
Mitcheldeania distans (Conil and Lys 1964). Visean-Serpukhovian, Belgium,
France, Czechoslovakia, Nova Scotia.
Suprageneric attribution. Nodular codiacean for Mamet and Roux (l975b).
Garwoodiaceae for Luchinina (1987).
Stratigraphic range and distribution. Common in Dinantian-Namurian
lagoons, often associated with Garwoodia and Ortonella. Cosmopolitan, Eur-
asia, North Africa, North America.
83. Nanopora Wood 1964

Type of genus. Nanopora anglica Wood 1964. Visean, Great Britain.
Diagnosis. Thallus minute, subcylindrical. Branches simple, euspondyl, per-
pendicular to the cylindrical. continuous, unconstricted medulla. In equatorial
sections, branches are elliptical, oval and alternate.
Nanopora [ragilissima (Maslov 1939). Lower Carboniferous, Kazakhstan,
Taimyr, Kolyma, Omolon (see Plate 3c).
Suprageneric attributions. Tribe Uraloporellae Shuysky 1985 for Shuysky
(1987). Salpingoporellinae Bassoullet et al. 1979. according to Deloffre (1988).
Stratigraphic range and distribution. Visean-Early Namurian. Cosmopolitan,

Algeria, Morocco, Belgium, Great Britain, France, Germany, Donbass, Poland,
Kazakhstan, Taimyr, Kolyma, Omolon, Alaska, Nova Scotia, Newfoundland.
84. Nansenella Mamet and Roux 1987

Type of genus. Nansenella multifurcata Mamet and Roux 1987. Bashkirian,
Ellesmere Island (see Plate If).
Diagnosis. Thallus nodular, composed of micritized sheaths. Tubes very ir-
regular, sinuous, dichotomous in all direction, sometimes cylindrical and
continuous, then constricted ampuliform. Simple, double or triple
ramifications.
Suprageneric attribution. Nodular codiacean (Mamet and Roux 1987).
Cyanobacteria?
Stratigraphic range and distribution. Bashkirian to Sakmarian. Canadian Arctic.
85. Neoanchicodium Endo in Endo and Kanumai 1954

Type of genus. Neoanchicodium catenoides Endo in Endo and Kanumai 1954.
Lower Permian, Japan (see Plate Ij).
Diagnosis. Thallus composed oflarge blades, erect on substratum. Blades end
in cylindrical "fingers". Central zone with dichotomous poorly calcified tubes,
contains one row of ellipsoidal "utricles"(?) with internal and external openings.
The reconstruction ofShuysky (1987), inspired by Chuvashov (1974), indicates
two rows of "utricles", which are not observed in the type material.
Suprageneric attributions. "Phylloid algae" (vernacular). New family An-
chicodiaceae Shuysky 1987.
Stratigraphic range and distribution. Forms the "Neoanchicodium community"
of Fliigel (1977). Not as widespread as Eugonophyllum or Ivanovia, but im-
portant in formation of phylloid banks. Moscovian - Lower Permian.
Yugoslavia, Carnic Alps, Urals, China, Japan, Canadian Arctic.
Nigriporella Rigby 1958

See Tubiphytes.
86. Nostocites Maslov 1929 sensu Maslov 1956

Type of genus. Nostocites vesiculosa Maslov 1929, Middle Carboniferous,
Donbass.
DiagnOSis. Thallus composed of cells serving as substratum to a palisade of
elongate cylindrical cells containing an axial micritized filament.
Note. The original drawings of Maslov (1929) are sketchy and have led to
numerous misinterpretations. The type-material is preserved in Moscow and
Maslov's (1956) reinterpretation is correct.
Suprageneric attributions. Hormogonales for Maslov in Orlov (1963). Pleu-
rocarpales? (Chroococcidiaceans) for Mamet and Roux (1983).
Stratigraphic range and distribution. Occurs as Early as the Middle Devonian
414 B. Marne!
(Eifelian of Belgium). Quite common throughout the Carboniferous. Spain,

Poland, Great Britain, Belgium, Donbass, Russian Platform, China, Australia,
Canadian Arctic, Alaska, New Mexico, Arkansas, Oklahoma, Newfoundland.
87. Orthriosiphon Johnson and Konishi 1956

Type of genus. Orthriosiphon saskatchewanensis Johnson and Konishi 1956.
Tournaisian-Visean boundary, Saskatchewan.
Diagnosis. Thallus very large and thick. Cortex club-shaped, poorly calcified,
fragile, often abraded. Cortical tubes dichotomous, diverging slowly a number
of times and terminating is small funnels at the outer periphery. Conceptacles
prominent, subspherical, irregularly arranged, subcortical.
Note. The taxon shows affinities with Cummingsella and Orthriosiphonoides.
Othriosiphon turnhouti Mamet, Mortelmans and Roux 1979. Visean, Belgium.
Suprageneric attributions. At the creation of the genus, thought to belong to the
Codiaceae? Attributed to the Triploporelleae (Pia 1920) Bassoullet et al. 1979
according to Shuysky (1987) (cortical tubes interpreted as dasycladacean
"branches"). Udoteaceae for Bassoullet et al. (1983).
Stratigraphic range and distribution. Late Tournaisian-Late Visean. In the
Canadian Arctic extends up to the Bashkirian. Belgium, Alaska, Canadian
Arctic. Very prolific in the American Cordillera (Alberta, Wyoming, Idaho,
Montana), Saskatchewan, Tennessee, Alabama. The type Orthriosiphon sas-
katchewansis is endemic to North America.
88. Orthriosiphonoides Petryk in Petryk and Mamet 1972, non Vachard 1981
Type ofgenus. Orthriosiphonoides salterensis Petryk in Petryk and Mamet 1972.
Visean, Alberta (see Plate 21).
Diagnosis. Thallus large, club-shaped, segmented, "annulated". Large central
stem. Segments outwardly rounded. Sutures deep. Cortical tubes thin, nume-
rous, straight to slightly curved, of constant diameter, dichotomous. Concep-
tacles spherical to ellipsoidal, randomly distributed.
Orthriosiphonoides tenuiramosa Mamet and Rudloff 1972. Visean, Alberta,
British Columbia (see Plate 3v).
Suprageneric attributions. Udoteaceae for Bassoullet et al. 1983 (see also cri-
tique of Va chard 1981). Attributed to the Triploporelleae (Pia 1920) Bassoullet
et al. 1979 by Shuysky (1987) (cortical tubes interpreted as dasycladacean
"branches").
Stratigraphic range and distribution. Endemic to North America (Eurasiatic
reports of Va chard (1981) may be due to erroneous taxonomy). Visean to Early
Permian. Can be very common in high energy channelized grainstones. Can-
adian Arctic, American Cordillera (Alaska, Yukon Territory, British Columbia,
Alberta, Idaho, Wyoming, Montana, Utah), Tennessee.
89. Ortonella (Garwood 1914) Mamet and Roux 1981

Type ofgenus. Ortonellafurcata Garwood 1914. Visean, Great Britain.
Diagnosis. Thallus nodular, composed of closely-packet radiating, dicho-
tomous micritized sheaths. Even-diameter tubes branch at relatively low angle
(100 to 45°).
Ortonella balbinia (Poncet 1974). Siegenian-Visean, France, Morocco.
Orton ella capnostyloides (Derville 1931). Visean, France
Ortonella coloradoensis Johnson 1945. Lower to basal Middle Carboniferous,
Urals, Alaska, Alberta, Saskatchewan, Colorado.
Ortonella gracilis Johnson 1951. Late Carboniferous-Permian, Canadian
Arctic, Texas. (Probable in Germany and Russian Platform as aff.)
Ortonella kershopensis Garwood 1931. Lower Carboniferous, Ireland, Great
Britain, France, Donbass, Kuznetsk, Alberta, Colorado.
Ortonella maksimovae Maslov 1956. Tournaisian, Donbass.
Ortonella moscovica Maslov 1956. Lower Carboniferous, Belgium, Russian
Platform.
Ortonella myrae Racz 1964. Moscovian, Spain.
Ortonella tenuissima Garwood 1931. Famennian - Lower Carboniferous,
Great Britain, Donbass, Australia.
Orton ella tyrrelensis Mamet and Rudloff 1972. Tournaisian-Visean, Belgium,
Alberta, Nova Scotia.
Ortonella upensis Birina 1948. Devonian-Carboniferous transition to Tour-
naisian, Russian Platform, Urals, Kazakhstan.
Suprageneric attributions. Nodular codiacean (Mamet and Roux 1981). Gar-
woodiaceae (Johnson) Shuysky 1973 according to Luchinina (1987).
Stratigraphic range and distribution. Common in lagoons (see Garwoodia).
Cosmopolitan in Northern Hemisphere. Very long ranging from Ordovician to
Mesozoic.
"Osagia" Twenhofel1919
Type of the ''form-genus''. Osagia incrustata Twenhofel1919. Lower Permian,
Kansas.
Diagnosis. Oncoid formed by the intimate intergrowth of algal filaments, sessile
foraminifers and other organisms such as sponges or bryozoans. Carboniferous
"Osagia" often produced by Girvanella sheaths, encrusting apterrinellids and
sponges. The association of Girvanella, Archaeolithophyllum, phylloid blades
and attached tuberitinids is common in the Moscovian. In the Bashkirian,
"Osagia" is formed by Berestovia-cyanophytes, by Iberiaella-Girvanella-
Donezella or by Bevocastria-sponges. As "Osagia" is formed by associations of
different organisms, the generic and specific concepts have little taxonomic
value.
Suprageneric attributions. See diagnosis.
Stratigraphie range and distribution. The" Osagia" community of Fliigel (1977)
is composed of Girvanelle-Hedraites-apterrinellid oncoids formed in shallow
416 B. Mamet
near-coast or in off-shore environments. Doubtful reports in the Riphean

(where it was probably formed by cyanobacteria). "Osagia" is scarce in the
Lower Carboniferous and has a sudden outburst in the Bashkirian. Common
from the Middle Carboniferous to the Permian. Cosmopolitan in the Northern
Hemisphere.
"Ottonosia" Twenhofel1919
Type of «form-genus". Ottonosia laminata Twenhofel 1919. Lower Permian,
Kansas.
Diagnosis. Asymmetrical encrustations on the upper surface of a substratum.
Microftora and microfauna are the same as observed in "Osagia". Needs
revision.
Suprageneric attributions and stratigraphic range. See "Osagia".
Pachysphaera Conil and Lys 1964 OB]

see Calcisphaera.
Pachysphaerina Conil and Lys 1967 OB]

see Calcisphaera.
90. Palaeoberesella Mamet and Roux 1974

Type ofgenus. Nodosinella lahusenivon Moller 1879. Visean, Russian Platform.
Diagnosis. Thallus cylindrical, long, with rare ramifications. Medulla sub-
divided by coarse conical projections forming irregular medullary cavities; they
appear spherical in the centre of the thallus and angular in a section near the
cortex. Branches straight, simple, rare, perpendicular to the cortex. Concep-
tacles spherical, cortical.
Synonym. Stylaella Berchenko 1981.
Suprageneric attributions. Palaeoberesellaea Mamet and Roux 1974.
Family Palaeoberesellaceae trans. Shuysky 1985 for Shuysky (1987).
Stratigraphic range and distribution. Originally described from the Tournaisian
to the Namurian. Known to occur as early as the Middle Devonian. Very
abundant in restricted platforms where it forms baffiestones. Cosmopolitan.
Morocco, Algeria, Spain, France, Great Britain, Belgium, Germany, Russian
Platform, Donbass, Urals, Siberia, Kuzbass, Kolyma, Omolon, China, Aus-
tralia, Alaska, American Cordillera (Alberta, Wyoming, Montana, Idaho),
American midcontinent and Newfoundland.
91. Palaeomicrocodium Mamet and Roux 1983

Type of genus. Palaeomicrocodium devonicum Mamet and Roux 1983. Fras-
nian, Australia.
Diagnosis. Thallus(?) composed of an aggregate of spheres of radial plates. The
formal emendum of Mu and Zhang (1988) claiming that the plates are com-
posed of radiate fibrous calcite is incorrect.
Suprageneric attributions. An algal incertae sedis belonging to the

Microcodiaceae Maslov 1960. Related to Microcodium and Baccanella.
Stratigraphie range and distribution. Eurasia, Australia, Canadian Arctic.
Doubtful as early as Late Silurian (Gotland). Middle Devonian (Eifelian) to
Middle Carboniferous. Same ecological indicator as Microcodium and
Baccanella.
92. Parachaetetes Deninger 1906 emend Mamet and Roux 1977

Type of genus. Parachaetetes tornquisti Deninger 1906. Bathonian, Sardinia.
Diagnosis. Thallus nodular, composed of closely packed, elongated filaments,
subpentagonal in equatorial section, subdivided by horizontal, continuous,
slightly concave partitions. Filaments also divided by longitudinal "septiform
processes". Wall micritic.
Synonym. Tomilithon Maslov 1962 (as subgenus).
Parachaetetes concentrica (Maslov 1956). Devonian-Lower Carboniferous,
Germany, Russian Platform.
Parachaetetes donbassica Berchenko 1981. Devonian-Carboniferous transition,
Donbass.
Parachaetetes garwoodi (Hinde in Garwood 1912). Tournaisian-Visean, Wes-
tern Europe, Arctic Siberia, British Columbia, Alberta (see Plate 3e).
Parachaetetes glenwoodensis (Johnson 1945). Tournaisian, Saskatchewan,
Colorado.
Parachaetetes (Tomilithon) johnsoni Maslov 1962. Late Devonian-Late Tour-
naisian, Belgium, Great Britain, Poland, Donbass, Urals, Arctic Siberia, Kuz-
bass, Australia, Colorado.
Parachaetetes paleozoicus (Maslov 1935). (= Solenopora nexa Maslov 1956).
Lower Carboniferous. France, Donbass, Urals, Arctic Siberia, Alaska, Kansas,
Quebec.
Parachaetetes regularis Konishi 1958. (= Parachaetetes intermedius Konishi
1958). Upper Devonian-Upper Tournaisian. Belgium, Poland, Donbass,
Australia, Alaska, Alberta.
Parachaetetes russiensis (Maslov 1956) (= Solenopora koivense Chuvashov
1965). Devonian to Devonian-Carboniferous transition, Russian Platform,
Urals, Donbass.
Parachaetetes velbertiana (Paul 1938). Tournaisian, Germany.
Suprageneric attribution. Solenoporaceae Pia 1927.
Stratigraphic range and distribution. Cosmopolitan. Very long ranging from
Ordovician to Mesozoic.
"Paradella" Maslov 1956

Type of genus. Paradella recta Maslov 1956. Carboniferous, Kazakhstan.
Diagnosis. Needs revision. The type and the associated Paradella adunca
Maslov 1956 and Paradella ftstulosa Maslov 1956 are probably Carboniferous
phylloid algae belonging to the Anchicodiaceae. A fourth Carboniferous
418 B. Mamet
species, Paradella arcuata Maslov 1956, is a coelosporellid. A fifth species,

Paradella alveata Maslov 1956, is a dissolved thallus of an udoteacean.
In 1981, Kulik published photographs of Paradella recta which are unusable.
They are probably weathered blades of Eugonophyllum.
Paradella adunca re-illustrated by Kulik (1981) (not the type of the genus) can
be differentiated form Eugonophyllum while the re-illustration of Paradella
fistulosa bears no resemblance to Maslov's original drawing.
Paradella arcuata in the concept ofRauser-Chernoussova and Korolyuk (1981)
also differs considerably from Maslov's illustration. The addition of a new
Devonian udoteacean Paradella crassa by Shuysky (1973) and the illustration of
a Devonian Paradella? by Deloffre (1988) (after Kochansky-Devide and Giisic
1971) render the genus unusable without formal emendation.
Suprageneric attributions. Maslov 1956 erected the tribe Paradellae. Kulik
(1981) includes Paradella and Eugonophyllum in that tribe. Shuysky (1987)
attributes Eugonophyllum to his new tribe Ivanoviae while restricting Paradella
to the Paradellae. Deloffre (1988) questionably places Paradella among the
Aciculelleae, but with a Devonian age, which is implausible. Unless the generic
status of Paradella is clarified, its taxonomy remains open to discussion.
Stratigraphic range. See above.
93. Paraepimastopora Roux 1979 (Epimastopora auct)

Type of genus. Epimastopora kansasensis Johnson 1946. Late Carboniferous,
Kansas (see Plate 20).
Diagnosis. Thallus cylindrical, large, unsegmented, with a thick stem and a
slender poorly calcified cortex. Cortex with projections, perforated on both sides
by numerous simple, isodiametric branches. In tangential sections these are
sub cylindrical to subpolygonal (see Roux in press).
Paraepimastopora kanumai Endo in Endo and Kanumai 1954. Upper Car-
boniferous - Permian, Japan, China.
Paraepimastopora urtazymensis (Chuvaskov and Anfimov 1988). Moscovian,
Urals.
Suprageneric attributions. Mastoporae (Pia 1920) trans. Deloffre 1988.
Stratigraphic range and distribution. Bashkirian to Permian. Carnic Alps,
Afghanistan, Urals, Japan, China, Yukon Territory, Canadian Arctic, Amer-
ican midcontinent, New Mexico.
94. Parakamaena Mamet and Roux 1974

Type of genus. Kamaena(?) tenuisepta Mamet and Rudloff 1972. Visean,
Alberta.
Diagnosis. Thallus elongate, cylindrical, rarely ramified by simple bifurcation.
Medulla divided by thin, straight partitions, perpendicular to the cortex. Cells
rectangular, height greater than width. Branches thin, few, simple.
Para kama en a exilis Ivanova 1988. Siberian Platform (Omolon Massif).
Parakamaena irregularis Berchenko 1981. Tournaisian, Donbass, Siberian

Platform.
Suprageneric attributions. Palaeobereselleae Mamet and Roux 1974 according
to Deloffre (1988). Unconvincingly reported to the Anthracoporellopsiae by
Shuysky (1985).
Stratigraphie range and distribution. Scarce. Middle Devonian to Visean.
Belgium, Great Britain, Donbass, Siberian Platform, Omolon Massif, Alaska,
Alberta. The type species is endemic to North America.
95. Parastacheia Marnet and Roux 1977

Type ofgenus. Parastacheia iglii. Mamet and Roux 1977. Late Visean, Algeria.
Diagnosis. Thallus fusiform, encrusted on a central support. Rows of subpa-
rallel laminae with oblique insertion on support. Cells subrounded. Wall
calcareous, yellowish, hyaline.
Suprageneric attributions. Stacheiinae Loeblich and Tappan 1961. New tribe
"Aoujgaliae Chuvashov 1987".
Stratigraphie range and distribution. Scarce. See type of genus.
96. Pekiskopora Marnet 1974

Type of genus. Pekiskopora macqueeni Mamet 1974. Middle Tournaisian,
British Columbia.
Diagnosis. Thallus, a succession of spherical units. Thick stem and meta-
spondyl verticils sharply protruding from the cortex. Medulla expands in
numerous composite vestibules followed by crowns of sixteen branches.
Branches expand slowly in diameter.
Suprageneric attributions. U nconvincingly reported to the Triploporellinae Pia
1920 emend Bassoullet et al. 1979 by Shuysky (1987). Albertaporelleae (Gilven<;
1979) for Deloffre (1988).
Stratigraphic range and distribution. Locally abudant in Tournaisian-Visean.
Endemic to North America. British Columbia, Alberta, Tennessee, New-
foundland.
97. Peristacheia Marnet and Roux 1983

Type of genus. Peristacheia jonesi Mamet and Roux 1983. Late Tournaisian,
Australia.
Diagnosis. Lemon-shaped stacheiin with differentiated thallus. Hypothallus
with rows of quadratic cells. Perithallus composed of ovoid cells. Wall calca-
reous, yellowish hyaline.
Suprageneric attributions. Stacheinaceae Loeblich and Tappan 1961.
Stratigraphic range and distribution. Scarce. Late Tournaisian-Early Visean,
Australia.
98. Petschoria Korde 1951

Type ofgenus. Petschoria elegans Korde 1951. Middle Carboniferous, Urals.
420 B. Marnet
Diagnosis. Thallus ramified, cylindrical, unsegmented. Hypothallus promi-

nent, composed of long longitudinal filaments formed by rectangular cells.
Perithallus poorly developed but of even thickness, composed of very fine
filaments of rectangular cells, perpendicular to the hypothallic tissue.
The genus is restricted to Petschoria elegans. Petschoria antiqua Berchenko 1981
and Petschoria(?) tenuicrustata Shuysky 1973 do not belong to the genus.
Suprageneric attribution. New tribe Petschoriae Chuvashov 1987. Ungdarel-
laceae Maslov 1962.
Stratigraphic range and distribution. Moscovian, Urals, Yukon Territory, New
Mexico. Other reports are incorrect (France, Algeria, Morocco).
Pokorninella Vachard in Perret and Vachard (1977)

See Proninella?
"Polymorphocodium" Derville 1931

Type of genus. Polymorphocodium lapparenti Derville 1931. Visean, France.
Diagnosis. Thallus composed of encrusting asymmetrical, cystoid tubes dis-
posed in more or less regular rows and separated by micritized layers displaying
sponge canals, Bevocastria and birdseyes. Needs revision. Very similar to
"Stylocodium" .
Suprageneric attributions. A codiacean at origin. Pia (1937) thought it was a
consortium. Spongiostromidae Giirich 1906.
Stratigraphie range and distribution. Middle Visean-Basal Bashkirian. France,
Belgium, China, Alberta, Wyoming.
Praedonezella Kulik 1973

See Donezella.
Principia Brenckle 1982

See A rchaolithophyllum.
99. Proninella Reitlinger in Menner and Reitlinger 1971

Type of genus. Proninella tamarae Reitlinger in Menner and Reitlinger 1971.
Givetian, Siberia.
Diagnosis. Thallus encrusting, sinuous. irregular, branching. Tube compressed
or inflated. su bcylindrical, divided by irregular partitions. Wall yellowish, clear,
with thin pores (?)
Proninella gracilis (Vachard in Meissami et al. 1978) (= Proninella enigmatica
Mamet and Roux 1978). Visean. Morocco. France. Tennessee. Alabama.
Proninella strigosa (Vachard in Perret and Vachard 1977). Visean-Early
Bashkirian, Morocco, Spain, France. Iran. Alaska. Oklahoma. Kansas,
Alabama, Tennessee.
Suprageneric attributions. A foraminifer at origin. Transferred to the sponges

by Vachard in Perret and Vachard (1977). An algal incertae sedis for Groves
(1983).
Stratigraphic range and distribution. Occurs in lagoonal conditions associated
with Palaeosiphonocladales and calcispheres. Givetian-Bashkirian. Morocco,
Spain, France, Iran, Afghanistan, Urals, Siberia (Verkoyansk), Alaska, Alberta,
British Columbia, Oklahoma, Kansas, Alabama, Tennessee.
100. Protoumbella Mamet 1970

Type ofgenus. Umbella saccaminiformis Bykova in Bykova and Polenova 1955.
Famennian, Russian Platform.
Diagnosis. Utricle (?) spherical. Exterior smooth, without ornamentation. No
basal fibrous projection. Opening at end of small neck. No known apical lid.
Wall two-layered. A micritic layer and a thick yellowish prismatic layer.
Synonym. Grozdilovella Chermnykh 1972.
Protoumbella decurta (Reitlinger 1966). Devonian-Carboniferous transition,
Armenia.
Protoumbella elliptica (Reitlinger 1966). Devonian-Carboniferous transition-
Visean, Armenia, Urals, American midcontinent.
Stratigraphic range and distribution. Famennian-Visean, Russian Platform,
Urals, Armenia, Kazakhstan, American midcontinent. This is the youngest
known representative of the umbellids.
101. Pseudochaetetes Haug 1.883, emend. Mamet and Roux 1977

Type of genus. Pseudochaetetes champagnensis Peterhans 1929, Jurassic,
France.
Diagnosis. Thallus nodular, composed or closely packed, elongated filaments,
subpentagonal or subhexagonal in equatorial section, sparsely subdivided by
horizontal partitions. Wall micritic.
Pseudochaetetes elliotti (Bilan and Golonka 1973). Lower Carboniferous,
Poland, China.
Pseudochaetetes similis (Paul 1938). Tournaisian, Germany.
Stratigraphie range and distribution. Very long ranging. Ordovician (?) -
Silurian-Jurassic. Cosmopolitan in Northern Hemisphere.
Pseudoepimastopora Endo 1960

See Epimastopa pars and Epimastoporella pars.
102. Pseudogyroporella Endo 1959

Type ofgenus. Pseudogyroporella mizziaeformis Endo 1959. Permian, Japan.
Diagnosis. Thallus moniliform. Stem important. Cortex perforated by very
422 B. Marne!
large vesiculiferous primary branches. They have a trapezoidal form toward the
exterior. Needs revision.
Pseudogyroporella mizziaeformis is present in the Upper Carboniferous of the
Canadian Arctic archipelago.
Suprageneric attribution. Coniporelleae Bassoullet et al. 1979 according to
Shuysky (1987) and to Deloffre (1988).
Stratigraphie range and distribution. Upper Carboniferous-Lower Permian.
Yugoslavia, China, Japan, Canadian Arctic.
103. Pseudohedstroemia Mamet and Roux 1978

Type ofgenus. Pseudohedstroemia polyfurcata. Mamet and Roux 1978. Visean,
Tennessee.
Diagnosis. Nodule formed by radially disposed cylindrical, micritized sheaths
of two types. First group of variable tubes is ramified by small angle dicho-
tomies. It contains thin wavy tubes with double or triple branching.
Suprageneric attribution. Nodular codiacean? Cyanobacteria?
Stratigraphie range and distribution. Tournaisian-Namurian. Alaska, Alberta,
Utah, Tennessee.
104. Pseudoissinella Marnet and Rudloff 1972

Type of genus. Pseudoissinella alaskaensis Marnet and Rudloff 1972. Visean,
Alaska.
Diagnosis. Thallus cylindrical, erect, with multiple branchings at right angle.
Cortex, in fibro-radial yellowish calcite, perforated by scarce thin unrarnified
branches.
Suprageneric attribution. Tribe Issinelleae Deloffre 1987 according to Deloffre
(1988).
Stratigraphic range and distribution. Scarce in Devonian. Tournaisian to Lower
Namurian. Belgium, Donbass, Alaska, Alberta, British Columbia.
105. Pseudokamaena Marnet in Petryk and Marnet 1972

Type ofgenus. Pseudokamaena armstrongi Marnet in Petryk and Marnet 1972.
Visean, Alaska.
Diagnosis. Thallus cylindrical, dichotomous, with faint outer constrictions.
Stem divided by short transverse partitions. Cells ovoid to spherical. Wall
calcareous, yellowish, hyaline, perforated by thin, unrarnified straight pores.
Pseudokamaena boulderensis Marnet in Petryk and Marnet 1972. Visean,
Alberta.
Suprageneric attributions. Palaeobereselleae Mamet and Roux 1974. Exvota-
riselleae Shuysky 1985 according to Shuysky (1987).
Stratigraphie range and distribution. Mostly in Lower Carboniferous lagoons
associated with Palaeosiphonocladales and calcispheres. Scarce in Devonian
(Eifelian-Frasnian). Mostly Tournaisian-Lower Bashkirian. Belgium, France,
Donbass, Urals, Siberia, Afghanistan, Kolyma, Omolon Massif, Australia,

Canadian Arctic, Alaska, Alberta, Montana, Wyoming.
106. Pseudokomia Racz 1965

Type of genus. Pseudokomia cansecoensis Racz 1965. Moscovian, Spain.
Diagnosis. Thallus cylindrical, highly calcified. Hypothallic cells rather regular.
Perithallic cells perpendicular to hypothallus, heavily calcified and very regular.
The genus forms the morphological link between Epistacheoides and Petschoria.
Suprageneric attribution. U ngdarellaceae Maslov 1956. New tribe Petschoriae
for Chuvashov 1987.
Stratigraphie range and distribution. Moscovian, Spain, Canadian Arctic, New
Mexico.
107. Pseudonanopora Mamet and Roux 1975

Type of genus. Pseudonanopora stockmansi Mamet and Roux 1975. Middle
Visean, Belgium.
Diagnosis. Thallus tubular, cylindrical, with slight constrictions. Stem divided
by rudimentary pseudo-septa. Cortex perforated by randomly disposed,
straight, simple branches.
Suprageneric attribution. Anthracoporellopsiae Shuysky 1985 according to
Shuysky (1987).
Stratigraphic age and distribution. Devonian-Carboniferous transition to Ser-
pukhovian. Algeria, Morocco, Belgium, Donbass, Siberia (Kolyma).
108. Pseudosolenopora Mamet and Roux 1977

Type of genus. Solenopora owodenkoi Chanton-Giivenc; 1972. Late Visean,
Morocco.
Diagnosis. Thallus nodular, composed of closely packed elongated, contin-
uous, filaments, subpentagonal to sub hexagonal in equatorial section. Ho-
rizontal partitions absent. Wall micritic.
Suprageneric attributions. Solenoporaceae Pia 1927. New tribe Pseudosolen-
oporaceae Chuvashov 1987 (ill-founded).
Stratigraphie range and distribution. Late Devonian-Lower Namurian. Mo-
rocco, Belgium, Poland, Germany, Afghanistan, Australia, Alberta, British
Columbia.
109. Pseudostacheoides Petryk and Mamet 1972

Type ofgenus. Pseudostacheoides loomisi Petryk and Mamet 1972. Late Visean,
Alberta.
Diagnosis. Thallus encrusting, irregular, composed of closely-packed thick,
porous, undulating, horizontal elements and of thin and scarce vertical
elements. Cells elongated, fiat to undulating. Wall calcareous, yellowish, per-
forated by thin unramified pores. Central part ofthallus usually poorly calcified
and micritized.
424 B. Marne!
Suprageneric attribution. Stacheiinae Loeblich and Tappan 1961 emend. New

tribe Pseudostacheoideae Chuvashov 1987.
Stratigraphic range and distribution. Early Visean-Lower Namurian.
"Cosmopolitan". Morocco, Algeria, Great Britain, Belgium, Afghanistan,
Alaska, Alberta, British Columbia, Alabama.
Pseudoumbella Berchenko 1971

See Biumbella Mamet 1970.
llO. Pseudovelebitella Mamet and Roux 1978

Type of genus. Velebitella simplex Kochansky-Devide 1964. Early Permian,
Yugoslavia.
Diagnosis. Thallus composed of a succession of barrel-shaped units, with a
central cylindrical stem. Medulla expands into short vestibules. Tufts of five to
six primary phloiophore branches are localized in the central part of the units.
Pseudovelebitella pareyni (Mamet and Roux 1975). Visean, Algeria, Great
Britain, Russian Platform.
Pseudovelebitella simplicissima (Vachard in Massa and Vachard 1979). Mos-
covian, Algeria.
Suprageneric attribution. Velebitelleae (Vachard 1977) Mamet and Roux 1983.
Stratigraphic range and distribution. Visean-Early Permian. Algeria, Great
Britain, Yugoslavia, Russian Platform.
"Pycnostroma" Giirich 1906

Type of genus. Pycnostroma densius Giirich 1906. Visean, Belgium.
Diagnosis. Oncolite? formed by a spongiostromid encrusting a central
uncalcified support (now filled by cement or by sediment). Tissue composed of
micritized and dissolved sheaths and a loosely interwoven meshwork of at least
two types of algal tubes. Needs redescription.
Other Carboniferous species. The micritized part of Pycnostroma spongilliferum
Giirich 1906 has in situ sponge spicules.
Suprageneric attribution. Spongiostromidae. An association of cyanobacteria
and sponges.
Stratigraphic range and distribution. Known from Ordovician to Carboniferous
(Namurian). Morocco, Belgium, France, Alberta, Nevada, Tennessee.
Ill. Quasiumbella Poyarkov 1965

Type of genus. Umbella rotunda Bykova III Bykova and Polenova (1955).
Famennian, Russian Platform.
Diagnosis. U tricle(?) nearly spherical. External wall smooth. without or-
namentation. No basal projections. Apical lid present.
Quasiumbella cutis (Conil and Lys 1964). Famennian, Devonian-Carboniferous

transition, Belgium.
Quasiumbella fragilis Berchenko 1981. Devonian-Carboniferous transition,
Donbass.
Quasiumbella gravis (Coni1 and Lys 1964). Famennian, Devonian-Carbonife-
rous transition, Belgium, Donbass.
Stratigraphie range and distribution. As is the case for most umbellids, common
in the Late Devonian and peters out in the Tournaisian. Belgium, Poland,
Russian Platform, Donbass, Urals, Armenia, Iran, Siberia, Kolyma, Australia,
Alberta.
Quasiumbelloides Berchenko 1971

see Biumbella.
112. Queenslandella Mamet and Roux 1983

Type of genus. Queenslandella jenkinsi Mamet and Roux 1983. Visean,
Australia.
Diagnosis. Metaspondyl dasyclad composed of a succession of annular units.
Central stem subcylindrical with expansions at the junction of each unit.
Composite vestibule. First order ramifications. Branches numerous, slightly
arched and ph1oiophore (see Plate 1b).
Suprageneric attribution. Albertaporelleae (Guvenc; 1979) Mamet and Roux
1981 according to Deloffre (1988).
Stratigraphic range and distribution. Middle to Late Visean. Endemic to
Australia.
"Radiosphaera" Reitlinger 1957

(or "radiospherid calcispheres" Stanton 1967)
Type ofgenus. Radiosphaera basilica Reitlinger 1957. Devonian-Carboniferous
transition, Russian Platform.
Diagnosis. When disarticulated, observed as calcareous hollow spheres. Wall
two-layered, a thin internal micritic layer and a thicker pseudo-fibrous wall, with
coarse tapering prisms. No aperture. In situ, forms grape-like clusters of
loosely-packed calcispheres around a noncalcified axis (see Mamet and Rudloff
1972). At origin (1957) considered as "undetermined spheres" without clear
reference to botanical or zoological origin. (Most Russian authors consider the
taxon as a foraminifer and the genus is thus a homonym of the Protozoan
Radiosphaera Jorgensen 1905). A diagenetic stage ofTuberitinidae for Vachard
and Tellez-Giron (1986).
Carboniferous species. There are many extant "species" in the literature which
appear as ill-defined as the genus. Needs revision.
Suprageneric attributions. See Calcisphaera .for discussion. The original Cal-
cisphaera sol Williamson 1881 is a radiosphaere.
426 B. Mamet
Stratigraphic range and distribution. Givetian-Visean. Very common in res-

tricted environments. Cosmopolitan in Northern Hemisphere.
113. Rectangulina Antropov 1959

Type ofgenus. Syniella tortuosa Antropov 1950. Frasnian, Russian Platform.
Diagnosis. Algal clast composed of bundles of subparallel, angular, unseg-
mented, straight tubes. Wall(?) micritic.
Suprageneric attributions. Controversial. Foraminifer (Chuvashov 1965), co-
prolite (Conil et al. 1969), alga (Antropov 1959, Mamet et al. 1987, Luchinina
1987), incertae sed is (Vachard 1981).
Rectangulina geniculosa (Bykova 1961). Frasnian-Early Namurian. Great
Britain, Belgium, Russian Platform, Urals, Siberia, Tennessee, Arkansas.
Stratigraphic range and distribution. Ordovician? - Carboniferous. Quite
scarce. Libya, Morocco, Algeria, France, Belgium, Great Britain, Germany,
Poland, Russian Platform, Urals, Kazakhstan, Afghanistan, Siberia, Australia,
Canadian Arctic, Arkansas, Alabama, Tennessee, Georgia, Nova Scotia.
114. Renalcis Vologdin 1932 emend. Mamet and Roux 1983

Type ofgenus. Renalcis granosus Vologdin 1932. Cambrian, Siberia.
Diagnosis. Thallus an aggregate of hollow, inflated cells. Forms simple,
straight, rectilinear colonies to complex botryodal aggregates. "Wall" micritic,
but not directly secreted, dark, often fissured by retraction.
Synonyms. Chabakovia Vologdin 1939, Nubecularites Maslov 1937 (pars),
Eniseiella Maslov in Malakhova 1959, Izhella Antropov 1955, Nephelostroma
Dangeard and Dore 1957 and Shuguria Antropov 1959.
Suprageneric attributions. Controversial. Foraminifer (family Rena1cidae
Riding and Brasier 1975), red alga (solenoporid with "quadrangular cells" of
Korde 1959), cyanophyte (as chroococcales, Chabakoviaceae Korde 1969),
chroococcophyceae for Luchinina (1987). Consensus is usually to classify
Renalcis among cyanobacteria (see Saltovskaya 1984 for discussion).
Stratigraphic range and distribution. Proterozoic. Very abundant from Cam-
brian to Devonian. Peters out in Lower Carboniferous. Cosmopolitan, but plays
no important sedimentological role after the Frasnian.
115. Retephycus Johnson and Konishi 1956

Type of genus. Retephycus globosus Johnson and Konishi 1956. Late Tour-
naisian, Saskatchewan.
Diagnosis. Thallus(?) composed of closely packed algal filaments. Tubes are
interwoven, appear to branch and are reported to have an irregular outline in
transverse section. Needs revision.
Suprageneric attribution. Originally placed in the chlorophytes.
116. Richella Mamet and Roux 1987

Type of genus. Richella incrustata Mamet and Roux 1987. Mid-Moscovian,
Canadian Arctic (Ellesmere) (see Plate lk).
Diagnosis. Thallus encrusting, composed of semi-cylindrical, erratically in-
terwoven, heavily calcified tubes, attached at their base, and perforated by
straight, coarse, undivided branches.
Suprageneric attribution. Algal incertae sedis. In growth form, similar to
Wetheredella.
Stratigraphie range and distribution. Namurian-Moscovian. Tennessee, Can-
adian Arctic.
Roquesselsia Termier, Termier and Vachard 1977

See Epistacheoides.
Samarella Maslov and Kulik 1956

See Uraloporella.
117. Sandoella Mamet and Roux 1978

Type of genus. Sandoella fowleri Mamet and Roux 1978. Late Visean,
Tennessee.
Diagnosis. Thallus subcylindrical, verticillated, a succession of irregularly
shaped units. Medulla important. Cortex with ovoid cavities followed by
bundles of straigh t branches.
Suprageneric attribution. Does not fit any extant tribe of Diploporaceae.
Stratigraphic range and distribution. Visean, Alberta, Tennessee.
Shamovella Rauser-Chernoussova 1951

See Tubiphytes.
Shartymophycus Kulik 1973

See Fasciella.
"Shermanophycus" Johnson 1940
Type of genus. Shermanophycus gouldi Johnson 1940. Upper Carboniferous,

Colorado.
Diagnosis. An algal biscuit. Needs revision. See Stylophycus, Gouldina.
Shuguria Antropov 1950

See Renalcis.
428 B. Marnet
118. Sinarella Chuvashov in Chuvashov and Anfimov (1988)

Type of genus. Sinarella delicata Chuvashov and Anfimov 1988. Moscovian,
Urals.
Diagnosis. Thallus important, cylindrical with poorly calcified thin cortex and
big stem. Wall perforated by triangular pores, protruding from the outside
periphery.
Suprageneric attribution. Cyclocrinaea Pia 1920.
Sinustacheoides Termier, Termier and Vachard 1977

See Stacheoides.
119. Solenopora Dybowski 1877 emend. Mamet and Roux 1977

Type of genus. Solenopora spongioides Dybowski 1877. Ordovician erratic
block, Baltic.
Diagnosis. Thallus nodular, composed of closely packed elongated filaments
subangular in equatorial section, subdivided by septiform processes. Wall
micritic. Conceptacles ovoid to spherical.
Solenopora dionantina Pia 1937. Visean. Great Britain, Poland, Alaska.
Most Carboniferous species of the literature ought to be transferred to Para-
chaetetes.
Stratigraphic range and distribution. Very long ranging. Ordovician-Mesozoic.
Cosmopolitan. Forms the "Solenopora community" ofFltigel (1977).
120. Sphaerinvia Vachard 1981

Type of genus. Sphaerinvia piai Vachard 1981. Devonian-Carboniferous
transition, Afghanistan.
Diagnosis. Hollow calcitic sphere, with functional opening (basal or apical?)
attached to a calcified tube. Wall two-layered, an inner micritic layer with
radiating parietal expansions, and a segmented yellowish lamellar outer layer.
Suprageneric attribution. Incertae sedis. Vachard claims that the genus has no
opening, but it is conspicuous in the material published by Bogush (1984).
Similar material was considered as belonging to the Charophytes by Pia (1937)
and a reasonable case for the same attribution has been made by Bogush (1984).
However, Sphaerinvia's wall is different from that of bona fide Palaeozoic
charophytes.
Stratigraphic range and distribution. Devonian-Carboniferous transition -
Early Visean, Belgium, Great Britain, Urals, Afghanistan, Kazakhstan, Siberia,
Omolon. Very common in Alaska, Yukon Territory, British Columbia, Alberta,
New Mexico, Arizona, Utah.
121. Sphaerocodium Rothpletz 1890 emend Wray 1967

(= Rothpletzella Wood 1948 OBJ.)
Type ofgenus. Sphaerocodium bornemanni Rothpletz, Trias, Austria (redefined

as Sphaerocodium gotlandicum Rothpletz 1908).
Diagnosis. Thallus composed of palisades of highly dichotomous cylindrical
tubes. Bifurcation is at low angle, always in the same plane and the tubes remain
welded. In axial section, tubes are ovoid and their succession forms a charac-
teristic "rosary". Wall micritic.
Synonyms. Rothpletzella Wood 1948, Coactilum Maslov 1956.
Suprageneric attributions. Algal incertae sedis. Wray attributed it to the Po-
rostromata Pia 1927. Could be linked to the Proauloporales Luchinina 1975.
Uncorrectly attributed to the Girvanellaceae by Luchinina (1987) (as
Rothpletzella ).
Stratigraphic range and distribution. Very common in Lower and Middle
Palaeozoic. Forms bindstones in association with megafauna. Also forms
baffiestones in mud-mounds. Last common occurrence is Frasnian and the
genus is uncommon above that level. (Illustrated in Mamet and Roux 1975b
from the Visean of Germany).
122. Sphaeroporella Antropov 1967

Type of genus. Sphaeroporella aksubaica Antropov 1967. Devonian-Carbon-
iferous transition, Russian Platform.
Diagnosis. Thallus(?) encrusting, an aggregate of ovoid or reniform cystoid
cells. Wall two-layered, outside wall micritic with an inner pseudo-fibrous
perforated layer.
Sphaeroporella conchatiformis (Malakhova and Chuvashov 1965). Devonian-
Carboniferous boundary, Urals, Siberia (Kuzbass), China.
Sphaeroporella tchussovensis (Malakhova and Chuvashov 1965). Devonian-
Carboniferous boundary, Urals, China.
Suprageneric attribution. Algal? incertae sedis. Wetheredellidae Vachard in Dil
et al. (1977), trans. nov., Wetheredellaceae, emend. Berchenko in Shuysky
(1987). Attributed to the new tribe Wetheredelleae Berchenko 1987. Akin to
Asphaltina Mamet in Petryk and Mamet 1972, Cateniphycus Maslov 1960 and
Wetheredella Wood 1948. Needs revision.
Stratigraphic range and distribution. Uncommon. Described from Early
Devonian to Visean. Ireland, Great Britain, France, Belgium, Siberia
(Kuzbass), China, Australia, Alberta, Tennessee.
123. Sphinctoporella Mamet and Rudloff 1972

Type ofgenus. Sphinctoporella lisburnensis. Mamet and Rudloff 1972. Middle
Visean, Alaska.
Diagnosis. Thallus elongated, subpentagonal in cross-section with slight outer
constrictions and much more accentuated divisions ofthe stem. Poorly calcified
cortex contains two or three rows of spherical cavities ("sporangial cavities",
"endospores").
Suprageneric attribution. Aciculelleae Bassoullet et al. 1979 for Shuysky (1987)
and Deloffre (1988).
430 B. Mamet
Stratigraphic range and distribution. Endemic to Alaska where it is restricted to

the Visean.
Spinumbella Platonov 1974

See Elenia.
"Spongiostroma" Gurich 1906

Type o[genus. Spongiostroma maeandrinum Gurich 1906. Visean, Belgium.
Diagnosis. Algal biscuit with dissolved tissue of Bevocastria, leaving a pelle-
toidal meshwork counterprint. Needs revision.
Other Carboniferous "species".
"Spongiostroma" bacilli[erum, "S", ovuli[erum and "S", granuli[erum (all by
Gurich 1960) are former exposed algal mats, heavily burrowed and cracked.
Suprageneric attributions. Spongiostromid Gurich 1906. Tentatively attributed
to the Rivulariaceae by Ischenko (1985).
Stratigraphic range and distribution. Needs revision.
124. Stacheia Brady 1876

Type o[genus. Stacheia marginuloides Brady 1876 (= Stacheia pupoides Brady
1876). Visean, Great Britain.
Diagnosis. Thallus cylindrical to fusiform, encrusting a central support, and
formed by a succession of overlapping arched rows of subquadratic cells. Wall
hyaline, yellowish, unperforated.
Stacheia acervalis Brady 1876. Visean, Great Britain.
Suprageneric attributions. Originally described as a foraminifer. Incorrectly
transferred to the aoujgaliid sponges by Vachard in Perret and Vachard (1977).
Stacheiinae for Mamet and Roux (1977). Reported to the new tribe Mametellae
by Chuvashov (1987).
Stratigraphic range and distribution. Common from Visean to Bashkirian.
Reaches the Moscovian in the Canadian Arctic. Morocco, Great Britain,
France, Belgium, Germany, Poland, Donbass, Alaska, Canadian Arctic, Al-
berta, Arkansas.
125. Stacheoidella Mamet and Roux 1987

Type o[ genus. Stacheoides(?) spissa Petryk and Mamet 1972. Visean, Alberta.
Diagnosis. Thallus encrusting on support. Perithallic and hypothallic cells well
differentiated. H ypothallus thin, composed of one or two rows of subspherical
cells. Perithallus, continuous rows of irregular cells. Wall yellowish, hyaline.
Suprageneric attribution. Stacheiinae Loeblich and Tappan 1961 emend.
Stratigraphic range and distribution. Visean to Moscovian. Endemic to North
America. Canadian Arctic, Alaska. American Cordillera, American Mid-
continent.
126. Stacheoides Cummings 1955 emend. Petryk and Mamet 1972

Type ofgenus. Stacheia polytrematoides Brady 1876. Visean, Great Britain.
Diagnosis. Thallus encrusting, very irregular, with protuberances. Grossly
concentric arrangement of irregular rows of cells. Cells also irregular with walls
of uneven thickness. Wall yellowish, hyaline, unperforated.
Other Carboniferous species
Stacheoides cannindahensis Mamet and Roux 1983. Late Visean, Australia.
Stacheoides meandriformis Mamet and Rudloff 1972. Abundant from Visean to
Moscovian. Extends to Permian in Canadian Arctic. Spain, Siberian Platform,
Kuzbass,Alaska, Yukon Territory, British Columbia, Alberta, Idaho, Montana,
Utah, American midcontinent.
Stacheoides papillata Cummings 1955. Visean. Morocco, France, Great Britain.
Stacheoides tenuis Petryk and Mamet 1972. Visean-Namurian. Algeria, Mo-
rocco, Spain, Russian Platform, Donbass, Urals, Siberia (Kuzbass, Ver-
koyansk), China, Alaska, Alberta, British Columbia, Idaho, Utah, American
midcontinent, Tennessee. Extends up to the Permian in the Canadian Arctic.
Suprageneric attributions. Originally described as a foraminifer. Incorrectly
transferred to the aoujgaliid sponges by Vachard in Perret and Vachard (1977).
Belongs to the Stacheiinae for Mamet and Roux (1977) and to the new tribe
Aoujgaliae for Chuvashov (1987).
Stratigraphic range and distribution. Doubtful in Late Tournaisian? Abundant
from basal Visean (Zone II) to the Bashkirian. Still observed in the Early
Permian of the Canadian Arctic Archipelago. Algeria, Morocco, Spain, Great
Britain, France, Belgium, Russian Platform, Urals, Donbass, Afghanistan,
China, Australia, Canadian Arctic, Alaska, Yukon Territory, British Columbia,
Alberta, American Cordillera, American midcontinent, Tennessee.
"Stipulella" Maslov 1956

Type of genus. Stipulella fascicularis Maslov 1956. Lower Carboniferous,
Russian Platform.
Diagnosis. Algal clast composed of bundles of straight, unsegmented, calcified,
cylindrical sheats. Each bundle has an angular outline. Similar to Rectangulina
but the tubes here are cylindrical.
Suprageneric attribution. Algal incertae sedis. Ranged among the cyanophytes
by Luchinina (1987).
Stratigraphic range and distribution. Scarce, Late Tournaisian to Early
Namurian. Great Britain, Belgium, France, Russian Platform, Australia.
Stylaella Berchenko 1981

See Palaeoberesella.
"Stylocodium" Derville 1931

Type of genus. Stylocodium rhopaloides Derville 1931. Visean, France.
Diagnosis. Thallus columnar, composed of concentric rows of cystoid tubes
432 B. Marnel
(Aphralysia? or Sphaeroporella?). Needs revision.

Suprageneric attribution. Wetheredelleae Berchenko in Sh uysky (1987).
Stratigraphic range and distribution. Scarce. Ireland, France, Belgium, Alaska.
Visean. Needs revision.
"Stylophycus" Johnson 1940

Type ofgenus. Stylophycus carbonarius Johnson 1940. Upper Carboniferous,
Colorado.
Diagnosis. Algal biscuit of uncertain affinity. Needs revision.
Stylophycus calcarius Johnson 1940. Upper Carboniferous, Colorado.
Suprageneric attribution. See Shermanophycus and Gouldina.
Stratigraphie range and distribution. See type of genus. Needs revision.
Subkamaena Berchenko 1981

See Kamaena
127. Sycidium Sandberger 1849

Type ofgenus. Sycidium reticulatum Sandberger 1849. Devonian, Germany.
Diagnosis. Gyrogonite oblate spheroidal, non-coronulate, with meridional ribs
and rectangular to hexagonal polygons. Small basal and important apical
openings. Wall foliated, protruding at each rib.
Carboniferous species
Sycidium faveatum Peck 1934. Basal Carboniferous, Iowa, Missouri.
Also doubtful reports of Sycidium melo Sandberger 1880 and Sycidium panderi
(Ehrenberg 1858) at the Devonian-Carboniferous transition in the Donbass
(Berchenko 1981).
Stratigraphic range and distribution. Abundant in the Middle- Late Devonian of
Eurasia and North America. Still present at the base of the Carboniferous in the
same regions.
128. Taldykites Chuvashov 1988

Type of genus. Ta/dykites mugodjaricus Chuvashov and Anfimov 1988. Mos-
covian, Urals.
Diagnosis. Needs further study. The type material does not show the compo-
sition of the central tubes. Lateral tubes disposed as radiating fans. Tubes
bifurcate at various angles and show some constrictions. They terminate as thin
peripheral branches. Needs revision.
Suprageneric attribution. According to original description, belongs to the
Dimorphosiphoneae Shuysky 1985. The description is that ofa udoteacean.
Tomilithon Maslov 1962

See Parachaetetes (as subgenus).
Trinodella Maslov and Kulik 1956

See Dvinella (as subgenus).
129. Tubiphytes Maslov 1956

Type of genus. Tubiphytes obseurus Maslov 1956. Lower Permian, Urals.
Diagnosis. Thallus(?) massive, cylindrical, branching, encrusting on central
support, now preserved as cement-filled cavity. Tissue heavily micritized, with
darker and clearer concentric "zones". Although filaments have often been
reported the tissue is probably composed of rectangular elongated cells.
Synonyms. Shamovella Rauser-Chernoussova 1951, NOM. NUD, Nigriporella
Rigby 1958, "Hikorocodium" sensu Flilgel non Endo 1951.
Supragenerie attributions. Controversial. Hydrozoan for Rigby 1958. Incertae
alga for Maslov 1956. Cyanobacteria (Girvanellaceae Luchinina 1975) ac-
cording to Luchinina (1987). Red alga for Kochansky-Devide (1970).
Stromatolite for Chuvashov and Dyupina (1973). An algal incertae sedis for
Mamet et al. (1987). See also Roux (1985).
Stratigraphic range and distribution. Forms the "Tubiphytes community" of
Flilgel (1977). Often associated with phylloid algae in organic build-ups.
Common on open shelf. Known from Middle Carboniferous to Triassic. Car-
boniferous (to Early Permian) occurrences are in Spain, Yugoslavia, Turkey,
Urals, Kazakhstan, Iran, Afghanistan, China, Japan, Siberia, Canadian Arctic,
American Cordillera, American midcontinent, Mexico.
130. Ungdarella Korde 1951 non Maslov "1950" or 1956

Type of genus. Ungdarella. eonservata Korde 1951 (non" Ungdarella uraliea
Maslov 1950", nee "Ungdarella uraliea Maslov 1956"). Upper Carboniferous,
Urals.
Diagnosis. Thallus elongated, ramified. Hypothallic tissue composed of a row
of rectangular cells. Perithallic tissue, much more conspicuous, composed of
numerous, thin, long rows of subquadratic cells, oblique to the hypothallus.
Wall calcareous, yellowish, hyaline.
Ungdarella(?) americana Toomey and Johnson 1968. Namurian-Bashkirian,
Algeria, Russian Platform, New Mexico.
Ungdarella gracilissima Kulik 1973. Middle Carboniferous, Urals.
Ungdarella peratrovichensis Mamet and Rudloff 1972. Namurian-Early Per-
mian, Alaska, British Columbia. Canadian Arctic.
Ungdarella uraliea Maslov 1956 (= Ungdarella deceanglorum Elliott 1970).
Common from Late Visean-Moscovian. Scarce up to the Permian. Libya,
Morocco, Algeria, Spain, France, Great Britain, Austria, Turkey, Irak, Russian
Platform, Urals, Donbass, Afghanistan, Kazakhstan, Australia, China, Can-
adian Arctic Archipelago, Alaska, British Columbia, New Mexico, Arizona (see
Plate 3r).
Suprageneric attributions. Ungdarellaceae Maslov 1956 or new tribe Ungda-
relleae Chuvashov 1987.
434 B. Mamel
Stratigraphic range and distribution. Very common. Forms boundstones in the

Middle Carboniferous in a manner similar to Komia. Latest Visean to Mos-
covian. Scarce up to the Permian. Libya, Algeria, Morocco, Spain, France,
Great Britain, Austria, Turkey, Irak, Russian Platform, Donbass, Urals,
Kazakhstan, Afghanistan, Australia, China, Canadian Arctic, Alaska, British
Columbia, New Mexico.
131. Unjaella Korde 1951

Type ofgenus. Unjaella ovata Korde 1951. Late Carboniferous, Northern Urals.
Diagnosis. Thallus spindle-shaped, with thin cortex and important medulla.
Pores subhexagonal in transverse section with honey-comb arrangement and
regular interpores.
Suprageneric attribution. Tribe Cyclocrineae Pia 1927, subtribe Mastoporinae
Pia 1927 according to Berchenko and Saltovskaja in Shuysky (1987).
Stratigraphic range and distribution. Only known by the type.
132. Uraloporella Korde 1950

Type of genus. Uraloporella variabilis Korde 1950. Middle Carboniferous,
Northern Urals (see Plate 2h).
Diagnosis. Thallus elongated, cylindrical, unsegmented. Diaphragms present.
Cortex perforated by numerous short, thin, "blind" cortical branches. The
thallus was originally surrounded by a mucilaginous coating, now preserved as
a continuous cement layer.
Synonym. Samarella Maslov and Kulik 1956.
Uraloporella sieswerdai (pars) Racz 1965. Moscovian, Spain. Part of Racz's
original material belongs to Uraloporella and part to Zidella Saltovskaya 1984.
Suprageneric attributions. Unconvincingly classified as hypercalcified sponge
by Termier et al. (1977), as foraminifer (Riding and Jansa 1976) or as micro-
problematicum. (Wright 1982). Belongs to the Palaeosiphonocladales of
Shuysky (1985). Placed among the Beresellaceae (Maslov and Kulik 1956)
Shuysky 1985 by Shuysky (1987) and by Deloffre (1988). The tribe Uralopo-
relleae was also proposed by Shuysky (1985).
Stratigraphic range and distribution. Very common and often associated with
Beresella or Dvinella. Total range Namurian to Early Permian. Very abundant
from Late Bashkirian to Moscovian. Devonian and Early Carboniferous
references are to be discarded. Spain, Turkey, Russian Platform, Urals, Oon-
bass, Kazakhstan, Siberia, China, Canadian Arctic, Alaska.
Velebitella Kochansky-Devide 1964

Type ofgenus. Velebitella triplicata Kochansky-Oevide 1964. Middle Permian,
Yugoslavia. Although this genus is sometimes mentioned in the Carboniferous,
it is restricted to the Permian. Carboniferous taxa such as "Velebitella" simplex
Kochansky- Oevide 1964 ough t to be transferred to Pseudovelebitella.
133. Vermiporella Stolley 1893 sensu Pia 1927

Type ofgenus. Vermiporella fragilis Stolley 1893. Ordovician erra tic, Germany.
Diagnosis. Thallus cylindrical, irregularly ramified. Stem unconstricted. Cor-
tex aspondyl, perforated by numerous straight cylindrical branches.
Vermiporella(?) shartymensis Kulik 1973. Middle Carboniferous, Urals.
Suprageneric attributions. New tribe Vermiporelleae Saltovskaja in Shuysky
1987. Dasyporellinae (Pia 1920) according to Deloffre (1988).
Stratigraphic range and distribution. Abundant in Ordovician-Silurian-
Devonian. Very scarce in the Carboniferous. Doubtful in the Visean of Spain,
Bashkirian of the Urals and in the Moscovian of the Canadian Arctic.
134. Wetheredella Wood 1948

Type of genus. Wetheredella silurica Wood 1948. Wenlock, Great Britain (see
Plate 3m).
Diagnosis. Thallus(?) encrusting, an aggregate of intertwined, flexuous, un-
divided regular tubes. Perforated two-layered wall, external layer micritic,
internal layer pseudo-fibro-radial.
Wetheredella starobeshevense Berchenko 1983. Serpukhovian-Bashkirian,
Donbass.
Suprageneric attributions. The original family Wetheredellaceae Vachard 1976
is a composite association offoraminifers, algae and incertae sedis. It has been
emended by Berchenko (in Shuysky 1987) to fit the Dasycladales. Doubtful alga
for Riding 1977, cyanophyte for Copper 1976, cyanobacterium for Ischenko
(1985).
Stratigraphic range and distribution. Abundant in Silurian-Devonian in car-
bonate build-ups. Very scarce in the Carboniferous. Isolated occurrences in the
Visean (Great Britain and Turkey) and in the Middle Carboniferous (Don bass ).
135. Windsoporella Mamet and Rudloff 1972

Type of genus. Windsoporella radiata Mamet and Rudloff 1972, Namurian
Zone 17, Maritime Provinces of Canada (see Plate 3j).
Diagnosis. Thallus cylindrical, composed of a succession of units separated by
constrictions. Annulated stem. Projections of medulla in the centre of each unit
form around a vestibule from which expand tufts of coarse, simple, phloiophore
branches.
Suprageneric attribution. Velebitelleae (Vachard 1977) Mamet and Roux 1983
according to Deloffre (1988). Unconvincingly reported to the Diploporeae (Pia
1920) Bassoullet et al. 1979 by Shuysky (1987).
Stratigraphic range and distribution. Endemic to the latest Visean-earliest
Namurian of the Maritime Provinces of Canada (Nova Scotia, Newfoundland
shelf, Magdalen Islands of Que bec).
436 8. Mamet
136. Yukonella Mamet and Rudloff 1972

Type of genus. Yukonella bamberi Mamet and Rudloff 1972. Visean, Yukon
Territory.
Diagnosis. Calcareous irregular tube with some external and internal con-
strictions. Wall micritic, perforated by numerous erraticaly disposed pores.
Suprageneric attribution. Algal incertae sedis (Shuysky 1987).
Stratigraphic range and distribution. Early Carboniferous. Endemic to the
Yukon Territory, Alaska and the Omolon Massif (Siberia).
137. Zaporella Racz 1965

Type of ten us. Zaporella cantabriensis Racz 1965. Moscovian, Spain.
Diagnosis. Thallus club-shaped. Stem cylindrical, undivided. Cortex heavily
calcified, perforated by randomly disposed cylindrical pores perpendicular to
the exterior. Secondary branches grow out of the ultimate ends of the primary
branches. Dichotomy is variable.
Remark. Deloffre (1988) and Shuysky (I 987} put Zaporella in synonymy with
Anthracoporella. Both genera are recognized here as the branching of the pores
is different.
Suprageneric attribution. Dasyporellinae ex Dasyporelleae (Pia 1920) Bas-
soullet et al. 1979.
Stratigraphic range and distribution. Moscovian of Spain. Could occur in the
Early Permian of China (Zaporella baxoensis Mu 1982).
138. Zidella Saltovskaya 1984

Type of genus. Zidella maxima Saltovskaya 1984. Visean, Tadzhikhistan.
Diagnosis. Thallus, a long cylindrical tube, dichotomous, with slight internal
and external constrictions. Cortex perforated by numerous "blind", thin, cor-
tical branches. The thallus may have been surrounded by a mucilaginous
coating, now preserved as a continuous cement layer. The genus is close to
Uraloporella Korde 1950.
Zidella minor Ivanova 1988. Visean, Urals.
Suprageneric attribution. Uraloporelleae Shuysky 1985. Erroneously Beresel-
leae for Ivanova (1988).
Stratigraphic range and distribution. Visean-Moscovian, Spain, Urals,
Tadzhikhistan.
5 Stratigraphic Distribution
Figures 2 and 3 record the stratigraphic range of the microfiora. As previously

mentioned, all the simple cyanobacteria or Garwoodiaceae are very long
ranging, some already known in the Precambrian and often observed
throughout the whole Palaeozoic. As they do not evolve a pprecia bly they can be
proposed as reliable indicators for environmental comparison through the

Palaeozoic. The same applies to the solenoporids.
Other algal groups are more restricted in time.
The umbellinids are mostly Middle-Late Devonian. However a few genera
reach the "Strunian" and the Tournaisian (Biumbella, Braznikovia, Eoumbella)
and Protoumbella is the youngest known representative in Visean time.
Anchicodiaceae do not appear before the Moscovian, where they suddenly
become important mound-builders.
The most precise time index is Calcifolium which spans the latest Visean-
earliest Namurian and is endemic to the Western Tethys.
Most of the Dasycladales are long ranging, but some are restricted to a stage.
They are certainly not as precise as their Mesozoic counterparts and are too
facies sensitive to be used in routine correlation. However, a number of
characteristic floras are recognizable, some endemic to a continental block.
The Tournaisian of the American Cordillera is characterized by Colum-
biapora. The Visean-Early Namurian in the Western Tethys has Cabrieropo-
ra-laphetella-Kulikia; in the Eastern Tethys, Queenslandella; in North Amer-
ica, Dutroella, Sandoella, Spinctoporella, Windsoporella; Koninckopora is
known from all the continental blocks.
The Bashkirian Dasycladaceae are not diagnostic and the stage is recog-
nizable on the red alga Masloviporidium and the first abundance of Bereselleae
(Beresella, Dvinella).
The Moscovian witnesses the first occurrence of Clavaporella, Connexia,
Epimastopora, Herakella, Sinarella, Zaporella. They are associated with
Eugonophyllum and Ivanovia.
The latest Carboniferous and Early Permian flora are difficult to separate.
Palaeosiphonocladales are declining although Beresella is still common.
Epimastopora-Epimastoporella are abundant associated with Pseudogyro-
porella-Gyroporella.
While many widespread groups have identical stratigraphic distributions
from one basin to another, there are a few cases of strong heterochrony among
Fig. 2. Stratigraphic distribution of microftora. I Albertaporella; 2 Amorfta; 3 Anatolipora; 4

A nchicodium; 5 A nthracoporella; 6 Anthracoporellopsis; 7 Aoujgalia; 8 Aphralysia; 9 Archaeolith-
ophyllum; 10 Asphaltina; II Asphaltinella; 12 Atractyliopsis; I3 Beresella; 14 Berestovia; 15
Bevocastria; 16 Biumbella; 17 Borisovella; 18 Brazhnikovia; 19 Cabrieropora; 20 Calci[olium; 21
Calcisphaera; 22 Clavaporella; 23 Coelosporella; 24 Columbiapora; 25 Connexia; 26 Cribro-
kamaena; 27 Cribrosphaeroides; 28 Cummingsella; 29 Cuneiphycus; 30 Cylindro[olia; 31 Dasy-
cladophycus; 32 Dokutchaevskella; 33 Donezella; 34 Dutroella; 35 Dvinella; 36 Einorella; 37 Elenia;
38 Eokoninckopora; 39 Eolithoporella; 40 Eoumbella; 41 Eouraloporella; 42 Eovelebitella; 43
Epimastopora; 44 Epimastoporella; 45 Epiphyton; 46 Epistacheoides; 47 Eugonophyllum; 48 Evlania;
49 Exvotarisella; 50 Fasciella; 51 Foliophycus; 52 Fourstonella; 53 Frustulata; 54 Garwoodella; 55
Garwoodia; 56 Girvanella; 57 Gissarella; 58 Gouldina; 59 Gyroporella; 60 Halenopora; 61 Hed-
stroemia; 62 Herakella; 63 Iberiaella; 64 Intextulella; 65 Issinella; 66 Ivanovia; 67 Japhetella; 68
Kamaena; 69 Kamaenella (Illustration see page 438)
438 B. Marnet
/'VVV'VVVV DIACHRONIC
X CARBONATE PRODUCER
JURASSIC
•
ABUNDANT
"REEF" BUilDER
COMMON I-
TRIASSIC
------ SCARCE ~ STRATIGRAPHIC MARKER
12 15 22 29
M. + U. ".1
PERMIAN 4 5 9 13 ".I
ASSELIAN
GHZELIAN
+
ORENBURG IAN
21
I I
I I
I
I I
MOSCOVIAN I
I I
I
I I I
I I I
10 II
2 I 22 25
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I 8
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E. NAMURIAN
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I 13 14 I I
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+ I 19 20 I I
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t
9 26 I I
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I 3 6 30
TOURNAISIAN 12 23
24
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DEVONIAN 16 18
E. + M. II
DEVONIAN
10 , 21 27
SILURIAN
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15
Fig. 2. (see caption page 437)

56
59
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33 35 46 1
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74 81 89 92 101
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ASSELIAN I
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E. NAMURIAN I -t. I
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+ I
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82 83 88 I
75 I
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DEVONIAN I
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SILURIAN
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91 lOl
ORDOVICIAN
89
CAMBRIAN
Fig.3. (see caption page 442)

119
121
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102 I
110 130 137
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?
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121 133
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114 119
442 B. Mamet
Fig. 3. Stratigraphic distribution of microftora. 70 Komia; 71 Koninckopora; 72 Kulikia; 73

Lithostroma; 74 Macroporella; 75 Malakhovella; 76 Mametella; 77 Masloviporella; 78 Maslovipo·
ridium; 79 Mellporella; 80 Menselina; 81 Microcodium; 82 Mitcheldeania; 83 Nanopora; 84
Nansenella; 85 Neoanchicodium; 86 Nosticites; 87 Orthriosiphon; 88 Orthriosiphonoides; 89 Or-
tonella; 90 Palaeoberesella; 91 Palaeomicrocodium; 92 Parachaetetes; 93 Paraepimastopora; 94
Parakamaena; 95 Parastacheia; 96 Pekiskopora; 97 Peristacheia; 98 Petschoria; 99 Proninella; 100
Protoumbella; 101 Pseudochaetetes; 102 Pseudogyroporella; 103 Pseudohedstroemia; 104
Pseudoissinella; 105 Pseudokamaena; 106 Pseudokomia; 107 Pseudonanopora; 108 Pseudosolen-
opora: 109 Pseudostacheoides; 110 Pseudovelebitella; III Quasiumbella; 112 Queenslandella; 113
Rectangulina; 114 Renalcis; 115 Retephycus; 116 Richella; 117 Sandoella; 118 Sinarella; 119
Solenopora; 120 Sphaerinvia; 121 Sphaerocodium; 122 Sphaeroporella; 123 Sphinctoporella; 124
Stacheia; 125 Stacheoidella: 126 Swcheoides: 127 Sycidium; 128 Taldykites; 129 Tubiphytes; 130
Ungdarella; 131 Unjaella; 132 Uraloporella; 133 Vermiporella; 134 Wetheredella; 135 Windsopo-
rella: 136 Yukonella; 137 Zaporella; 138 Zidella
possible udoteaceans and stacheiins when one compares the Tethyan and North
American blocks with the Arctic.
All known Orthriosiphon-Orthriosiphonoides are Lower Carboniferous
with the exception of the Arctic where they are Middle Carboniferous and even
reach the Lower Permian. The same heterochronism is observed for stacheiins
such as Stacheia. Epistacheoides. and Stacheoidella-Stacheoides which are
"normally" Visean-Namurian but are Moscovian to Early Permian in the
Arctic. This distribution suggests that the Tethyan, and to a lesser degree
American, seas were the breeding ground from which the microflora repeatedly
invaded the Arctic block during periods of warming.
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Chapter 20
Permian Marine Calcareous Algae
R. RIDING and L. Guo 1
Abstract
The Permian is characterized by development of distinctive gymnocodiacean non·reeffloras and of

the problematic reefal genera Archaeolithoporella and Tubiphytes. Lower Permian calcareous algae
are dominated by dasycladaleans and phylloid algae. The importance ofphylloids declines towards
the middle Permian, as does that of problematic groups common in the Carboniferous such as
Stacheia, Ungdarella and similar genera. Calcified cyanobacteria and genera traditionally referred
to the Solenoporaceae are, overall, of minor importance during the period. Key locations for the
study of Permian algal floras in New Mexico and Texas, the Eastern Alps and Yugosla via, the central
Urals, the Caucasus, northern Iran and Afghanistan, south China, and Japan, reflect their originally
broadly equatorial distribution.
1 Introduction
An aura of mystery surrounds much that we know concerning Permian calca-

reous algae. There is a residue of the Carboniferous flora in the Early Permian,
where phylloid algae are prominent in reefformation, continuing a role already
well-established in the Pennsylvanian. This is not to deny that phylloid algae
present problems of identity (some are probably green algae, others red), but
their decline during the Lower Permian coincides with the rise in importance of
the even more intriguing reef building genera Archaeolithoporella (red alga?)
and Tubiphytes (cyanobacterium?, hydrozoan?, sponge?), together with the very
successful, and also enigmatic, Gymnocodiaceae. The latter is the single most
distinctive group of Permian algae, and it combines morphological features of
both rhodophytes and chlorophytes. Gymnocodiaceans, together with dasy-
cladaleans, are common constituents of Permian shallow marine non-reefal
limestones.
Nevertheless, it is in the reefal facies, particularly in the middle Permian
when Archaeolithoporella and Tubiphytes were most common, that problems
are concentrated. Uncertainties concerning affinity make it difficult to determine
whether these reefs are dominated by algae or sponges. Diagenetic problems
make it difficult to say whether the reef fabric is altered skeleton or cement.
These doubts concerning reef com position and structure are well observed in the
history of research on the Capitan Reef Complex of New Mexico and west
Texas, but there is a suite of other less famous reefs, particularly in the
1 Department of Geology. University of Wales College of Cardiff. Cardiff CF I 3 YEo UK

Permian Marine Calcareous Algae 453
Mediterranean region, which further emphasizes the problems centred on

middle Permian reefs. Only in the very latest Permian reefs, such as those in
southern China, do Archaeolithoporella and Tubiphytes become subordinate to
calcisponges, and a transition towards Triassic floras occurs. Latest Upper
Permian (Palaeo[usulina zone) reefs also occur in Skyros, Greece, where at least
two reef types are present, characterized by the predominance of Ar-
chaeolithoporella and Tubiphytes, and by the predominance of caIcisponges,
respectively (Flugel, pers. comm. 1989).
Fundamental research on Permian calcareous algae is summarized by Pia
(1937, pp. 813-837), Johnson (1963), Babcock (1979), and Fliigel (1984). The
principal features which are beginning to emerge are:
1. Decline during the Permian of Carboniferous groups, notably phylloid
algae and algal microproblematica such as Stacheia Brady, Ungdarella
Maslov, etc;
2. Middle Permian importance ofdasycladaleans, but their decline in the Late
Permian;
3. Stepwise progressive increase in the importance of gymnocodiceans during
the Permian;
4. Importance, especially in the middle Permian, of the reef-associated
problematica Archaeolithoporella Endo and Tubiphytes Maslov;
5. Broadly equatorial distribution, from North America to the Tethyan belt,
although a northward trend in east Asia (particularly Japan) is anomalous
and may indicate error in current continental reconstructions;
6. Two principal environmental associations: (a) reefal, dominated by
phylloid algae, Tubiphytes and A rchaeolithoporella, and (b) shallow normal
marine non-reefal, dominated by Gymnocodiaceae and Dasycladales.
In addition, the evaporitic Zechstein deposits of north-west Europe contain a
distinctive stromatolitic facies, although this is not dealt with here.
2 Algal Facies
2.1 Shelves
Fragments of dasycladaleans and gymnocodiaceans are conspicuous in both

inner and outer platform environments throughout the Permian. In the Lower
Permian (Asselian and Sakmarian) of the Carnic Alps, for example, Girvanella
and Efluegelia occur in lagoonal environments whereas the inner platform is
characterized by the dasycladaleans A nthracoporella Pia, Epimastopora Pia and
Mizzia Schubert, plus the problematic Ungdarella, with a more diverse dasy-
cladalean-dominated outer platform association (p. 573 and Fig. 2 in Flugel
1979). A similar distribution can be seen in rocks of Artinskian age in central
Afghanistan where the dasycladalean A tractyliopsis Pia emend. Accordi occurs
near the seaward margin of oncoid shoal deposits in association with the
gymnocodiacean Dzhul[anella Korde and the phylloid red alga Archaeolitho-
454 R. Riding and L. Guo
phyllum Johnson, while the cyanobacterium Zonotrichites Bornemann occurs in

more restricted environments shorewards of oolitic banks (Fig. 10 of Vachard
and Montenat 1981). In the Upper Permian gymnocodiaceans generally
become more abundant than dasycladaleans in both inner and outer platform
environments (Elliott 1968).
2.2 Reefs
Wolfcampian (Early Permian) reefs, such as those of the southwestern United

States which are characterized by calcareous sponges, phylloid algae, and
bryozoans bound by Archaeolithophyllum,Archaeolithopore/la, and Tubiphytes,
may have their origins in late Pennsylvanian "thicket communities" on the
margins of phylloid algal banks (Wahlman 1986). Nevertheless, phylloid algae,
particularly Eugonophyllum Konishi and Wray and Archaeolithophyllum,
continue as dominant organisms in reef formation in the Early Permian.
Phylloid algal mounds with thicknesses of approximately 5-10 m form exten-
sive complexes up to more than 120 m thick and several kilometres across in
shelf environments in New Mexico and west Texas (Cys 1985). By Sakmarian
and Artinskian time phylloids had declined and the Tubiphytes-Archaeolitho-
porella association dominated reefs such as those at Trogkofel in the Carnic Alps
of southern Austria [FIUgel1966 (with Archaeolithoporella not named directly,
but referred to as a stromatolite), FIUgel 1981a]. However, the Guadalupian
Capitan Reef Complex of New Mexico and Texas shows a more complicated
zonation, with Tubiphytes Maslov, A rchaeolithopore/la Endo and phylloid algae
all involved in reef construction, although phylloids remain generally
subordinate.
In the late Permian reefs of Sichuan and Hubei, China, phylloid algae are
not noticeable and even Tubiphytes and Archaeolithoporella are subordinate to
sponges, "hydrozoans" (which, as in the Triassic, may also include sponges,
FIUgel 1981 b, p. 307), tabulozoans and bryozoans.
3 Algal Groups
Despite the intricate problems of affinity in Permian calcareous algae, we

attempt here to outline a preliminary classification based on existing infor-
mation (Table I). This provides a guide to uncertainties which need to be
clarified by research, as well as to the emerging facts concerning calcareous algae
at the end of the Palaeozoic.
3.1 Rhodophyta
Gymnocodiaceae, "ancestral corallines", genera attributed to the Solenopo-

raceae, and Archaeolithoporella may all be placed tentatively in the Rhodo-
Table 1. Classification of Permian marine calcareous algae and cyanobacteria. listing principal
generaa
PROKAR YOTES
I Cyanobacteria Garwoodia Wood 1941 ". Girvanella Nicholson and Etheridge 1878.
Hedstroemia Rothpletz 1913. Ortonella Garwood 1914. Zonotrichites
Bornemann 1887.
ALGAE
I Rhodophyta a) Gymnocodiaceae. Gymnocodium Pia 1920. Nanjinophycus Mu and

Riding 1983, Permocalculus Elliott 1955 (= Dzhulfanella Korde 1965).
Succodium Konishi 1954", Tauridium GUvenc 1966".
b) "Ancestral corallines". Archaeolilhophyllum Johnson 19561'", Cunei-
phycus Johnson 1960 (probably not an alga), Efiugelia Vachard 1979.
Lysvaella Chuvashov 1971.
c) "Solenoporaceans". Parachaeleles Deninger 1906. Pycnoporidium
Yabe and Toyama 19281". "Solenopora" Dybowski 1878b
2 Phylloid algae'H ' Anchicodium Johnson 1946, Eugonophyllum Konishi and Wray 1961.
Hikorocodium Endo 1951 (probably not an alga). Ivanovia Khvorova
1946.
3 Chlorophyta a) Dasycladales
A Iberlaporella Johnson 1966
Amicus Maslov 1956
Anatolipora Konishi 1966
A nthracoporella Pia 1920
Archaeocladus Endo 1956
A trac/yliopsis Pia 1937
Clavaphysoporella Endo 1958
Clavaporella Kochansky and Herak 1960
Clypeina Michelin 1845
Diplopora Schafhautl 1863
Dvinella Khvorova 1949
Endoina Korde 1965
Eogoniolina Endo 1953
Eovelebitella Vachard 1974
Epimastopora Pia 1922 (=
Globuliferoporella Chuvashov 1974)
Epimastoporella Roux 1979
Goniolinopsis Milanovic 1965
Gyroporella GUmbel 1871
Imperiella Elliott 1975
lohnsonia Korde 1965
Kochanskiella Milanovic 1965
Likanella Milanovic 1966
Macroporella Pia 1912
Mizzia Schubert 1907
Nanjinoporella Mu and Elliott 1982
Nipponophysoporella Endo 1959
Oligoporella Pia 1912
Paraepimas/opora Roux 1979
Permopora Elias 1947
Permoperplexella Elliott 1968
Physoporella Steinmann emend Endo
Pseudopimastopora Endo 1960
Table 1. (Continued)
ALGAE
Pseudogyroporella Endo 1959

Pseudovermiporella Elliott 1958
Pseudovelebitella Marnet and Roux 1978
Salopekiella Milanovic 1965
Shigaporella Endo 1961
Sinoporella Yabe 1949
Teutloporella Pia 1912
Thailandoporella Endo 1969 (= Gyroporella GUmbel)
Triploporella Steinmann 1880
Velebitella Kochansky-Devide 1964
Velomorpha Horiguchi 1961
Vermiporella Stolley 1893
Xainzanella Mu 1984
Zaporella Racz
b) Halirnedaceae. Aphroditicodium Elliott, 1970, Neoanchicodium Endo
1954, Orthriosiphonoides Petryk 1972, Siamporidium Endo 1969
4 Problernatica a) Archaeolithoporella Endo 1959"
b) Caicispheres'"
c) Collenella Johnson 1942pb
d) Receptaculitids p,
e) Stacheia Brady 1876 group'"
f) Tubiph ytes Maslov 1956"b
g) Ungdarella Maslov 1956 and Komia Korde 1951 group'"
a A, Probably algal; H, may belong with the Halimedaceae; Ph. phylloid; Pb; probably notalgal; Ps,
possibly not algal; RC, probably include rhodophytes and chlorophytes; Archaeolithophyllum is
also phylloid.
b Genuine Solenopora is restricted to the Ordovician.
phyta, but this necessitates careful discussion because very few of these can be
regarded with certainty as red algae.
3.1.1 Gymnocodiaceae
This extinct group includes representatives that resemble the weakly calcified
Recent chaetangiacean red alga Galaxaura. The family Gymnocodiaceae was
erected by Elliott (1955) to include Gymnocodium Pia emend. Elliott (Fig. 1) and
Permocalculus Elliott (Fig. 2). Mu and Riding (1983) referred Nanjinophycus to
the Gymnocodiaceae. Succodium Konishi and Dzhul[anella Korde (see Korde
1965), as well as Tauridium Guvenc may also belong in the family (but also see
Halimedaceae, below). Dzhul[anella is probably a junior synonym of Per-
mocalculus (Xinan Mu, pers. comm. 1989).
These are cylindrical or sack-like fossils, typically occurring as fragments a
few millimetres in size. with a filamentous internal medulla and a relatively thin
cortex with pores. The plants may have been segmented and sporangia may be
present. This latter detail is important for the affinities of the group: vegetatively
Fig. 1. Gymnocodium. Changxing Formation . Uppermost Permian, Sichuan, China . x50

Fig. 2. Permocalculus. Changxing Formation. Uppermost Permian, Sichuan, China. x20
the Gymnocodiaceae somewhat resemble halimedacean green algae, but they

can also be compared with chaetangiacean red algae such as modern Galaxaura
and, in addition, some gymnocodiaceans possess reproductive conceptacles
similar to those of Galaxaura (see Mu and Riding 1983, pp. 270-274). At present
the most likely affinity of the Gymnocodiaceae is with the red algae, which is the
view arrived at by Pia (1937, pp. 830-831).
Gymnocodium and Permocalculus first appear in the Lower Permian.
Gymnocodium is restricted to the Permian with the exception of a possible
Eocene occurrence (Massieux 1966, pp. 119-121). Permocalculus is known to
range to the Cretaceous, and has also recently been reported from the Lower
Miocene of Papua New Guinea (M.D. Simmons, pers. comm. 1989). Nanjino-
phycus is, so far, only known from the Permian of China. Gymnocodiaceans are
particularly abundant in the Upper Permian (Fig. 22), and Gymnocodium and
Permocalculus are geographically widespread; occurring in North and Central
America, southern Europe, Tunisia, Turkey, the Middle East, Pakistan,
Thailand, Vietnam, China, and Japan.
Gymnocodiaceans typically occur as sand and gravel-size fragments in
ramp and outer shelf environments, associated with dasycladaleans and
Pseudovermiporella.
3.1.2 "Ancestral Corallines"
The origins of the coralline red algae are obscure. Several Upper Palaeozoic
genera have been suggested to be ancestors of the Mesozoic Corallinaceae and
Wray (1977, pp. 71-74) termed these ancestral corallines. In the Permian these
include A rchaeolithophyllum, Cuneiphycus Johnson, and Lysvaella Chuvashov.
Archaeolithophyllum and Lysvaella are similar in internal structure, and possess
hypothallial-perithalial tissue differentiation and conceptacle-like bodies, both
of which suggest relationship to extant calcareous red algae.
Archaeolithophyllum has a thin crustose thallus similar to that of
squamariacean (Peyssonneliaceae) red algae and to some phylloid algae (see
below), whereas Lysvaella is cylindrical in form. Cuneiphycus has large cells
arranged in anastomosing sheets and does not appear to have differentiated
tissue. Its affinities are more problematic.
Lysvaella occurs in the Lower Permian of the Urals (Chuvashov 1971) and
Cuneiphycus, best known from the Pennsylvanian, is present in the Lower
Permian of the Carnic Alps (FIGgel 1977, p. 320). Archaeolithophyllum is an
important component oflimestones, including reefs, in the Pennsylvanian and
Permian of the central and south-western USA (OUe and Parks 1963; Wray
1964; Heckel and Cocke 1969; Mazzullo and Cys 1979; Toomey and Cys 1979).
James et al. (1988, pp. 300-301) tentatively compare Archaeolithophyllum with
sq ua mariaceans (peyssonneliaceans).
3.1.3 Solenoporacean Genera
Massive skeletons composed of narrow closely packed cellular filaments which

do not show tissue differentiation or conceptacles have traditionally been
assigned to the Solenoporaceae. Genera grouped into this family have been
reported to be geographically widespread in the Permian, although they do not
appear to be very abundant.
Johnson (1942) reported Solenopora Dybowski from the Capitan Reef
Complex and, later, S. centurionis and S. texana from the nearby Apache
Mountains (Johnson 1951). Johnson (1951) and Rigby (1957) considered S.
texana to occur as an intergrowth with the bryozoan Acanthocladia
guadalupiensis. However, the bryozoan-algal consortium hypothesis in general,
wh ich originated with Condra and Elias (1944), was criticized by Duncan (1957,
p. 789) who, according to Bain (1967, p. 213) maintained that "Acanthocladia
represents bryozoan growth entirely". In line with this, Babcock (1974) sug-
gested that S. texana, rather than being algal, may actually represent the outer
surface of Acanthocladia, an observation supported by Reinhardt (1988, p. 256).
Babcock (1977, p. 19) noted that Parachaetetes Deninger "the only other
solenoporacean red alga in the Capitan" is restricted to the Upper Capitan-
massive, i.e. the shallower part of the reef complex. Solenopora and/or Para-
chaetetes have also been recorded from the Southern Alps (Fig. 18) (FIGgelI979,
p. 575), southern Tunisia (Driggs 1977, p. 44), Saudi Arabia (Rezak 1959), the
eastern slope ofthe Ural Mountains (Chuvashov 1969), south China (Mu 1981),
and Japan (Konishi 1954; Endo 1959). Vachard 1980, p. 381) records Pseudo-
chaetetes from the Upper Artinskian of Afghanistan. Fossils in which cross-
partitions are often absent and which have been referred to Solenopora occur
abundantly in the Zechstein limestones of the Holy Cross Mountains, Poland
(Belka 1979).
Pycnoporidium Yabe and Toyama has been described from the Upper
Permian of Japan (Endo 1956, 1961). Despite uncertainty concerning the
affinity of Mesozoic examples of Pycnoporidium, Johnson (1963, p. 97) regarded
the Permian specimens as being solenoporaceans.
3.1.4 Ungdarella Maslov, Komia Korde
These problematic genera show broad similarities. They are rodlike, branched
fossils with a fibrous, apparently cellular, construction. They have commonly
been regarded as rhodophytes (Korde and Maslov 1963, pp. 258-259) and both
genera are present in the Permian. Komia is reported to range to the top
of the Asselian, and Ungdarella throughout the period (Fig. 39 in Roux 1985),
although it is most common in the Lower Permian.
3.1.5 Archaeolithoporella Endo
A rchaeolithoporella (Figs. 15, 16) is a light-dark laminated encrusting organism

which Endo (1959) originally described from the Permian of Japan. Babcock
(1977, p. 15) verified Endo's report that cross-partitions are absent and discussed
Archaeolithoporella in detail. The genus is, together with Tubiphytes, both
abundant in the Permian and enigmatic. It makes crustose masses up to several
millimetres thick in reefs in the Wolfcampian of Texas (Wahlman 1985), and
Lower Artinskian of the Alps (Flugel 1981 a), and also in Upper Permian reefs
in Tunisia (Boyd et al. 1975), Sicily (Flu gel 1984), south China (Fan et al. 1982),
the northern Caucasus (Pisera and Zawidzka 1981), and the Capitan Reef
Complex (Babcock 1977; Toomey and Cys 1977, pp. 136-138).
Although Endo (1959) failed to recognize any cross-partitions, the presence
of conceptacle-like bodies led him to place A rchaeolithoporella in the red algae.
Johnson (1963, p. 98) agreed and went further by asserting that Endo's pho-
tograph does show partitions across the layers. He placed A rchaeolithoporella in
the Solenoporaceae and compared it with "masses of poorly preserved Tertiary
Lithocodium". However, a cement origin has also been proposed for Ar-
chaeolithoporella (Dunham 1972). This was critically assessed by Babcock
(1977, pp.16-17) who concluded in favour of an organic origin for Ar-
chaeolithoporella. Babcock drew attention to the regularity of the laminae, the
palaeoecology of Archaeolithoporella, and the stratigraphic restriction of the
genus, but also emphasized the need for further studies. Subsequently (Babcock
1986, pp. 17-19), he restated the difficulties of interpreting Archaeolithoporella
but concluded that it is probably a red alga. On balance, at present it seems likely
thatArchaeolithoporella is organic and Johnson's (1963) broad comparison with
corallinacean crusts remains valid, if tantalizing.
However, a likely complication in elucidating the nature of Archaeolith-
opore lla is the fact tha t this name may have been applied too widely. A particular
problem in this respect concerns confusion between Archaeolithoporella and
stromatolites in earlier work on Permian reefs (see p. 52 and Fig. 9 in FlUgel
1966). The important binding role of algae was also noted, for example, by Bain
(1967, pp. 213-214, 226) in Leonardian (Artinskian-Kungurian) patch reefs of
the Glass Mountains, Texas. However, apart from Tubiphytes, Bain did not
identify the algae, which he described as "stromatolitic". These could be
non-skeletal stromatolites, but it is also possible that they may be crusts of
Archaeolithophyllum, other phylloids, or Archaeolithoporella, all of which are
common encrusters in mid-Permian reefs. Vachard (1980, p. 344, PI. 5, Fig. 2)
mentions "Archaeolithoporella stromatolites" in the Upper Permian of Afgha-
nistan. Laminated encrustations in the Zechstein of northern Europe resemble
Archaeolithoporella, but may not necessarily be the real thing (Smith 1981,
p. 178), although Peryt (1986) has suggested that Archaeolithoporella could be
regarded as a junior synonym of Stroma ria Walther (see also FlUgel 1989,
pp. 127-128). Caution is necessary!
4 Phylloid Algae
Pray and Wray (1963, p. 209) introduced the term phylloid algae to refer to a
variety of leaflike genera which are especially common in the Pennsylvanian
and Permian. Phylloid algae in the Permian include Anchicodium Johnson,
Eugonophyllum Konishi and Wray (Fig. 14), Ivanovia Khvorova, and Ar-
chaeolithophyllum Johnson. The group is, however, morphological rather than
biological and may include both red and green algae. However, the term
phylloid algae is useful descriptively because the internal structure of these
fossils is often lost during diagenesis, so that it can be impossible to distinguish
them in many cases.
Early studies led to the view that genera such as Anchicodium, Ivanovia and
Eugonophyllum were codiacean green algae (Konishi and Wray 1961; Johnson
1963, pp. 126-132). However, as work progressed, a rhodophyte affinity also had
to be considered. The phylloid alga Archaeolithophyllum has a cellular internal
structure which relates it to red algae (see "ancestral corallines", above).
Wray et ai. (1975) compared Recent squamariaceans with Permian material
and suggested that some phylloid algae are Squamariaceae. FlUgel (1977, Table
2) listed Anchicodium and Eugonophyllum as possible ancestral squama-
riaceans. But Eugonophyllum has internal, or partly internal, conceptacle-like
structures which suggest that it is not a squamariacean. Ivanovia has been
mentioned as the phylloid genus which compares best with Squamariaceae
(Wray pers. comm., in Riding 1977). However, it is also possible that some
Palaeozoic phylloid algae, apart from Archaeolithophyllum are halimedacean
green algae (see Johnson 1963) and James et al. (1988, p. 300) regard Eugono-
phyllum and Ivanovia as both belonging to this latter group.
Phylloid algae are best known from the southwestern United States.
Phylloid algal mounds are the dominant type of reef in the Wolfcampian ofN ew
Mexico and Texas, and mound complexes are major hydrocarbon reservoirs in
the region. Ivanovia, Eugonophyllum and Archaeolithophyllum are especially
important genera here (Toomey 1976; Toomey and Cys 1979; Cys 1985;
Wahlman 1985). Phylloids are also common until the upper Sakmarian in the
Canadian Arctic (Mamet et al. 1979) and there is a similar stratigraphic
distribution in the Southern Alps (FlUgel 1977). A nchicodium occurs in the
Lower Permian of Tibet (M u 1982) and Eugonophyllum has been recorded in the
Lower Permian of Guizhou, China (Zhang and Zhen 1985). In Japan A n-
chicodium occurs in the Lower Permian (Endo 1953).
Phylloid algae retain their importance from the Pennsylvanian into the
Early Permian, but they suffered rapid decline during the Sakmarian. Phylloid
genera continue to be present in the Trogkofel Limestone (Sakmarian and
Lower Artinskian) of the Southern Alps (Edwards, pers. comm. 1988) and are
locally important in the Capitan Reef Complex. In the latter, Babcock (1979)
reported Eugonophyllum and Archaeolithophyllum. But phylloids are relatively
rare in the Late Permian, although they are now known from the Norian (Late
Triassic) of the Yukon (Reid 1986).
5 Chlorophyta
5.1 Dasycladales
Permian dasycladaleans (erect stick-like algae with relatively thick walls pierced
by pores originally occupied by branches) are more diverse in the Lower
Permian than in the Upper Carboniferous (Fig. 40 in Roux 1985). At least 47
genera (Table 1), which belong to at least seven tribes, have been described for
the period as a whole. The most common genera are Diplopora, Eogoniolina,
Epimastopora (Fig. 5), Epimastoporella (Fig. 8), Gyroporella (Fig. 10), Mizzia
(Fig. 9), Macroporella (Fig. 4), Paraepimastopora (Fig. 6) and Velebitella.
Genera especially common in the Lower Permian include Anthracoporella (Fig.
7), A tractyliopsis (Fig. 3), and Globuli[eroporella. The latter genus is regarded as
a junior synonym of Epimastopora by Roux (1979), whose taxonomy is followed
here (see Figs. 5, 6 and 8). But for further details concerning Epimastopora also
see Endo (1960) and FIUgel and FlUgel-Kahler (1980, pp. 138-142). Bebout and
Coogan (1964) revised A nthracoporella using Wolfcampian material from
Texas. Pseudovermiporella (Figs. II, 12) is particularly common in the Upper
Permian. There is no current agreement concerning the affinity of this latter
genus, although Elliott (1968, p. 70) concluded that it is probably algal. Bere-
sell ids (recently shown by Skompski (1987) to be dasycladaleans) range from the
Carboniferous up to the Asselian, as do the still problematic donezellids (Fig. 39
in Roux 1985).
Fig.3. A Iracly iiopsis. Trogkofel Formation. Lower Permian, Trogkofel, Carnic Alps, Austria. x20
Fig.4. Macroporeiia. Trogkofel Formation. Lower Permian, Trogkofel, Carnic Alps. Austria. x20
Fig. 5. Epimaslopora. Trogkofel Formation. Lower Permian. Trogkofel, Carnic Alps, Austria. x20
Fig.6. Paraepimaslopora. Trogkofel Formation. Lower Permian , Trogkofel. Carnic Alps, Austria.
x20
Fig. 7. Anlhracopore/la. Trogk ofel Formation. Lower Permian, Trogkofel, Carnic Alps, Austria.
x7.2
Fig. 8. Epimaslopore/la. Upper Pseudoschwagerina Limestone. Lower Permian. Trogkofel, Carnic
Alps. Austria. x20
Fig. 9. Mizzia. Capitan Formation. Upper Permian, Guadalupe Mountains. New Mexico. x2l
Fig. 10. Gyroporella. Trogkofel Formation. Lower Permian, Trogkofel, Carnic Alps. x2l
Fig. 11. Pseudovermiporella. Changxing Formation. Uppermost Permian, Sichuan , China . x50
Fig. 12. Pseudovermiporella soda/iea Elliott. Changxing Formation. Uppermost Permian. Sichuan.
China. x63
Details of Permian dasycladalean associations are best known from Iraq

(Elliott 1968), Japan (Endo 1956, 1958, 1959), the Carnic Alps (Flugel 1966,
1977, 1985; Homann 1972; Flugel and Flugel-Kahler 1980), Afghanistan
(Vachard 1980, Vachard and Montenat 1981), Tibet (Mu 1982), southern China
(Mu and Elliott 1982, Zhang and Zhen 1985), Tunisia (Glintzboeckel and
Rabate 1964, Vachard 1985), and New Mexico and Texas (Johnson 1942, 1951).
Rezak (1959) reported a few genera from the Permian of Saudi Arabia.
Dasycladalean diversity during the period is greatest during the middle
Permian (Fig. 2 in Flugel 1985), but declines sharply in the latest Permian
(Xi-nan Mu, pers. comm. 1988) although individual genera continue to be
abundant.
5.2 Halimedaceae
Apart from those phylloid algae (see above) which may be halimedaceans, this
group of green algae, characterized by internal filamentous structure, is
represented in the Permian by Aphroditicodium Elliott and Orthriosiphonoides
Petryk. Neither is particularly abundant. Neoanchicodium Endo (Fig. 13) may
be a halimedacean (Xinan Mu, pers. comm. 1989). Two additional genera,
Tauridium Giivenc and the more common Succodium Konishi, could be
halimedaceans, but may also belong to the Gymnocodiaceae (see above).
Succodium has been reported from Japan (Konishi 1954a, Endo 1957), the
Caucasus (Korde 1965), the Southern Alps (Homann 1972), Thailand (Endo
1969), and Vietnam (Nguyen 1970). Korde (1965) first suggested that Succodium
could be a gymnocodiacean, and this has been followed byVachard (1977). Mu
and Riding (1983) placed S. endoi Nguyen in the gymnocodiacean genus
Nanjinophycus.
6 Tubiphytes Maslov
Tuhiphytes (Fig. 17) is commonly irregularly cylindrical or encrusting in form

and has a characteristic dark, flocculent, internal structure, generally in a series
of uneven bands. Internal tubes forming a central canal system are usually
present. It occurs as small (a few millimetres) individ ual masses and also as thick
Fig. 13. Neoanchicodium. Trogkofel Formation. Lower Permian, Trogkofel, Carnic Alps, Austria.
xl8
Fig.14. Eugonophyllum. Upper Pseudoschwagerina Limestone. Lower Permian, Trogkofel, Carnic
Alps, Austria. x6
Fig. 15. Archaeolithoporella. Changxing Formation. Uppermost Permian, Sichuan, China. x20
Fig. 16. Archaeolithoporella encrusting Tubiphytes. Capitan Formation. Upper Permian,
Guadalupe Mountains, New Mexico. x20
Fig. 17. Tubiphytes. Changxing Formation. Uppermost Permian, Sichuan China. x36
(up to several centimetres) encrustations on sponges and bryozoans. It ranges

from Carboniferous to Cretaceous. It is especially abundant in the Permian,
being a major element of reefs such as Capitan (Babcock 1977) and Trogkofel
(FIUgelI98Ia).
Maslov (1956) described Tubiphytes as a possible cyanobacterium
(Schizophyta) from the Kungurian of the central Urals. Previously, Rauser-
Chernousova (1950) had named this fossil Shamovella, but without providing a
description (see Maslov 1956, p. 21). Konishi (1959) recognized Nigriporella
Rigby as a junior synonym of Tubiphytes. Babcock (1977, pp 17-18) briefly
reviewed the affinities which had been proposed and concluded that neither a
hydrozoan nor a cyanobacterial affinity for Tubiphytes seems likely. He has
again (Babcock 1986, pp. 15-17) reviewed the genus in some detail. Ernst Ott (in
Kraus and Ott 1968, p. 271) has suggested that Tubiphytes may be a sponge. A
somewhat similar genus, Plexoramea Mello from the Triassic, has been com-
pared with both chlorophyte algae and fungi (FlUgel et al. 1988).
Tne question of the nature of Tubiphytes is made especially significant by
the abundance of this genus in Permian reefs. It is a major component in the
middle Permian and is also abundant in the Early and Late Permian. It now
seems that earlier views, such as that of Rigby (1958), comparing Tubiphytes
with hydrozoans are nearer the truth than suggestions of a cyanobacterial or
algal affinity. Indeed, an invertebrate affinity seems likely, and if Tubiphytes
should prove to belong to the calcisponges (the view favoured here) then this will
only reinforce the already recognized importance of this group in middle to
Upper Permian reefs generally.
7 Cyanobacteria
Girvanella Nicholson and Etheridge, Hedstroemia Rothpletz, and Ortonella

Garwood, together with some other non-septate genera attributed to the
Solenoporaceae, have been fairly widely reported from the Permian as a whole,
but they only occassionally appear to be locally important. FlUgel (1979, p. 573)
notes Girvanella as a characteristic element of restricted lagoonal environments
in the Lower Permian of the Carnic Alps.
Osagia Twenhofel and Ottonosia Twenhofel, oncoids which usually include
Girvanella together with other encrusting organisms, have been reported from
the Lower Permian (Twenhofel 1919; Johnson 1963, p. 134). Toomey et al.
(1988, 1989) demonstrate that Ottonosia and Osagia nodules from the Lower
Permian of Kansas and Oklahoma are dominated by "Girvanella, intimately
associated with various calcareous and agglutinated encrusting tubiform fo-
raminifers" (Toomey et al. 1989, p. 51), but note that Girvanella tends to be
diagenetically degraded to micrite. They point out that the principal difference
between Ottonosia and Osagia is one of size, with Ottonosia being larger and
containing larger attached epizoans including worm tubes and bryozoans.
Toomey et al. (1989, p. 61) propose that the formal names Ottonosia and Osagia
be abandoned.
Two Permian genera which have been attributed to the cyanobacteria,

Collenella Johnson and Tubiphytes, almost certainly do not belong in this group.
Collene/fa, which is important in the upper part of the Capitan Reef Complex
(Johnson 1942, 1963, pp. 136-137; Toomey and Cys 1977, p. 136) has, like
Tubiphytes (see Sect. 6 above) been compared with metazoans such as
stromatoporoids and bryozoans (see Babcock 1979, p. 424).
Hedstroemia, a cyanobacterium of rivulariacean type, is present quite
commonly with Collene/fa in the upper Capitan-massive (Babcock 1977, Fig.
3b). However, other cyanobacterial genera are more rare. Zonotrichites Bor-
nemann, a probable rivulariacean, is reported from the Lower Artinskian of
eastern Afghanistan by Vachard (1980, p. 335). Vachard (1980, pp. 344-345)
also reports Bacinella RadoiCic from the Upper Permian of Afghanistan. This is
probably the earliest record of this problematic meshwork-like fossil, which is
best known from the Triassic-Cretaceous, and has variously been referred to
chlorophytes and prokaryotes, as well as to invertebrates.
Renalcis Vologdin and Tharama Wray have been described from the Upper
Permian of Tunisia (Vachard and Razgallah 1988), but these reports require
clarification. One illustrated specimen (Vachard and Razgallah 1988, their Fig.
1) closely resembles Izhella Antropov, hitherto not known from rocks younger
than Lower Carboniferous.
The relative lack of importance of calcified cyanobacteria in the Permian is
in contrast to both the Lower Carboniferous and the Middle to Upper Triassic.
8 Receptaculitids
Receptaculitids are relatively large dish or vase-shaped fossils which have been
compared with green algae, as well as with sponges. The group is most
prominent in the middle Palaeozoic. Carboniferous and Permian examples are
rare (Nitecki 1972, p. 306). Parona (1933) reported them from the Permian of
Sicily and, if confirmed, this appears to be among the last known occurrences of
this problematic group.
9 Regional Distribution
Low sea-level stands characterize the Permian but, nevertheless, extensive

shallow marine sequences developed in some areas, notably western and Arctic
North America, the Ural region, and the margins of Tethys including south
China, and Japan (see Ross and Ross 1979). All these areas contain calcareous
algae (Fig. 19). In addition, stromatolites are common in the evaporitic Zech-
stein area of northern Europe. There are at least seven key areas for the study of
normal marine Permian calcareous algae worldwide, and these reflect a broadly
equatorial distribution for these sequences during both the Lower and the
Upper Permian (Figs. 20, 21).
Fig. 18. "Solenopora" . Trogkofel Formation . Lower Permian. Trogkofel. Carnic Alps. Austria. x6
Fig. 19
Fig.20. Distribution of principal localities with marine Lower Permian calcareous algae. Base map
from Smith et al. (1981)
...
Fig. 19. Main localities of described marine Permian calcareous algae, with principal references.
I Capitan reef complex, Guadalupe Mountains, Texas and New Mexico (Babcock 1977);
2 southern Tatum Basin, southeastern New Mexico (Cys 1985); 3 northern Sacramento Mountains,
south-central New Mexico (Toomey and Cys 1979); 4 Motley County, Texas (Wahlman 1985);
5 Apache Mountains, Texas (Johnson 1951); 6 Northern Midland Basin, Texas (Toomey 1985);
7 Kansas (Johnson 1946); 8 Gautemala (Johnson and Kaska 1965); 9 and 10 Sverdrup Basin,
Canada (Davies et al. 1986); 11 Northern Franklin district, Canada (Mamet et al. 1979); 12 Sexten
Dolomites, Italy (FliigeI1977, Fig. I); 13 Carnic Alps (FliigeI1977, Fig. I); 14 Sava Folds, Slovenia.
Yugoslavia (FliigeI1977, Fig.I); 15 Velebit Mountains, Croatia, Yugoslavia (Fig. Iin FliigeI1977);
16 Biikk Mountains, Hungary(Herakand Kochansky 1963); 17 Greece (Pia 1937); 18 Sicily(Fliigel
and Stanley 1984); 19 Djebel Tebaga, Tunisia (Elliott 1958; Emberger 1958; Glintzboeckel and
Rabate 1964; Vachard 1985); 20 Anatolia, Turkey (Bilgiitay 1959); 21 Labariver Basin, northern
Caucasus (Korde 1965; 1967; Pisera and Zawidzka 1981); 22 Mosul Liwa, northern Iraq (Elliott
1968, p. 85) ; 23 Isfa han, Iran ; 24 Dammam, Saudi Arabia (Rezak 1959); 25 Oman (Elliott 1958);
26 Cherdyn, western Urals (Chuvashov 1983); 27 Sterlitamak, western Urals (Chuvashov 1983);
28 Bashkiria, USSR (Kulik 1978); 29 Salt Range, Pakistan (Rao and Varma 1953); 30 Pondu, Tibet
(Mu 1982); 31 Kara mulun, Tibet (Mu 1982); 32 Lago, Markam, Toba, Tibet (Mu 1982); 33 Khao
Ph long Phrab, Thailand (Endo 1969); 34 Vietnam, Cambodia, Laos (Nguyen 1970); 35 Guizhou ,
southern China (Mu 1981); 36 Sichuan, southern China (Guo and Riding 1988); 37 Nanjing, China
(Mu and Elliott 1982 ; Mu and Riding 1983); 38 Kyushu , Japan (Konishi 1954); 39 Hiroshima,
Japan (Endo 1957); 40 Gifu-ken, Japan (Endo 1954); 41 Hokka ido, Japan (Endo 1957)
i ~j,.,
~
~ ....s'
Fig.21. Distribution of principal localities with marine Upper Permian calcareous algae. Base map
after Smith et al. (1981)
9.1 New Mexico-Texas
Shallow carbonate shelves surrounding deep-water embayments in what is now

the southwestern United States were sites of reefal sedimentation at several
times during the Lowerand Middle Permian (Fig. 19, localities 1-6; Figs. 20, 21),
and phylloid algae, Tubiphytes and Archaeolithoporella are important additions
to sponges in these reef environments. Phylloid algal mounds are well-
developed in the Wolfcampian (Toomey and Cys 1979; Cys 1985). These Early
Permian mounds contrast, in both morphology and algal flora , with the Upper
Permian Guadalupian Capitan Reef Complex. The latter contains locally
abundant Archaeolithoporella, Eugonophyllium, and Tubiphytes, distributed
patchily mainly in the upper 50-60 m of the Capitan massive limestone
(Babcock 1977, 1979). In addition, Babcock notes Parachaetetes and Hed-
stroemia in the uppermost parts of the reef, and the dasycladaleans Macroporella
and Mizzia in the bedded back-reef facies. An important feature of the algal
distribution at Capitan is the vertical succession, from bottom to top: (I)
Archaeolithoporella and Tubiphytes , (2) phylloid algae, (3) Parachaetetes and
H edstroemia.
9.2 Eastern Alps and Yugoslavia (Fig. 19, Localities 12-15)
The Sakmarian to Artinskian Trogkofel Limestone of southern Austria and

northern Italy is slightly younger in age than the Wolfcampian bioherms of the
southwestern USA. Structural complications and patchy dolomitization ob-
scure the morphology and setting of the Trogkofel reef limestones, but they
contain abundant Archaeolithoporella and Tubiphytes in addition to Ar-
chaeolithophyllum, bryozoans, calcisponges, cyanobacteria, dasycladaleans and
other algae (Flugel 1966, 1981a; Homann 1972; Flilgel and Flugel-Kahler
1980). Edwards and Riding (1988) have suggested that sparite fabrics inter-
preted as marine cement at Trogkofel (and at Capitan) may be diagenetically
altered algal skeletons. Calcareous algae are also present nearby in the Sexten
Dolomites of northern Italy (Ogilvie-Gordon 1927; Pia 1937; Accordi 1956;
Praturlon 1963). Here, and in adjacent parts of northern Yugoslavia, the
Bellerophon Formation of Upper Permian age contains a non-reefal shelf flora
dominated by Gymnocodiaceae and Dasycladales, with gymnocodiaceans
more important in the more restricted inner shelf facies (Noe 1987). The
Permian-Triassic boundary is located in the overlying Tesero Horizon and is
marked by disappearance ofthe Permian algae and foraminifers due to climatic
change (Noe 1987, p. 115).
The Permian calcareous flora, particularly the gymnocodiaceans and da-
sycladaleans, of Yugoslavia are known mainly through the work of Pia (1937),
Herak and Kochansky (1960), and Kochansky and Herak (1960).
9.3 Central Ural Mountains
The Lower Permian of the central western Urals contains reefal facies with
calcareous algae (Chuvashov 1983) (Fig. 19, localities 26, 27). In the Asselian
and Sakmarian dasycladaleans (Anthracoporella, Epimastopora, Glob u life-
roporella, Gyroporella), phylloid algae (Eugonophyllum), and the possible
halimedacean Neoanchicodium are common, and Tubiphytes is very abundant.
However, floral diversity decreased during the Artinskian and Kungurian as
evaporitic and non-marine conditions developed.
9.4 Caucasus
The Caucasus area (Fig. 19, locality 21) is comparable with both the Urals and
with more thoroughly marine Tethyan areas. Korde (1965, 1967) described
Guadalupian and Dzhulfian (partly equivalent to Kazanian, see Harland et al.
1982, p.23) algae, which are dominated by dasycladaleans and gym-
nocodiaceans. These probably are from non-reef facies. Marine conditions
persisted longer than in the Urals, but nevertheless from the late Guadalupian
onwards there was reduced algal diversity, and by the mid-Dzhulfian no
calcareous algae were present in the area.
9.5 Northern Iraq and Afghanistan
Shelflimestones of Permian age in northern Iraq (Fig. 19, locality 22) have been
studied by Elliott (1968), who reported abundant Mizzia, Gymnocodium and
Permocalculus throughout the 800 m thick Permian sequence. Dasycladaleans
less common than Mizzia include Anthracoporella, Atractyliopsis, Permoplex-
ella, Pseudoepimastopora, Pseudovermiporella (possibly not a dasycladalean),
plus very scarce Clypeina. Solenopora, in contrast to the dasycladaleans and
gymnocodiaceans, is uncommon (Elliott 1968, p. 93).
Vachard (1980) has described in detail Artinskian and Upper Permian
calcareous algae from east-central Afghanistan. Common genera include
Tubiphytes, numerous dasycladaleans (particularly Mizzia, Clavaporella,
A tractyliopsis) , Orthriosiphonoides, Archaeolithophyllum, and the gymnoco-
diacean Permocalculus (also reported as its junior synonym Dzhul[anella).
9.6 South China
The Permian of Tibet (Fig. 19, localities 30-32) and south China (Fig. 19,
localities 35-37) contains shallow marine limestones at many levels throughout
the sequence, which is relatively complete. Calcareous algae have been described
from the Lower Permian of southern and eastern Tibet (Mu 1982, 1984) and the
Lower Permian ofNanjing (Mu and Elliott 1982; Mu and Riding 1983). But the
best known flora is from the Lower and Upper Permian of southern Guizhou
(Kweichow) in southwest China (Mu 1981; Zhang and Zhen 1985). In Tibet,
Nanjing, and Guizhou the sequences are essentially in non-reefal platform
facies and the flora, as is that of Iraq, is dominated by Dasycladales and
Gymnocodiaceae. An interesting feature of the Chinese Permian is the presence
in eastern Sichuan and western Hubei of very late Permian (Changxing Stage)
reefs. These contain abundant Archaeolithoporella and Tubiphytes, in addition
to calcisponges and "hydrozoans", and are associated with dasycladaleans,
gymnocodiaceans, and Pseudovermiporella in the off-reeffacies (Fan et a1. 1982;
Reinhardt 1988; Guo and Riding 1988).
9.7 Japan (Fig. 19, Localities 38-41)
In numerous papers. mainly published between 1951 and 1961, Riuji Endo
described calcareous algae from the Permian of Japan (see for example Endo
1959 and references therein and in Johnson 1963). Most of the algae are
dasyc\adaleans and they appear to be in bedded limestones associated with
fusuline foraminifers. Endo erected A rchaeolithoporella. Neoanchicodium.
Hikorocodium (probably not an alga), and the dasyc\adaleans Archaeocladus,
Clavaphysoporella. Eogoniolina. Nipponophysoporella. Pseudogvroporella. and
Shigaporella. In addition he reported occurrences of Epimastopora. Girvanella.
Gymnocodium, Macroporella, Oligoporella, Physoporella, Solenopora, Teut-

loporella, and Vermiporella (actually Pseudovermiporella).
Kenji Konishi also contributed significantly to description of Japanese
Permian algae (see references in Johnson 1963), including erection of Suc-
codium from the late Permian (Konishi 1954).
10 Conclusions
The most abundant and widespread fossils referred to the calcareous cyano-
bacteria and algae in the Permian are Archaeolithoporella and Tubiphytes
(although it is likely that the latter is neither a cyanobacterium nor an alga),
gymnocodiaceans, dasycladaleans, and phylloid algae (Table 1). In addition,
the red alga A rchaeolith ophyllum (also phylloid in form), and the problematic
genera Cuneiphycus, Stacheia and Ungdarella are present. Calcified cyano-
bacteria such as Girvanella, Hedstroemia and Ortonella occur, although rarely
abundantly, throughout the period. Beresellids and donezellids which, like
stacheiids and members of the Ungdarella group, are common in the Upper
Carboniferous, have also been reported to persist into the Asselian (Fig. 39 in
Roux 1985). Genera referred to the Solenoporaceae occur, but are not common,
and receptaculitids make a final, very rare, appearance.
Although detailed information is still lacking from many areas, it is clear
that there are marked changes in the importance of major groups of calcareous
algae during the period (Fig. 22). The Early Permian is dominated by the
persistence of Carboniferous groups, notably phylloid algae and problematical
algae (Roux 1985, p. 670). Decline of these groups is counterbalanced from the
Artinskian onwards by the rapid, stepwise expansion of the Gymnocodiaceae,
which is dominant in non-reef shelf limestones in the Upper Permian, and by
dramatic increase in importance in the reef-associated problematica Ar-
chaeolithoporella and Tubiphytes. Dasycladaleans are also common until at least
the middle Permian, and probably longer. The result of these changes is a
marked contrast between Early and Late Permian algal floras, with those ofthe
middle Permian being intermediate in composition. These developments are
seen just as clearly in non-reefal as in reefal facies. Thus, (l) Early Permian reefs
are dominated by phylloid algae with dasycladaleans in bedded shelf lime-
stones; (2) Trogkofel, Capitan, and other reefs of the middle Permian are
characterized by Archaeolithoporella and Tubiphytes, with subordinate phyl-
loids; while (3) Late Permian reefs in China have fewer of these organisms and
more sponges, with gymnocodiaceans abundant in associated bedded
limestones.
Permian calcareous algae are best known from reef environments. The
Capitan Reef Complex, ofGuadalupian age, is a good example both of vertical
and lateral variations in algal distribution, and of problems of ecological and
diagenetic interpretation of the algae. In the same area of the southwestern
United States, Wolfcampian phylloid algal mounds are well developed. Per-
-J
CHANGXINGIAN ""'"
.-----'TATAR IAN ""'"
lie
w IRAQ
~ IKAZANIAN
a. JAPAN
~ 0
~
UFIMIAN CAPITAN ( ~
..••
>-
~
KUNGURIAN 0 Go
III
• :a
~
lie ARTINSKIAN ~ ...
w ~ )1K~'Y1
~ TROGKOFEL
0 SAKMARIAN
..J NEW MEXICO/
ASSELIAN S> TEXAS
REEF OFF-REEF )
Fig. 22. Abundance estimates for major groups of marine calcareous algae and possible algae, plus sponges, tabulozoans and "hydrozoans".
Stratigraphic positions and ranges of areas with notable reef, and off-reef sequences are shown on the left and right respectively. The Urals region contains
both reef and off-reeffacies. Time scale based on Harland et al. (1982, chart 2.7)
1'"
is:
'5·"
(JQ
.,
::s
p..
r
Cl
c::
o
mian reefs are also seen in the Tethyan belt. In the Mediterranean region there
are notable examples in Austria, Greece, Italy, Tunisia and Yugoslavia (Fliigel
and Stanley 1984) of which possibly the best known are in the Artinskian ofthe
Carnic Alps (e.g. Trogkofel) close to the Austrian-Italian-Yugoslav border. In
China, Late Permian (Changxing) reefs (e.g. in Hubei and Sichuan) are im-
portant for understanding developments immediately prior to the end of the
period. More needs to be known concerning the algal floras of Lower Permian
reefs in Arctic regions, such as Canada, Svalbard and the Barents Sea, and the
central Ural Mountains.
Shallow normal marine non-reefal limestones with calcareous algae, no-
tably dasycladaleans and gymnocodiaceans, are best known in northern Iraq,
the Caucasus Mountains, southern China and Japan. The lithofacies of these
deposits are, however, less well documented than those of the reeflimestones.
The Zechstein stromatolitic deposits of northern Europe represent a quite
different facies, related to developments also seen in the Urals, in which
evaporitic conditions progressively eliminated normal marine algae, as well as
invertebrates.
Permian calcareous algae present contrasts, as well as some similarities,
with those of both the Carboniferous and Triassic. Several groups which are
important in the Pennsylvanian decline rapidly in the Lower Permian while
groups, such as calcified cyanobacteria, which are common in the Triassic are
not conspicuous during much of the Permian. Gymnocodiaceae, and the
problematic genera Archaeolithoporella and Tubiphytes, dominate the period,
and it is the uncertainties concerning the affinities of these taxa which are
responsible for much of the confusion, as well as not a little of the interest,
characterizing the study of Permian calcareous algae.
Acknowledgements. Li Guo's work has been supported by the South Western Petroleum College in
Nanchong, Sichuan, China, and TOTAL, Paris. We are grateful for Andre Maurin's support and
encouragement. David Edwards very kindly provided us with examples of algae from Trogkofel in
the Carnic Alps. We are particularly indebted to Xi-nan Mu for tilling in gaps in our knowledge.
Comments on the manuscript by Jack Babcock, Erik Flugel, and Xi-nan Mu have been most helpful
and we appreciate their willingness to share their expertise with us.
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Chapter 21
Triassic and Jurassic Marine Calcareous Algae:
A Critical Review
Abstract
The information on benthic Triassic and Jurassic marine algae is scattered in about 1600 articles,
most of which have been published during the past two decades. The Triassic algal floras yield
porostromate "blue-green algae". udoteacean and dasycladacean green algae as well as soleno-
poracean and gymnocodiacean red algae. These groups also occur in the Jurassic, except for the
gymnocodiaceans; in addition the first corallinaceans are recorded from the Upper Jurassic. About
50 genera and 200 species have been described from the Triassic. and about 60 genera and 130
species from the Jurassic.
Lower Triassic algae (except spongiostromates) are lacking on a global scale. The extinction
rate near the Permian/Triassic boundary is 56% for dasycladacean gepera and 53% for all algal
genera. Because of an absence of originations and high extinction rates at the end of the Middle
Permian, Upper Permian algal floras are characterized by low-diverse assemblages consisting
predominantly of gymnocodiaceans and some dasycladaceans. Most dasycladaceans became
extinct prior to gymnocodiaceans (as demonstrated by Late Permian floras from the Southern Alps
and China), probably because gymnocodiaceans were better adapted to fluctuating salinity con-
ditions during periods of regression.
The first Triassic skeletal algae are recorded from the early Middle Anisian of the western
Tethys. Reef associations composed of encrusting spongiostromates, porostromates and soleno-
poraceans can be compared with Upper Permian algal associations described from "algal/cement
reefs". Anisian lagoonal and platform environments are characterized by high-diverse dasy-
cladacean floras which indicate a rapid post-Permian diversification. Diversity decreases from the
Anisian to the Upper Triassic. Taxonomic turnover is high during Middle Triassic and Late Triassic
but low at the Ladinian/Carnian boundary. Decreasing diversity is also evident within the four
dasycladacean assemblages characteristic for Anisian to Rhaetian time intervals.
Liassic and early Middle Jurassic dasycladacean floras appear to be low-diverse subsequent to
the Hettangiian-Sinemurian, but the information about Upper Liassic and Middle Jurassic algae is
scanty. A significant increase in diversity during the Upper Jurassic reflects diversification during the
Kimmeridgian and Portlandian (Tithonian).
Most Triassic and Jurassic time intervals are characterized by high origination rates at species
level. Extinction rates are high during the Triassic and Liassic, low during the Middle Jurassic and
increase significantly during the Upper Jurassic. Majorextinction events, noted for animal biota, can
be recognized also on generic and partly on specific level for the dasycladacean algae.
Dasycladaceans and udoteaceans exhibit a Tethyan, predominantly pan-tropical dispersal
pattern since the Middle Triassic. Isocrymal distributional patterns have been proved for Jurassic
algae but might have also existed in the Triassic (indicated by Middle Triassic non-Tethyan
dasycladacean floras and perhaps also by the first Triassic dasycladaceans found in western North
America).
Utilization of Triassic and Jurassic algae in biostratigraphy is restricted to dasycladaceans and
strongly affected by environmentally controlled distribution patterns. Algal assemblages, rather
than the time-range of species, enable the recognition of time intervals which might correspond to
several ammonite or conodont zones in the Triassic. Jurassic platform and platform-edge car-
1 Institut flir Palaontologie, Universitat Erlangen. LoewenichstraBe 28. 8520 Erlangen. FRG
482 E. FlUgel
bonates can be subdivided into facies-controlled time units ("coenownes") which only partly might
be regarded as synchronous within a global context. The stratigraphical range of Upper Jurassic
dasycladaceans is relatively short for more than 50% of the species, especially during the Kim-
meridgian and Portlandian.
The study of Triassic and Jurassic algae offers new insights into algae/sediment interactions,
such as accumulation of bioclastic carbonate sands by halimediform green algae, evolution of
spongiostromate and porostromate oncoids, together with the as yet not well understood
significance of microbial crusts for the formation of carbonate buildups.
1 Introduction
This paper provides a summary of the current state of research in the field of
marine Triassic and Jurassic calcareous algae, together with an assessment of
future avenues of investigation. Not included are non-benthic as well as
fresh-water algae, and the controversial information about evolutionary trends,
e.g. of dasycladaceans (see Kamptner 1958; Herak et al. 1977). The review starts
with a discussion ofthe extent and significance ofthe data base (number oftaxa,
taxonomic problems), followed by a discussion of taxonomic diversity which
might be used to elucidate extinction and origination rates of Triassic and
Jurassic algae. Despite the continued expansion of our knowledge on sys-
tematics and morphology there are still obstacles with regard to the relations
between Late Permian and Triassic algae. The development of Upper Permian
algal floras, therefore, is also considered. Speculations about the causes of
changes in the composition of algal floras during time are only peripherally
discussed. The groups dealt with here comprise "blue-green algae" (or cyano-
bacteria) including spongiostromate and porostromate taxa (which might
include green algae as well, see Roux 1985); green algae (udoteaceans, dasy-
cladaceans) and red algae (solenoporaceans, gymnocodiaceans, corallina-
ceans). Mesozoic phylloid algae (squamariaceans?) are only known from a
unique occurrence in the Late Triassic of Northwestern Canada (Reid 1983).
The information on Triassic and Jurassic calcareous algae is scattered
through about 1600 papers, published between 1820 and today. Sixty percent of
the articles on Triassic algae and about 80% of the information about Jurassic
(and Cretaceous) algae have been published during the last three decades. This
situation is comparable with the strong increase in the knowledge of many plant
protist groups (Tappan 1980). The increase in the number of "new" species
generally depends on the efforts of only a few scientists as indicated by the
distinct increase in the number of Triassic algal taxa in the years between 1910
and 1940, predominantly due to the fundamental work of Julius von Pia, and by
the peak between 1960 and 1980, caused by Jan Bystricky's excellent con-
tributions, and by the work of Milan Herak, Ernst Ott and new investigations of
Triassic reef carbonates (Fig. I). The striking expansion of our knowledge of
Jurassic and Cretaceous algae since 1960 (Fig. 2) originates from the enormous
work of the "Groupe fran~aise d'etude des algues fossiles" and from the
contributions by Harlan Johnson and Graham F. Elliott. The common use of
Jurassic and Triassic algae in microfacies studies and paleoecological analysis is
Triassic and Jurassic Marine Calcareous Algae: A Critical Review 483
35
30
188
~ . -54
Number of paper~
25 (Total sum 654)
20
New species ,
IS
(Total sum 198)"'"
"
"23
10
,
1840 18501860 1870 18801890 19101920 1930 1940 1950 1960 19701980
. 1900
Fig. 1. Interest in the study of Triassic algae expressed by the number of relevant papers and by the
number of species proposed within a decade. Source: Emberger (1979) and additions
another reason for the increase in algal literature. Excellent bibliographies are
available for Triassic algae (Emberger 1979), Jurassic and Cretaceous dasy-
cladaceans (Bassoullet et al. 1978) and udoteacean green algae (Bassoullet et al.
1983). Much of the following review is based on the data gathered in these
critical compilations.
2 Data Base
The number of genera and species proposed for Triassic and Jurassic algae is
tabulated in Table 1. A taxonomic census is always difficult because of the
different state of the taxonomic concepts used and also because ofthe possibility
that important papers have been overlooked. Table 1 is based on probably more
than 90% of the relevant literature and , therefore , should receive a high degree
of confidence with regard to the completeness ofthe data. Problematical or poor
taxonomic concepts, however, will strongly influence the conclusive results of
the census. Taxonomy and classification of Mesozoic solenoporacean red algae
and especially of porostromate algae are highly artificial. Porostromate
classification depends on the possibility of recognizing inter- and intraspecific
morphological variability (Dragastan 1985), of differentiating various fos-
484 E. Fliigel
40
284 ..a9
35
30
25
20
15
New spec ies

10 (Total sum 235)
18101820 1830 1840 1850 186018701860 1890 191019201930 194019501960 1970 1980
1900
Fig. 2. Interest in the study of Jurassic and Cretaceousdasycladacean algae as shown by the number
ofrelevant papers and by the number of new species proposed per decade. Source: Bassoullet et al.
(1978) and additions
Table l. Number of genera and species attributed to the major algal

groups occurring in the Triassic and Jurassic"
Triassic Jurassic
Genera 12 8
Porostromata Species 36 24
Genera 4 4
Udoteaceae Species 4 7
Genera 25 44
Dasycladaceae Species 136 82
Genera 5 4
Solonoporaceae Species 19 12
Genera 3
Gymnocodiaceae Species 3
Genera 2
Corallinaceae Species 2
aThe table is based on probably more than 90% of the relevant literature.
Because of taxonomic problems the numbers of the porostromate algae
are less confident than the data of the other algal groups.
silization patterns (Leinfelder 1985), and of finding Recent counterparts

(Schafer and Senowbari-Daryan 1983). Difficulties arising from these obstacles
as well as from varying calcification patterns may even influence the taxonomy
of fossil algae having Recent descendants such as dasycladacean and udo-
teacean green algae (Bassoullet et al. 1975, 1983, 1984; Deloffre 1988). The
species concept offossil green algae is still a matter of discussion. A biometrical
approach, as exemplified for some Triassic and Jurassic dasycladaceans (Hurka
1967; Remane 1969; Zorn 1974, 1977; Bystricky 1984), might assist us to
overcome some of these difficulties.
There is considerable discussion concerning whether counting and
tabulating palaeontological taxa will provide sufficient information for recog-
nizing origination and diversification phases (see Valentine 1985), because of
the discrepancies between true biological species and "species" created on
minor morphological differences or even on different preservation or on-
togenetic stages of one and the same biological species. A census of Triassic and
Jurassic dasycladacean species, therefore, can only reflect very rough trends of
the overall morphological variability within the whole group. Because a modern
revision of Triassic and Jurassic dasycladacean algae is lacking, the recognition
of these trends (e.g. times of pronounced extinctions) is certainly based on a
mixture of well-defined morphological species and questionable "species"
which cannot be used for taxonomic comparisons because of insufficient
definitions. The elimination of those species from the census, however, does not
strictly influence the results shown in Figs. 5 and 6, probably because most of
these questionable "species" have been described from the Middle Triassic and
Upper Jurassic substages which, even after the subtraction of these taxa (about
20% of the total number of species, both in the Middle Triassic and the Upper
Jurassic), represent high-diverse time units.
Most Phanerozoic diversity patterns have been described in terms of
generic diversity or higher taxa diversity. Doing this with dasycladacean algae
would result in a distinctly different pattern for diversity and origination as
compared with the pattern indicated by species number. The substages of the
Middle and Upper Triassic are characterized by corresponding numbers of
genera, the number of new dasycladacean genera per substage remaining more
or less constant. The number of genera increases during the Lower Jurassic,
seems to be lower in the Middle Jurassic, and has a distinct peak in the Upper
Jurassic (31 genera!). The origination rates are low during the Triassic (about 20
to 30%) and Liassic (31%) but high for the Middle Jurassic and Upper Jurassic
(more than 80%). One reason for the difference between generic and specific
diversity is the more intensive study of Upper Jurassic dasycladacean algae as
compared with Triassic algae. The existence of time-dependent species groups
(associations, Ott 1972b), characterized by morphologically related species,
points to a rather broad generic concept applied to Triassic dasycladaceans.
The distribution of the major algal groups in time is shown in Fig. 3. All
Triassic algal groups already existed in the Late Paleozoic. Jurassic groups
continue into the Cretaceous. Dasycladaceans exhibit a high taxonomic coin-
cidence of uppermost Jurassic and lowermost Cretaceous species; about 44% of
the species known from the Tithonian continue to the Berriasian. A similar
486 E. FlUgei
CARBON-
IFEROUS
PERMIAN Triassic Jurassic RETACEOUS
L U L M U L M U L M U L U
CYANOPHYTA
"SPONGIOSTROMATA"
POROSTROMATA
GIRVANELLID ALGAE
FILAMENTOUS ALGAE
CHLOROPHYCOPHYTA
UDOTEACEAE ~
DASYCLADACEAE
BERESSELID ALGAE
PHYLLOID ALGAE
~
RHODOPHYCOPHYTA
UNGOARELLACEAE
SOLENOPORACEAE
ANCESTRAL CORALLINES
CORALLINACEAE
GYMNOCODIACEAE ~
Fig. 3. Stratigraphical range of algal groups occurring in the Triassic and Jurassic. Note the gap in
the paleontological record in the Lower Triassic, only non-skeletal algae are known from this
interval. Phylloid algae are represented by a unique occurrence only
pattern can be recognized within the porostromate algae, if one accept the
present classification of this group. Of special interest are the gaps in the record
of the gymnocodiacean red algae; rare Triassic species have been described
recently (H.W. Fluge11971; Senowbari-Daryan and Schafer 1980; Schafer and
Senowbari-Daryan 1983).
2.1 Relations Between Permian and Triassic Algae
A logical outcome of the data summarized in Fig. 3 is the extent to which Late
Permian potential precursors of Triassic algae have been affected by the biotic
crisis near the Permian/Triassic boundary. Recent discussions of the Phane-
rozoic diversity patterns do not consider calcareous algae (Valentine 1985). For
invertebrates and protozoans a Late Permian extinction rate of83% on generic
level and on 94 to 96% on specific level is estimated (Sepkoski 1986). The
extinction rates for calcareous algae seem to be different.
A survey of Late Permian algal floras from the southern Alps, Trans-
caucasia, Central Afghanistan, and China (Fig. 4) indicates a distinct change in
diversity and taxonomic composition during time (Table 2). Middle Permian
dasycladacean associations are characterized by high generic and specific
diversity. Twenty-four genera are known from the Middle Permian Neosch-
wagerina zone, but only 13 genera from the Upper Permian. About 58% of the
Middle Permian genera became extinct prior to the Upper Permian. Nearly all
Late Permian dasycladacean genera are already known from the Middle
Permian. There is a significant lack oftaxonomic originations. Algal limestones
from the Late Permian Palaeofusulina zone yield 9 genera and about 14 species
of dasycladaceans. At least 5 genera should have crossed the Permian/Triassic
boundary (Diplopora, Epimastopora, Gyroporella, Macroporella, Physoporella)
because they occur in Anisian limestones again. These recurring taxa suffered
a distinct reduction in species diversity and population size during the late
Permian. This is indicated by the development of algal floras within Late
Permian sections in the Southern Alps (Bellerophon and Tesero Formations,
Noe 1987) and in China (Wuchiapingian and Changhsiangian; Mu 1981; Rui
et al. 1984; Reinhardt 1987): (1) both in the southern Alps and in China algal
diversity decreases near the top of the sections; (2) dasycladaceans (with the
exception of Mizzia) disappear prior to gymnocodiacean red algae which
predominate in most Late Permian algal floras; (3) the algal floras are generally
low-diversity; (4) Gymnocodium bellerophontis, Mizzia and Atractyliopsis are
euryhaline taxa (Noe 1987) in contrast to Permocalculus tene/lus, M acroporella,
Vermiporella and Solenopora which seem to be adapted to normal or only
slightly changing salinities.
The distribution of calcareous algae at the end of the Permian period,
therefore, might have been strongly controlled by salinity fluctuations which, in
the Southern Alps and probably also in Guizhou caused a significant reduction
of dasycladacean diversity. Salinity control is evident for Uppermost Permian
algae of Sichuan (Reinhardt 1987).
A strong salinity control of dasycladacean distribution might also be
inferred from a comparison of algal associations described from Middle to
Upper Permian sections in the Transcaucasian region (Korde 1965): from the
Gnishinskian to the Kashikian and to the Dzhulfian (see Fig. 4) species diversity
of algae decreases from 25 to 11 to 5, no algae are known from the Dorashamian
of this region. Dasycladaceans disappear prior to gymnocodiaceans which are
common in the Gnishikian part of the section (about 54% of the algal species)
but rare in the Kashikian (27%) and Dzhulfian (20%). Low-diversity algal
assemblages occurring in alternation with higher diverse floras, as well as the
abundance of gymnocodiaceans within the low-diversity assemblages, might be
explained by salinity changes.
The extinction rate at the Permian/Triassic boundary is 56% with regard to
dasycladacean genera, and about 53% considering all algal genera known from
the uppermost Permian. Algae living in reef environments seem to have been
influenced by extinction events less than lagoonal and platform associations.
Encrusting algae (Archaeolithoporella, Tubiphytes) are effective in building
"algal/cement reefs" during the Late Permian (Pisera and Zawidzka 1981;
Peryt 1986) as well as during the Triassic (Flu gel et al. 1984; Fluge11989; Martin
and Braga 1987).
The problem of survivorship of Permian algae and of the post-Permian
diversification, indicated by the extremely high species diversity of Anisian
algae, becomes even more complicated if possible phylogenetic lines are taken
into consideration. Kochansky-Devide and Gusic (1971) have recognized that
.I>-
00
00
Neoschwagerina Lepidolina/ Codonofusiella Palaeofusulina
Zone Yabeina Zone Zone Zone
q A bad e h t
Abadeh.
s u r m a Ham bas Iran
Gnishilc Khachik Dzhulfa Dorasham Trans-
I
_____ _ _iF ~- ~=:~ I",omi'
Mu r 9 a b • Cap i t a n • Darvas
1_
- - - - - - - - - - - --0
Maolcuan
Wo"hi" i0 9 Ch'09· i0 9
Chl.na
Maolcuan
III
~
Lungtan Changsing
I·
-·0
-·0
--
CD
? Belleropon Tesero Southern
!1Intra- Alps
? Lower? Middle I Upper llili.te ~
? -(})
?
-0
- -0-----::- !'T1
'Tl
----@ 2i
~
I
Table 2. Stratigraphical range of Middle Permian and Upper Permian dasycladacean algal generaa
Middle Permian Upper Permian

N eoschwagerina Lepidolina/ Codonofosiella Palaeofusu-
Zone Yabeina Zone Zone lina Zone
Anthracoporella x
Atractyliopsis x x
Clavaphysoporella x
Cia va porella X
Diplopora X X
Endoina X
Eoclypeina X X
Eogoniolina X
Eovelebitella X X
Epimastopora X X X
Gyroporella X X X
Imperiella X
Johnsonia X
Kochanskiella X
Macroporella X X X
Mizzia X X X X
Pseudoepimastopora X X
Pseudogyroporella X
Pseudovermiporella X X
Salopekiella X
U ragiellopsis X
Velebitella X X
Vermiporella X X
Physoporella X x
Zaporella X
aNote the difference in taxonomic diversity between Middle and Upper Permian algae. At least five
genera known from the uppermost Permian (Palaeofusulina zone) re-appear in the Anisian .
...
Fig. 4. Distribution of Middle and Upper Permian algal floras used in the discussion of the relations
between Permian and Triassic algae. Sources: 1 Korde (1965); 2 Pisera and Zawidzka (1981),
northern Caucasus; 3 Vachard and Montenat (1981), central Afghanistan; 4 Mu (1981); southern
Guiszou, China; 5 Zhao et ai. (1981), Shangxi, China: 6 Not! (1987), Southern Alps; 7 Kochansky
(1954), Montenegro; 8 Herak and Kochansky (1963), Bukk Mountains, Hungary; 9 Termier et ai.
(1977), Djebel Tebaga, Tunisia; 10 lohnson and Danner (1966), Marble Canyon Formation, North
America
490 E. FlUgel
most morphological innovations characteristic of Triassic morphotypes evolved

during the Late Palaeozoic and not in the Triassic. This includes also the
development of metaspondylity in dasycladaceans (Giiven<; 1979).
Lower Triassic algae are lacking on a global scale, with the exception of
non-skeletal spongiostromate algae which seem to be common in lowermost
Triassic environments (Bissell 1970; Taraz et a1. 1981; Sheng et a1. 1984; Noe
1987). Boring algae responsible for the formation of coated grains in many
Scythian calcarenites are also common (e.g. Yun et a1. 1983). Spongiostromate
algae might have been even effective in the buildup of Scythian reef structures
(Nazarevya et a1. 1986).
There is no clear explanation for this gap in the paleontological record of
calcareous algae in the Triassic. The causes of this worldwide phenomenon have
been discussed in terms of rapidly changing transgression/regression patterns,
salinity crisis, and reductions in the size of shallow-marine shelf environments.
A change in skeletal mineralogy or in the degree of calcification (as discussed
for some invertebrate groups; Mistiaen 1984; Cuif et a1. 1987) at the Per-
mian/Triassic boundary might be another explanation. Interestingly enough,
the first algal associations of Triassic age consist only of algae which used
calcite or Mg-calcite. Aragonitic dasycladacean green algae seem to appear
within the Anisian after a short time lag. A change in the calcification potential
due to non-tropical settings of Lower Triassic algal biotopes would perhaps also
explain the lack in the palaeontological record.
2.2 Diversity Patterns of Triassic and Jurassic Algae
Summary of the species diversity of Triassic and Jurassic dasycladacean algae

(Fig. 5) is based on a total of 253 species, 138 of which are Triassic, and 115
Jurassic in age. "Subspecies", described especially from the Middle Triassic, are
counted as species. In addition, the number of species occurring in consecutive
time units is indicated. Several conclusions concerning evolutionary trends (as
well as the different state of knowledge) can be derived from Fig. 5:
(1) Dasycladacean diversity is highest at the beginning of the Mesozoic,

reflecting a strong post-Permian, probably Anisian, taxonomic diversification
proceeding from only a few Permian survivors; (2) diversity decreases from the
Anisian to the Upper Triassic, although four distinct algal floras can be dis-
tinguished in this time (Ott 1972b) which do not coincide with a change in
sedimentary facies and might, therefore, represent evolutionary innovation
which declined towards the end of Triassic: the Anisian Physoporella-
Oligoporella assemblage comprises the largest number of species, followed by
the uppermost Anisian and Ladinian Diplopora annulata- Teutloporella nodosa
assemblage. Even lower is the diversity of the Carnian Poikiloporella duplica-
ta-Cl)peina besici assemblage and of the Norian and Rhaetian Diplopora
phanerospora-Heteroporella assemblage; (3) the number of species continuing
to subsequent time units is high in the Middle Triassic but low in the Upper
54
"1
Triassic Jurassic
"~
40
41
36
35
V)
UJ 30
U
UJ 27
a..
V)
u.. 25
0
a:
UJ
'"2
::;,
20
20
15
10-
2 2 2 2 2 2 22 2 2 2 2 2 2 2 2 2
« ~ ~ ~ ~ ~ ~« ~
w
~ ~ ~ « ~ ~ ~
I en
~ 2 a: I- <!Ja: I U 2 u 2 > <!J C
a: C
I-
>
2
« C
«
a:
«
0
2
w
« 2::;,
«2
u
«a> « w
--'
0
:;: 0
I
0
--'
a:
0
ea: 2
«
u u I I-W 0 « u..
a: I-
«--'
...J
V) --' I- « a>
«a> w
~~
V)
I-
2 u
X
0 2 a:
IV) w 2 0
...J :.:: a..
a..
Fig.5. Diversity of Triassic and Jurassic dasycladacean species. Hatched areas indicate the number
of taxonomic turnovers. The figure reftectschanging diversities but also gaps in our information (e.g.
Lower and Middle Jurassic)
Triassic, except for the Norian-Rhaetian interval; (4) Liassic dasycladacean

floras are characterized by a strong decrease in species diversity after the
Hettanginian-Sinemurian. This pattern as well as the apparent low diversity of
early Middle Jurassic floras should not be overrated because of the very poor
information available for these time intervals. The very first dasycladaceans
from the Toarcian have just been described (Cubaynes and Deloffre 1984);
(5) Bathonian to Portlandian dasycladaceans are studied in much more detail.
The increase in species diversity, recognizable for the Upper Jurassic, should,
therefore, reflect new diversification phases during Kimmeridgian and Port-
landian. This is also indicated by the low taxonomic turnover from the Ox-
fordian to the Kimmeridgian.
492 E. Fliigel
Diversity patterns of other algal groups are difficult to determine because of

classification problems (porostromate algae) and because of the poor record
(red algae). Mesozoic udoteacean green algae appear in the Upper Triassic
(FlUgel 1987, 1989b) but do not become highly diverse until the Lower Cre-
taceous (see Fig. 7). Both Boueina and Arabicodium are minor components of
Upper Triassic and Jurassic algal assemblages.
2.3 Extinction and Origination Rates
Most Triassic and Jurassic time intervals exhibit high origination rates (Fig. 6).
Low origination rates are only known for Rhaetian and Pliensbachian dasy-
cladacean species, but the latter may be an artifact. The Bajocian and the
Callovian origination rates are about 50%, all other origination rates are
considerably higher.
The extinction rates are high in most parts of the Triassic and Liassic, low
in the Middle Jurassic and increasing during the Upper Jurassic. Major generic
extinction events recognized from animal biota (Sepkoski 1986) are also
reflected by dasycladacean extinction rates: the Carnian and the Late Norian
peak as well as the Pliensbachian/Toarcian peak are represented. The Late
Tithonian peak (caused by the extinction of marine invertebrates as well as
dinoflagellate algae) is even more significant for dasycladaceans than for other
biota if the species level is considered. At generic level the Upper Jurassic
origination rate for dasycladaceans is extremely high (approximately 80%).
2.4 Tethyan Dispersal Patterns
The first Triassic algal assemblages, similar in taxonomic composition to some

Late Permian reef assemblages (see above), appear in the early Middle Anisian
of the Southern Alps. Dasycladacean assemblages became common during the
Anisian and flourished in the tropical settings of the western Tethyan region
(Southern Spain, Alps, Carpathian Mountains, Yugoslavia, Hungary, Bulgaria,
Romania, western Turkey) but some occurrences have also been reported
outside the Tethys north of the Tethyan belt (e.g. Poland; Gadzicki and
Kowalski 1974). The distribution of Ladinian algae corresponds more or less to
that of Anisian algal assemblages. Carnian dasycladacean algae exhibit a
somewhat extended distribution including occurrences in northern Tibet
(FlUgel and Mu 1982). Norian and Rhaetian algae known from the western
Tethyan region as well from Asia. Recent discoveries have proved the existence
of Upper Triassic dasycladaceans also in western Northern America (Fltigel et
al. 1989).
Elliott (1977, 1981, 1984) stressed the fact that aragonitic green algae
(dasycladaceans and udoteaceans) have proliferated since Triassic times within
the warm-water Tethyan sea, whereas subtropical tolerance within non-Teth-
yan settings has developed only for a minority of Triassic and Middle Jurassic
Triassi c Ju rass ic
100 :;l
0;'
~
(S '
90
_______ Origination '"0-::>
~ rate '-
80 "...
~'"
(S'
70
s:
::;
:;'
60 n'"
,__ Extinction ()
'"
::;
rate
50 '";;;o
>
riQ
40
'"'"
>
30 n
2·
(s '
E.
20 ::0
'"(ii'<
~
10
0
2 2 2 2 2 2 22 z 2 2 2 2 2 2 2 2
<{ <{ <{ <{ w <{ <{ <{
::! ;;:; ::! ::!::! ~ ::! ~ ::! ::!
x 2 2 a: >- Cla: ::t: u 2 u 2 :; 0 <.!l 0
>- 2 a: a w u a: w a a a a: 0 2
0 <{ <{ z::> <{ <{ ....J ....J
>- <{ <{ 2 <{:!: X 0 <{
U U ::t: a:I a <{ ~ >- ....J U. iX ....J
V> ....J >-w <{ w
a: >- <{ x >-
>-2
w _ 2:
'" '" « u :!:
w 0 a:
xv> '" ~ 0
....J C>-
~
"-
.j>.
Fig.6. Origination rates (percentage of species originating within a time interval) and extinction rates (percentages of species 'C
w
becoming extinct at boundaries between time units) of Triassic and Jurassic dasycladacean algae, The figure is based on a
total of 253 species, High Liassic extinction rates may be due to insufficient information
494 E. Fliigel
ev4WL~:ZZ;W~ Reef
Environments
FREQUENT ABUNDANT
~
Lagoonal and
fZl Platform
Environments
RARE UNKNOWN
FREQUENT ABUNDANT
Ha limed iform algal b i ocl~sts

within
the sand
fraction
- - - - - 50% - - -- - - - of lagoonal
carbonates
UPPER TRIASSIC LOWER CRETACEOUS MIOCENE RECENT

THAILAND FRANCE MEDITERRANEAN PACIFIC CARRIBEAN
Fig. 7. Environmental distribution and sedimentary significance of halimediform udoteacean

green algae. Variou s so urces (see Fliigel 1988)
taxa. There is a strong evidence for isocrymal distribution of dasycladacean

algae in Europe, north Africa and southwestern Asia , at least since Liassic times
(Elliott 1977). Water temperature and, probably. also substrate conditions
(FliigeJ 1985) are regarded as principal factors governing the regional dis-
tribution of early and middle Mesozoic green algae.
3 Biostratigraphical Importance
Occurrence and distribution of ancient calcareous algae is strongly facies-

controlled (Wray 1977). Utilization of Triassic and Jurassic algae in biostrati-
graphy is restricted to the Oasycladaceae. Pia had a very optimistic view
concerning the value of dasycladaceans as Triassic index fossils (see Pia 1943).
Although many of the algal samples studied by Pia lacked stratigraphical
control. he recognized the general successional patterns of the dasycladacean
assemblages fairly correctly. A better understanding of the influence of facies
changes, and the study of algae from geological sections in the Alps, Carpathians
and the Oinarides, led to some modifications concerning the time range of
species (Bystricky 1964, Herak 1965). Ott (1972a,b) proposed a revised algal
biochronology based on four dasyc1adacean assemblages occurring in vertical
successions and separated by two major floral changes (within the uppermost
Anisian and near the Ladinian/ Carnian boundary). The absolute time-range of
these units is rather long and embraces between two and about six ammonite
zones, and two to five conodont zones, established for the Triassic. The prac-
ticability of this zonation, especially in lagoonal and backreef carbonates poor
in macrofossils has been, however, confirmed by recent studies of Middle and
Upper Triassic platform and reef carbonates (Fois 1979; Fois and Jadou11983;
Dragastan 1981).
Many attempts have been made to subdivide Jurassic platform and plat-
form-edge carbonates by the occurrence and/ or dominance of particular cal-
careous algae (and benthic foraminifera). Starting with the fundamental papers
by Sartoni and Crescenti (1962), Praturlon (1969), Farinacci and RadoiCic
(1964), RadoiCic (1966), Crescenti (1971) and Ramalhao (1971), many studies of
the zonation of Jurassic and Cretaceous platform carbonates have been pub-
lished (e.g. Chiocchini et al. 1979; Peybernes 1979) most of which are critically
summarized by Jaffrezo (1980), who provided a biozonation for the Jurassic and
Lower Cretaceous of the Mesogean area. Because shallow-marine facies belts
are easily affected by sea level fluctuations, which might change the position of
the facies belts in time, the boundaries between the "coenozones" based on
calcareous algae can hardly be regarded as synchronous time lines.
The stratigraphical range of Jurassic dasycladacean species varies, as
exemplified by Upper Jurassic species; about 54% of the species seem to be
rather shortlived (restricted to one substage, e.g. to the Kimmeridgian), about
20% ofthe species are known ftom two substages. The remaining species appear
to be longlived, but this might be partly due to inconsistency of taxonomic
determinations (e.g. Salpingoporella annulata, see Bassoullet et al. 1978).
The biostratigraphical value of Triassic and Jurassic calcareous algae is
strongly restricted by the existence of environmentally controlled distributional
patterns known from many Middle and Upper Triassic platform and reef
carbonates (e.g. Ott 1967, 1972c; Flilgell979; Senowbari-Daryan and Schafer
1979) as well as from Jurassic carbonates (e.g. FliigelI979). These patterns are
defined by the occurrence of peculiar species within a distinct biotope, by the
restriction of algal assemblages to special facies or microfacies types, or by the
quantitative predominance of algal groups, adapted to changing environmental
conditions (e.g. in lagoonal environments). Various parts of lagoonal and
back-reef environments can be well distinguished, but a spatial subdivision of
reef and fore-reef environments is also possible (using porostromate, soleno-
poracean and various encrusting algae in addition to green algae). An in-
structive example has been described by Schafer and Senowbari-Daryan (1983)
from the Upper Triassic Pantokrator limestone of Hydra Island, Greece. It must
be stressed, however, that considerable changes in the settings of algal as-
sociations might have occurred during time: encrusting algae are common in
reef environments during the Triassic, but are found also in open-shelflagoonal
environments in the Jurassic. Tubiphytes occurs in organic reefs in turbulent and
496 E. FlUgel
fore slope positions during the Triassic, but in reefs and open platform setting in
the Upper Jurassic. In contrast, filamentous blue-green algae (or green algae)
seem to have been rather conservative with regard to their situation within
Triassic and Jurassic environments.
4 Sedimentological Implications
Some of the many interrelationships between the formation of carbonate

sediments and the activity of benthic calcareous algae are outlined below.
4.1 Green Algae and Bioclastic Sedimentation
Middle Triassic dasycladacean algae might have had a role in the rapid
accumulation of sand-sized carbonate sediments similar to that ofudoteacean
green algae today. Grainstones and packstones of Ladinian lagoonal calca-
renites in the Northern Alps may consist up to about 80% of dasycladacean
fragments, but the proportion oflimestones rich in dasycladaceans is often low
(20 to 30%) as compared with other limestone types occurring in the sections
(algal bindstones, bioclastic wackestones, mudstones). Upper Triassic (Norian
and Rhaetian) shallow-marine carbonates are poor in mass-accumulations of
dasycladaceans (generally not more than 15% of the rock volume). Liassic and
Upper Jurassic shelf carbonates may be richer in dasycladacean remains (e.g.
Liassic Palaeodasycladus limestone, up to 60%; Kimmeridgian and Tithonian
Clypeina limestones, up to 80%).
A review of the paleontological record of udoteacean algae, related or
identical to Halimeda, shows that these sedimentologically very important algae
appeared during the Upper Triassic (Thailand, Northern Alps), are scarce
during the Liassic and Middle Jurassic, and remained rare elements also within
Upper Jurassic algal assemblages (FltigeI1987). Diversity increased during the
Lower Cretaceous, both for Boueina and Arabicodium, and during the Tertiary
for Halimeda which today embraces about 30 species (Fig. 8). The palaeoen-
vironmental setting of Mesozoic halimediform algae is partly comparable with
that of recent Halimeda; lagoonal and reefal environments are known already
in the Upper Triassic. The frequency, however, was low in the Upper Triassic
and Jurassic as compared with other algal groups (porostromates, dasy-
cladaceans). The high amount of halimediform algal bioclasts recorded from
Upper Triassic occurrences (Fig. 7) should, therefore. not be overrated.
4.2 Oncoids
Oncoids with different microfabrics might reflect different paleoenvironments

(Peryt 1981): Porostromate oncoids (skeletal oncoids. Riding 1977; cyanoids
p.p .. Riding 1983) are believed to have been formed from Cambrian to Jurassic
~~/I2?ZZZZZZZZZ~~
Boueina
Leckhamptonella
~//////////Za
Ara bic od ium
Hal imeda
Fig. 8. Halimediform udoteacean green algae: Diversity and distribution in time. Height of the
ashlar representing Leckhamptonella corresponds to one species. Pictures of the morphotypes after
Elliott (1982)
times within marine, subtidal environments, generally with low-energy con-

ditions, probably in somewhat deeper waters. Spongiostromate oncoids (non-
skeletal oncoids, Riding 1977) occurred in lacustrine and, in association with
stromatolites, in transitional marginal marine environments. Starting approx-
imately during the Middle Jurassic, spongiostromate oncoids replaced poro-
stromate oncoids, and became the only oncoid type in marine settings, but
continued to flourish in lacustrine environments. During the Eocene spon-
giostromate marine oncoids were replaced by corallinacean rhodoids. Recent
investigations of Triassic and Jurassic oncoids (e.g. Dahanayake 1977, 1978;
Catalov 1983; Tichy 1983) seem to confirm this concept. Derivations might be
explained by redeposition of oncoids (Gasiewicz 1983) or by obliteration of the
primary microfabric by recrystallization (Peryt 1980).
4.3 Microbial Crusts
Micritic and peimicritic or pelsparitic "crusts", sometimes with spongy, peloidal

or fenestral microtextures, are common in Triassic and Jurassic reef carbonates.
498 E. Flugel
The origin of these crusts is generally attributed to the binding and cementing
effect of cyanobacteria, various "algae", or unknown microcommunities.
Of special interest are those microbial crusts which have been formed within
subtidal environments and occur in close association with sessile invertebrates
which may contribute to the formation of reefs. Several types can be distin-
guished in Triassic carbonates:
1. Micritic crusts, associated with encrusting and low-growing organisms (e.g.
Tubiphytes, calcisponges);
2. Micritic and pelmicritic crusts, associated with a high-diverse fauna con-
sisting offrame-building organisms (e.g. corals, calcisponges);
3. Thin encrusting micritic laminae, alternating with submarine carbonate
cements which may be quantitatively more important than the crusts.
Type 1 is common in Middle Triassic, especially Anisian reef carbonates (e.g.
Fois and Gaetani 1984). Type 2 seems to be restricted to Norian and Rhaetian
reefs where spongiostromate crusts are important secondary framebuilders; no
recent counterparts comparable with these crusts are known. The same seems to
be true for type 3, which is common during the Permian and Middle Triassic
(FlUgel 1989a). An as yet undeciphered interaction between microbiota and
unusually rapid synsedimentary cementation seems to be one of the main
factors responsible for the formation of "algallcement reefs".
Jurassic "cryptalgal crusts" (Dromart and Elmi 1986), "pellet crusts" (e.g.
Brachert 1986) or "algal crusts" are other biogenic structures, probably caused
by microbial activity. These crusts are common in the Upper Jurassic sponge
reeffacies (cf. Matyszkiewicz 1989). Indications for microbial activity get more
and more strong. Photoacoustic studies of Oxfordian crusts from northern
Franconia resulted in the recognition of anoxic bacteria which seem to have
been responsible for the bacterially induced precipitation of carbonate and for
a rapid submarine cementation (Krumbein, Oldenburg, pers. commun.).
Comparable results have been obtained from bacteriological studies of Recent
sea water and lagoonal carbonate mud (Morita 1980; Castanier et al. 1984).
5 Future Research
Despite increasing interest in Triassic and Jurassic algae during the last decades
there is a strong need for an intensification of the investigations in several fields
of research:
1. Data base: The information about the taxonomic inventory of Triassic and
Jurassic calcareous algae is far from complete. Even better known groups,
e.g. dasycladaceans, should yield a considerable number of as yet unde-
scribed taxa judging from the gaps becoming visible in connection with
higher taxonomic classifications (e.g. Bassoullet et al. 1979; Deloffre 1988).
2. Taxonomic revisions, as well as critical discussions, of classification con-
cepts are necessary. The members of the "Groupe franc;aise d'etude des
algues fossiles" have made many very important contributions in this

direction, but are-study of Triassic dasycladaceans (based on type material)
and a revision of porostromate algae (including comparison with Recent
counterparts and considering changes in sea-water chemistry, cf. Pentecost
and Riding 1986) are still lacking.
3. For some time intervals the information available is conspicuously poor
(Liassic, early Middle Jurassic, Oxfordian) or completely lacking (Early
Triassic). Discussions of evolutionary patterns as well as extinction and
origination rates, can give just a very rough picture of main trends.
4. Studies dealing with the evolution of Triassic and Jurassic calcareous algae
should be focussed on the change of morphological criteria through time.
Those studies should also include investigations of the transition between
Permian and Triassic algae, about the plasticity within algal populations
(e.g. Hurka 1969) and about the possibility of a change in calcification
patterns in time (Leadbeater and Riding 1986).
5. N on-Tethyan calcareous algae are of special interest because they might
include offshoots of the Tethyan stock which could facilitate the dis-
crimination of isocrymal distribution patterns.
6. Algal community analysis seems to provide a powerful tool in supporting
paleoenvironmental reconstructions. A systematic investigation of algal
floras, therefore, should be paralleled by microfacies studies in order to
obtain better estimations of the importance of facies control for algal
distribution patterns.
7. Because petrographical and microfacies criteria of "crusts" supposed to be
of "algal" or microbial origin are controversial, biochemical and geo-
chemical studies of crusts, formed in different paleoenvironments, are
strongly needed.
Acknowledgements. This study is a part of the research program "Evolution of Reefs" supported by
the Deutsche Forschungsgemeinschaft (project FI42/49-1, 2).
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Chapter 22
Mesozoic and Cenozoic Marine Benthic Calcareous
Algae with Particular Regard to Mesozoic
Dasycladaleans
F. BARATTOL0 1
Abstract
During the Mesozoic, benthic calcareous marine green algae are represented principally by the
Caulerpales and Dasycladales. Starting from this era Caulerpaleans consist of genera similar to
extant ones. Dasycladaleans are represented by forms with primary branches only and by ones with
higher orders of branches. Furthermore, the branch arrangement is mainly euspondyle and the
reproductive organs are mainly cladosporous. On the basis of morphological characters and of the
fluctuation in the number of genera and species it is possible to recognize the following stages:
persistence of Late Palaeozoic forms into the Triassic, decline and renewal during the Lower- Middle
Jurassic, Upper Jurassic-Lower Cretaceous acme, and Upper Cretaceous collapse.
During the Cenozoic era, calcified cyanobacteria undergo a numerical reduction. As for the red
algae, gymnocodiaceans and solenoporaceans disappear, while there is an explosive diversification
of the Corallinaceae, and Solieriaceae and Squamariaceae appear.
Among the dasycladaleans, thalli with branches of higher orders in an euspondyle arrange-
ment and with choristoporous reproductive organs prevail. During the Palaeocene this order
shows the greatest number of species.
1 Introduction
The marine benthic calcareous algae are mainly represented by the divisions
Cyanophyta, Rhodophyta, and Chlorophyta.
The literature on calcareous algae has increased remarkably in the last 25
years. It deals mainly with specific subjects; syntheses are still rather poor and
there all still far too few publications on the general significance of the calca-
reous algae.
Among the syntheses, the monographs of Johnson (1952, 1958, 1966),
Johnson and Konishi (1956, 1959), Johnson and H0eg (1961), Roux (1985) all
deal with Palaeozoic algae, and the monograph of Korde (1973) concerns
Cambrian algae. Johnson (1964, 1969) deals with Jurassic-Lower Cretaceous
algae, and Johnson (1960, 1962, 1963, 1965) with the corallinaceans and
solenoporaceans. The syntheses of Bassoullet et al. (1978) and Deloffre and
Genot (1982) cover post-Triassic fossil dasycladaleans, while that of Bassoullet
et al. (1983) deals with the udoteaceans. Among the reviews on the calcareous
algae, the monographs of Pia (1927), Johnson (1961), Korde et al. (1963),
Maslov and Korde (1963), Maslov et al. (1963), Wray (1977b), Emberger (1968),
Dragastan (1980), and Chuvashov et al. (1987), are especially im portant.
'Dipartirnento di Paleontologia. Largo S. Marcellino 10,80138 Napoli, Italy

Mesozoic and Cenozoic Marine Benthic Calcareous Algae 505
The relatively recent bibliographical reviews of Emberger (1976, 1978,

1979) about Paleozoic and Triassic algae, and that of Emberger and Jaffrezo
(1975) about the Jurassic-Lower Cretaceous dasycladaleans, are particularly
useful.
For algal systematics, I refer to Bold and Winne (1978), but for the taxa
above ordinal level the most commonly used suffixes have been utilized. As
regards the geological time-scale, I refer to Odin (1982).
The diagrams shown in the text have tried to utilize, as far as possible, the
data of the most recent literature.
2 Division Cyanophyta
2.1 Stromatolites
Stromatolitic structures (stromatolites and oncolites) are cosmopolitan and

reached their maximum abundance in the Proterozoic and in the Lower
Palaeozoic. They are also well represented in the Mesozoic, although we witness,
in this period, their global gradual decrease. In the Mesozoic they are well
distributed in the carbonate platform facies of the Alpine Chain, particularly in
very shallow environments.
Triassic stromatolitic structures are described by Fischer (1964), D'Argenio
(1966a), Catalano et al. (1974), and liassic stromatolites by, for example,
Colacicchi et al. (1975) and D'Argenio (1966b).
Stromatolitic structures are less frequent after the Lias. Examples are
known from the Palaeogene of the USSR (Maslov 1962), the Miocene of Saudi
Arabia (Whybrow et al. 1987), and of Mazara del Vallo in Sicily (Esteban et al.
1982). Recent stromatolitic structures (laminated algal mats and algal pebbles)
grow mainly in shallow waters and terrestrial high-humidity conditions of
tropical and subtropical areas (Monty 1965, 1967; Gebelein 1969). Rare oc-
currences in temperate zones are also known (Passeri 1974). For further in-
formation see Monty (1977).
Stromatolitic structures, both stromatolites and oncolites, are also well
known in the pelagic facies of the Mesozoic, particularly in the Jurassic
Ammonitico Rosso of the Mediterranean area. Their presence here has
modified interpretations of the sedimentary environment that had been con-
sidered deep water (e.g. Massari 1983; Massari and Dieni 1983; Clari et al. 1984).
2.2 Calcified Cyanobacteria
Among the most common of these in the Mesozoic are the nod ular thalli made
up of branched filaments which radiate from a restricted area of the substratum
belonging to the genus Cayeuxia Frollo and, subordinately, to the genera
Garwoodia Wood, Bevocastria Garwood, Hedstroemia Rothpletz and Ortonella
Garwood.
506 F. Barattolo
This type offossil has also been reported, even ifrarely, after the Mesozoic:
in the Miocene by Reis (1923), who attributed them to Zonotrichites Bor-
nemann; in the Pleistocene by Richter et aI. (1979), who attributed them to
Rivularia haematites (De Candolle). Same of these genera were considered
green algae by Johnson (1961), Elliott (1965), and Emberger (1968). Other
authors ascribe these morphologically similar forms to cyanophytes and group
them in the Garwoodiaceae (e.g. Chuvashov et aI. 1987). Recently Dragastan
(1985) has revised the systematics of this group which he assigns partly to
cyanophyceans and partly to codiaceans; Cayeuxia Frollo in particular is
considered a junior synonymous of Rivularia Roth.
In the back-reef facies of the Upper Cretaceous, the fragments of
filamentous thalli of hormogonaleans can assume a certain local lithogenetic
value (e.g. A eolisaccus kotori RadoiCic andA. barattoloi De Castro: PI. I, 1-2; De
Castro 1975, 1989).
3 Division Rhodophyta
The red algae (subclass Florideophycidae) have a wider stratigraphic and

geographic distribution (Fig. 1). The orders and the families known as fossils
are: Epiphytales (Epiphytaceae), Cryptonemiales (Ungdarellaceae, Sole no-
poraceae, Corallinaceae and Squamariaceae), Gigartinales (Solieriaceae),
Nemalionales (Gymnocodiaceae).
3.1 Epiphytales
The systematic position of these algae is somewhat controversial (see Roux

1985). Chuvashov et aI. (1987) ascribe these algae to Cyanophytes.
Epiphytaleans become less frequent after the Cambrian; Mesozoic Epi-
phyton has been pointed out by Masse (1979) from the Cretaceous of France.
Plate I. Cyanophyta and Incertae sedis. I Aeolisaccus barattoloi De Castro. The distribution of A.
barattoloi in sediment often looks like that ofcyanophycean sheath fragments inside the older parts
oflaminated algal mats from Andros Island. Thin section A.5238.3, Upper Cretaceous (Senonian)
of Regia Piana near Cusano Mutri (Campania. Italy), ca. x120. 2 Aeolisaccus kOlori RadoiCic.
Fragments of sheaths similar to extant Scytonemataceae. Thin section A.1049.5, Upper Cretaceous
(Senonian) of Mt. Tobenna near Giffoni V.P. (Campania, Italy), ca. x120. 3 Paronipora (=
Microcodium) penicillala Capeder. Inside the rock cavities both entire specimens and minute
fragments of Paronipora can be seen. Thin section A.6990.2. Lower Palaeocene (Danian) of Colle
di Medea near Manzano (Friuli, Italy). ca. x40. 4-5 Tubiphyles morronensis Crescenti. Subtrans-
versa I (4) and longitudinal (5) sections. 4 Thin section A.6646.1. Uppermost Jurassic of Capri
(Campania. italy). t:a. x30. 5 Thin ,ection A.6402.1. Uppermost Jurassic-Lowermost Cretaceous of
Capri (Campania. Italy). ca. x30
Plate I
508 F. Bara ttolo
10
5 0.26-0.75
o
I /
0.026 - 0.25
< 0.025 Spec ies/M.A .
..
Ot her t a x a - - - -- - - - -- - - - - - - - - - - -to
-.
«
I- Volvoca l es - - ------- -------- - - -
>-
:r
0-
Caulerpa les
0
'"
0
:r Receptacul itales
...J
t)
Oasycladales
Squamar i aceae
«
>-
>- Corall i naceae
:r
Q.
0
0 Solenoporaceae
0
:r
'" Other taxa 1-----------------------..
Ep i phyta l es
Garwoodiaceae
Fig. I. Qu a ntitative distribution of fossil ma rine benthic calcareous algae during the Phanerozoic.
A rrows indicate extant categories; living species are not included. Time frequency of each group is
represented by the number of species per Ma. Volvocales includes. in the chart. Volvocales.
Thaumatoporellales a nd Tetraspora les. Time scale is from Odin (1982)
3.2 Solenoporaceae
In the Upper Cambrian the family Solenoporaceae appears, the more repre-
sentative genera of which are Solenopora Dybowski and Parachaetetes Den-
inger. The group reached its maximum development in the Palaeozoic and
disappeared during the Eocene. In the Mesozoic and the Cenozoic we find the
genera Solenopora, Parachaetetes and Pycnoporidium Yabe and Toyama; this
latter one, however, according to Emberger (1968) and Schafer and Senow-
bari-Daryan (1983), should be referred to the green algae. Guvencipora Vachard
et al. (1978) is, up to now, only known in the Trias; Marinella Pfender is mainly
known from the Upper Jurassic to the Cretaceous (see Barattolo and Del Re
1984), and Elianella Pfender and Basse in the Cretaceous-Eocene (Hagn and Ott
1975; Poignant 1980).
Neosolenopora Mastrorilli, from the French and Italian Miocene, according
to Tillier's studies (1975), is to be assigned to the Bryozoa. Solenomeris Douville,
from the Upper Cretaceous-Palaeogene reeffacies, according to some authors
(Perrin 1987), is to be ascribed to the foraminifers (Acervulinidae).
Solenopores lived mainly in reef environments or on platforms characte-
rized by rough waters.
3.3 Corallinaceae
The Corallinaceae, probably evolved from the Solenoporaceae, represents by

far the most abundant group of fossil rhodophytes.
The family shows a progressive development from the Upper Cretaceous
up to the present times especially with the genera Archaeolithothamnium
Rothpletz, Lithophyllum Philippi, Lithothamnium Philippi and Mesophyllum
Lemoine. The genera Kymalithon Lemoine and Emberger, Paraphyllum
Lemoine and Hemiphyllum Lemoine occur exclusively in the Cretaceous. Apart
from a Permian species (Endo 1969), the genus Distichoplax Dietrich is res-
tricted to the Paleocene-Eocene. Leptolithophyllum Airoldi is an Oligocene
genus, Aethesolithon Johnson and Pseudoaethesolithon Elliott are exclusively
Miocene genera.
The number of species per million years is about 0.02-0.11 in the Mid-
dle-Upper Palaeozoic, and about 0 and 0.08 respectively in the Triassic and
Jurassic. It increases in an explosive way from the Cretaceous (1.05 species per
Ma.) to Pliocene (9.52 species per Ma.). In the present, corallinaceans densely
populate numerous habitat among the supralittoral to circalittoral zones, and
are spread from Arctic to tropical seas between latitudes 83° Nand 73° S.
In the Mediterranean area they flourish spectacularly in the tidal zone with
the "trottoirs", in the infralittoral zone with the buildups of the "coralligene de
plateau" and in the circalittoral zone with the conspicuous deposits of "maerl"
(Peres and Picard 1964; Peres 1967); they also form banks ofrhodoliths in the
sublittoral areas with strong currents. Corallinaceans are mostly widespread in
510 F. BarattQio
the "Lithothamnium facies" ("Nullipore facies" auet.) of the tropical reefs,

between low-tide level and a depth of about 30 metres.
For further information about the ecology of coralline algae see: Adeyand
MacIntyre (1973), Littler and Doty (1974), Adey and Vassar (1975), van den
Hoek et al. (1975), Adey et al. (1982), Bosence (1985) and Minnery et al. (1985).
In the geological past corallinaceans probably occupied the same envi-
ronments; they are known mainly from high energy habitats. In central and
southern Italy the Lower Miocene Cusano Formation, considered of circa lit-
toral environment (Barbera et al. 1978), is represented by a massive limestone
sequence up to 40 m in thickness, constituted essentially ofrhodoliths, which
can grow up to 25 cm in diameter, together with ostreids, pectinids, bryozoans
and foraminifers (Ogniben 1958).
For the phylogeny, and a review of the main taxa of corallinaceans see, for
example, Poignant (1977, 1979b), and for some palaeogeographical details see
Poignant (1979a) and Poignant and Laville (1972).
3.4 Gymnocodiaceae, Squamariaceae and Solieriaceae
The Gymnocodiaceae, probably related to the living Galaxaura, are known as

fossils mainly from Gymnocodium Pia (Permian) and Permocalculus Elliott
(Permian-Palaeocene). The genera Nanjinophycus Mu and Riding and Oligo-
plagia Herak are restricted respectively to Permian and Triassic. According to
Chuvashov et al. (1987), Succodium Konishi and Dzhulfanella Korde are also to
be ascribed to this family; gymnocodiaceans are referred by these authors to
green algae (order Siphonales). Gymnocodiaceans lived in platform environ-
ments associated with caulerpaleans and dasycladaleans. For further infor-
mation see Elliott (1955, 1956, 1961), Mu and Riding (1983), and Emberger's
(1976, 1979) bibliographic reviews.
The Squamariaceae are known, starting in the Mesozoic, from the genera
Ethelia Weber Van Bossae (= Pseudolithothamnium Pfender and Polystrata
Hydrich) (Cretaceous-Present) and Peyssonnelia Decaisne (Eocene-Present).
F or details of the synonymy of Ethelia, Pseudolithothamnium and Polystrata see
Massieux and Denizot (1964), Denizot (1968) and Buchbinder and Halley
(1985).
The extant family Solieriaceae is known in the Mesozoic from Agardhiel-
lopsis Lemoine (1966), occurring in the Lower Cretaceous (Aptian-Albian) reef
facies in France and Spain.
4 Division Chlorophyta
The Chlorophyceae, like the Rhodophyceae, has left most evidence of its
existence through geologic time in the littoral environments. It is represented as
fossils mainly by the following orders and families: Wetheredellaceae, An-
chicodiaceae, Receptaculitales (Receptaculitidae), Caulerpales (U doteaceae,
Codiaceae), Volvocales (Volvocaceae), Thaunatoporellales (Thaumatoporel-

laceae), Tetrasporales (Chlorangiaceae), Ulotrichales, Chaetophorales (Chae-
tophoraceae), Dasycladales (Seletonellaceae, Beresellaceae, Diploporaceae,
Dasycladaceae, Acetabulariaceae).
4.1 Wetheredellaceae and Anchicodiaceae
Wetheredellaceae populated reef environments mainly from the Devonian to

the Permian and subordinately from the Middle Jurassic to the Upper Cre-
taceous. For further information on this family see Cherchi and Schroeder
(1985) and Chuvashov et al. (1987); the latter authors ascribe the Wethe-
redellaceae to the Dasycladales. During the Mesozoic the family is mainly
represented by Koskinobullina socia lis Cherchi and Schroeder (1979) which
shows close similarities to Aphralysia jurassica Elliott (1982).
The Anchicodiaceae Shuysky (1987b) (= phylloid udoteaceans auct.) lived
in Carboniferous-Permian platform environments, especially in the reeffacies.
This family has been assigned both to red algae (e.g. Wray 1977a) and to the
green algae (e.g. Roux 1985). Shuysky (1987b) and Chuvashov et al. (1987)
ascribe Anchicodiaceans to the green algae order Siphonales; however several
recent classifications do not include the Siphonales anymore, which have been
split into several orders.
4.2 Caulerpales
Udoteaceae
The Udoteaceae is represented in the Recent by fourteen genera; they are well
known in geologic time from about twenty-five genera of which one is still living.
Udoteaceans appeared in the upper Cambrian and became widespread in the
Silurian-Devonian, in the Permian, and subsequently in the Palaeocene-
Eocene. The more frequent Mesozoic-Cenozoic genera are Boueina Toula (PI.
II, Fig. 2), Arabicodium Elliott, Halimeda Lamouroux and Ovulites Lamarck
(see Bassoullet et al. 1983). Udoteaceans lived in environments similar to those
of the dasycladaleans, populating lagoon and also shelf margin facies (e.g.
Nipponophycus Yabe and Toyama and luraella Bernier 1984).
Codiaceae
About forty genera have been assigned to the Codiaceae but subsequently, with
the improvement of the systematics, have been transferred to other families.
Forms like Halimeda Lamouroux and Boueina Toula are generally placed in the
udoteaceans. There is today a certain consensus in reassigning the Gar-
woodiaceae (= Mitcheldeaniaceae auct.) to cyanophyceans (Riding 1977;
Schafer and Senowbari-Daryan 1983; Chuvashov et al. 1987). The An-
chicodium-like forms (= phylloid Udoteaceans auct.) are grouped together in
512 F. Barattolo
Plate II
the Anchicodiaceae (Siphonales) by Shuysky (1987b) and Chuvashov et al.

(1987).
Recently Dragastan (1985) attributed some new genera established by
himself from the Mesozoic of Romania to codiaceans, and also considered
Garwoodia Wood and Bevocastria Garwood to be codiacean algae.
4.3 Volvocales, Thaumatoporellales and Tetrasporales
The order Volvocales is represented as fossils by mobile colonies (Volvocaceae)

that lived particularly in shallow-water carbonate facies. According to Kaz-
mierczak (1979, 1981) it is known in the Early Proterozoic with Eosphaera
Barghoorn, according to Hofmann and Schopf(1983) Eosphaera is an incertae
sedis genus (prokaryote?). Volvocaleans occur in the Devonian and Carbon-
iferous with Eovolvox Kazmierczak and other taxa formerly attributed to
non-radiosphaerid calcispheres and parathuramminid Foraminifera (Kaz-
mierczak 1975, 1976, 1981).
The order Thaumatoporellales (PI. II, 1) is wide-spread in the Mesozoic
(from the Late Triassic onwards) carbonate platform facies. This order has been
recently established by De Castro (1988a); it resembles volvocaleans in some
characters and chladophoraleans in others.
The order Tetrasporales is represented as fossils by sessile dendroid forms
(Chlorangiaceae) that flourished in shelflagoons; they are well known in the
Cretaceous with Sgrossoella De Castro (1969) (PI. 11,4), related to the living
genus Ecballocystis Yengar, and probably by the genus Cretacicladus Luperto
Sinni (1979) (PI. II, 3) related to the living Prasinocladus Kuckuck.
4.4 Ulotrichales and Chaetophorales
The Ulotrichales is mainly known from the genus Vermiporella Stolley. This
genus, once attributed to dasycladaleans (Pia 1920), is referred by Kozlowski
and Kazmierczak (1968) to this order (family uncertain) while it is considered
a foraminifer by Vachard (1976). This Palaeozoic genus also doubtfully occurs
in the Lower Cretaceous (Conrad 1970).
The genus Heterotrichella established by Schafer and Senowbari-Daryan
(1983), is tentatively referred by the same authors to the Chaetophorales.
Plate II. Chlorophyta (Caulerpales. Thaumatoporellales and Tetrasporales). 1 Assemblage of

thaumathoporellaleans. Sections of cylindrical and globose thalli. sometimes with younger thalli
inside (middle part of the figure). can be seen. Thin section A.5580.20. Uppermost Cretaceous of
PliaCiaz (Mali Ston. Yugoslavia), ca. xlO. 2 Boueina sp. Longitudinal section. Thin section
A.1974.27. Upper Cretaceous (Cenomanian) ofMt. Cerreto near Caserta (Campania, Italy), ca. x30.
3 Cretacicladus minervini Luperto-Sinni. Longitudinal sections. Thin section A.1982.1. Cretaceous
(Late Albian-Early Cenomanian) of Mt. Cerreto near Caserta (Campania, italy). ca. x80.
4 Sgrossoe/[a partenopeia De Castro. Longitudinal section. Thin section A.3028.35. Upper Cre-
taceous (Senonian) of Cerreto Sannita (Campania. Italy). ca. x40 (De Castro 1969)
514 F. Barattolo
4.5 Dasycladales
The dasycladaleans are the most numerous Mesozoic fossil algae when com-
pared to other groups. They are in fact represented by nearly 200 genera and
little less than 900 species ranged from the upper Cambrian to the Recent. They
attained high abundances in the Carboniferous-Permian, in the Upper Juras-
sic-Lower Cretaceous, and in the Palaeocene-Eocene. The genera have an
average existence of about 53 Ma. The average number of species per genus is
4.7 (Fig. 2).
Understanding of this group of organisms has been made possible thanks
mainly to the numerous contributions by Julius Pia, above all his "Neue studien
iiber triadischen Siphoneae Verticillatae" (1912), the specifications given in Pia
(1920), and finally the systematic scheme (Pia 1927) contained in Hirmer's
Handbuch der PaHiobotanik. Other important contributions include the
pioneering work of Steinmann (1899, 1903), who was the first to show the
importance of the position of reproductive organs for the systematics and the
phylogeny of this order. Morellet and Morellet's studies (1913, 1922) on the
Cenozoic dasycladaleans of the Paris Basin also hold particular interest.
For reference to numerous and often valuable contributions which have
appeared in more recent times see Johnson and Konishi (1959), Johnson and
H0eg (1961), Korde (1973) and Emberger's reviews (1976, 1978) for the
Palaeozoic; Emberger (1979) for the Triassic and the synthesis of Bassoullet et
al. (1978) for the Jurassic-Cretaceous and, finally, Deloffre and Genot (1982)
and Genot (1987) for the Cenozoic. For recent classifications see Chuvashov et
al. (1987), Deloffre (1987,1988) and Shuysky (1987b).
General considerations of taxonomic criteria and concepts are supplied by
Pia (1920), and Bassoullet et al. (1977, 1984). Statistical and biometrical
methodologies have also been used to describe fossil dasyclad species (e.g.
Remane 1969; Zorn 1974, 1977a; De Castro and De Rosa 1977; DeCastro 1979;
Barattolo 1978, 1980, 1981, 1982a,b,c, 1983a,b, 1984).
Aspects of reproduction in fossil dasyc1adaleans and its evolutionary
significance are discussed by Steinmann (1903), Pia (1920), Elliott (1972),
Barattolo (1980, 1983a) and Bucur (1985). Considerations of evolution, are
supplied by Herak et al. (1977). The function of the branches within the
framework of the algal construction is discussed by Pia (1920), Zorn (1972),
Barattolo (1978), Elliott (I 979b), Skompski (1984).
Palaeozoic Dasycladaleans
It is relevant here to briefly review the pre- Mesozoic history of dasyc1adaleans.
Cambrian-Devonian dasyc1ada-leans belong mainly to the Seletonellaceae,
secondly to the Beresellaceae, occasionally to Diploporaceae and Dasy-
c1adaceae. In the Devonian, Acetabulariaceae seem also to be represented by
the genera Lulipora Shuysky (1986) and Lopsiella Shuysky (l987a). These
Lower Palaeozoic forms are mainly characterized by the following features:
irregular shaped or simple thallus, the presence of only primary branches which
44
•• 16-17
Number
..... 6 - 7 01
40
. 1
••• 1
Genera
36 .
:>
c: T 1.
L
since Ceno zoic
since Mesozoic
....
(!)
since Paleozoic
32 ..
a.
<.>
28
'"a.
'"
24
20
16
12
4
/-
o Life of Genera
o 40 80 120 160 200 240 M.A .
Fig. 2. Dasycladales. Number of species per genus plotted against the life of genera in Ma . Circles.
squares and triangles represent genera which appeared in the Palaeozoic, Mesozoic and Cenozoic
respectively; small symbols represent a single genus, medium-sized ones represent 6- 7 genera and
large ones represent 16- 17 genera . The number of species per genus and the life of genera seem to
be linked by an exponential law. The chart shows a high frequency of genera with few species and
a low frequency of genera with many species. Outside the two curves, towards the right. relatively
new and/or poorly known genera, together with others having doubtful species or an uncertain
stratigraphical range fall
are in aspondyle arrangement, and reproductive organs of endosporate-type.

Reproductive features can be only inferred for the majority of taxa with
relatively thin branches because these ones were presumably not suited to
contain reproductive organs. Fewer genera show twofold branches, euspondyle
arrangement, and reproductive organs of c1adosporate-type (Fig. 3). With the
genus Archaeobatophora Nitecki (1976) in the Ordovician the presence of a
choristosporous tribe (Batophoreae) has been noted. Nevertheless, this genus
has no trace of choristosporate reproductive organs. During the Lower
Palaeozoic and Devonian dasyc1adaleans are principally represented by a small
number of genera (45), mainly characteristic of each period, and by ca. 80
516 F. Barattolo
ARRANGEMENT OF POSITION OF
"'IEE
PRIMARY BRANCHES REPRODUCTIVE ORGANS
~~
.'.. ..... 0 ~ 00
'. :':tA :::::

• • ..... <> 0
~~
0
~
• p . hO t . 00 0
, : ; .:.::.:.: 0 ~ ~ 0 0
abc d e
1.8
:.. 0
'
0 P.
65
0
0
o 0
K
[]
o
'.~.
0
0
o .. IJ []
Tr
245
p
o
0
Ca
530
Fig. 3a-f. Dasycladales. Stratigraphic distribution of primary branch arrangement (a-c) and
position of reproductive organs (d-f) during the Phanerozoic. a aspondyle; b euspondyle; c
metaspondyle; d endosporate-type; e cladosporate-type; f choristosporate-type. A broken line
indicates doubtful or very rare presence. Time scale is from Odin (1982)
species. The number of species per genus is also rather low: about 1.4
(Fig. 4).
The Carboniferous-Permian represents the period of maximum Palaeozoic
expansion of dasycladaleans. They belong particularly to the Seletonellaceae,
BereseUaceae, Diploporaceae and Dasycladaceae. The order is represented by
about 80 genera and 300 species; the importance of the dasycladaleans in this
interval is also shown by the relatively high number of species (2.6 species per
Ma) related to the duration of the period (lIS Ma).
The morphological styles which appeared in the preceding periods
(coarsely branched thalli, aspondyle branch arrangement, endosporous re-
Mesozoic a nd Cenozoic Marine Benthic Calcareous Algae 517
0
1.8
ci 10 20 30 40 50
~
6
0 - - - genera
23 N
33 0
------
Z E
51 W
65
<.) Pa
--
<;:
K
<.)
-,
~
130 ~
--- ---
~
0 J
N
0 r- --
en I
,,
I
w I
204 ::!:
)
Tr
245 ....
---
_--- ?
P \
, )
290
<.)
-
C
---- - -- - --
360 0
N
0 D
---
-----;>.
---
~
400
w
4 18
---' S
<{
Q..
0 ,- >
,-
",
,-
495
Ca - - s pe c ie s/ M.A.
530
2 3 4 5 6 7 8 9
Fig.4. Dasycla d a les. N u m ber of ge ne ra pe rtime interva l and number o f ps ecies per tim e interva l
pe r M a during the Ph a nerozo ic. No Lower Tri assic spec ies are known. th e va lu e suppli ed i smerely
specul a tive . L owe r Triassic ge nera a re the o nes prese nt both in the Palaeozo ic a nd in th e Middle
T riass ic. Only stro ngly calcified li vin g s pecies a re taken in to consid era ton f orth e Qu a tern a ry va lues.
Time sca le is fro m Odin ( 1982)
productive organs) persisted in the Carboniferous-Permian, while the more

"evolved" ones th at were present, but not well represented in th e Ca mbria n-
Devonian, become important.
Th e principal l atter characters o f the th alli are:
I. A prevalently euspondyle distribution of th e branches, even though
aspondyle (e.g. Antrachoporella Pia, Gy roporella Gumbel) and metaspondyle
arrangements a re a sol well represented . Metaspondyle dasycladalea ns are
represented b yas many "vestibuled" taxa (e.g. Velebitella Kochansky-Devide
and A lbertaporella Johnson) as " non-vestibuled" ones (e.g. Diplopora Shafh auti
518 F. Bara ttolo
and Clavapora Guvenc). For details of the shape and the systematic value ofthe
vestibule in the metaspondyle forms, see Mamet and Roux (1981,1983);
2. Reproductive organs are mainly of endosporate type and, secondly, of
cladosporate type; the taxa with relatively swollen branches, that presumably
could have contained reproductive organs, are probably to be assigned to the
cladosporous forms: Gyroporella Gumbel, characterized by vesiculiferous
branches, Physoporella Steinmann characterized by piriferous branches and
Kulikia Golubtshov with swollen phloiophorous branches;
3. Prevalence of forms with only primary branches, even though twofold
branched genera with dichotomic (e.g. Antracoporella Pia) and the tuft-like
ramifications (e.g. Dasycladophycus Elliott) also occur. The branches are of
akrophorous (e.g. Queenslandella Mamet and Roux), trichophorous (e.g. Li-
kanella Milanovic and Teutloporella Pia) and phloiophorous (Mizzia Shubert)
type; vesiculiferous (e.g. Gyroporella Gumbel) and piriferous (e.g. Physoporella
Steinmann) morphological variations also occur.
Mesozoic Dasycladaleans
The Mesozoic dasycladaleans belong mainly to the Dasycladaceae, to a lesser
extent to the Acetabulariaceae and Diploporaceae, occasionally to the Sele-
tonellaceae. The Mesozoic is on the whole an era of great diversification of
dasycladaleans; these undergo remarkable variations both in the vegetative
parts and the reproductive organs. The vegetative parts show the disappearance
of the metaspondyle and aspondyle forms, although the metaspondyle arrange-
ment seems to reappear, almost like an accident, with the genus Furcoporella
Pia in the Palaeocene. The endosporous forms disa ppear in the Mesozoic; while
the cladosporous forms are widespread and the cladosporous reproductive
organs exhibit the greatest degree of diversification. Choristosporous forms
appear and attain a moderate importance in the Mesozoic.
The following phases can be identified during the era in the history of
dasycladaleans: persistence of Late Palaeozoic forms into the Triassic, decline
and renewal during the Lower-Middle Jurassic, Upper Jurassic-Lower Cre-
taceous acme, Upper Cretaceous collapse.
Persistence of Late Palaeozoic Forms into the Triassic. Contrary to what was
once believed, the Triassic represents a period of decrease of dasycladaleans
when compared to the Permian and especially to the Lower Permian, and in fact
no dasycladaleans are known to occur in the Lower Triassic. In the period as a
whole (41 Ma), altogether about 24 genera are present (16 in the Middle Trias
and 19 in the Upper), and 80 species (56 in the Middle Trias and 26 in the
Upper); the number of species per Ma is about 1.95.
Compared to the Permian and especially to the Upper Permian the number
of species per genus is the same (4.5) while the number of genera decreases
considerably (37 in the Upper Permian, 16 in the Middle Trias).
During the Triassic the characters of the thallus, and most of the taxa, are
still those of the Carboniferous-Permian. In the Middle Triassic the genera
Diplopora Schafuautl, Teutloporella Pia, Oligoporella Pia, Physoporella Stein-

mann and Macroporella Pia prevail and Kantia Pia, Clavapora Giivenc, Ani-
soporella Botteron, Griphoporella Pia and Neoteutloporella Bassoullet et al.
appear.
Diplopora annulata Schafbautl (Pl. III, 2) and Oligoporella chia Herak seem
to be species adapted to live in the middle parts of the lagoons (Ott 1972);
Diplopora nodosa Schafuautl (PI. III, 5) and Teutloporella herculea (Stoppani)
seem to be adapted to live in the outer lagoon facies (Ott 1972, Zorn 1976, De
Castro 1979); Teutloporella echinata Ott flourished in the reef facies (Zorn 1976).
Zorn (1976), in particular, has shown that the epicontinental platform facies
that received a terrigenous sedimentary contribution, were populated by Di-
plopora hexaster-like forms and Oligoporella and Physoporella species with
paired whorls. The epioceanic shelf lagoons were mainly populated by
Macroporella, Teutloporella and Oligoporella or Physoporella species with
simple whorls. Further information on the distribution of numerous dasy-
cladaleans in the lagoon facies is given by Dragastan (1981).
In the Upper Triassic most of the preceding genera disappear, there is a
rapid reduction of the dasycladaleans and at the same time same renewal. The
more frequent genera are Griphoporella Pia, Gyroporella Giimbel and Poi-
kiloporella Pia (= Andrusoporella Bystricky). Heteroporella Praturlon, Pen-
taporella Senowbari-Daryan and Placklesia Bilgutay appear; the last two rather
recently established genera do not seem to survive to the end ofthe period. With
the appearance of Heteroporella the Upper Trias marks the appearance of
choristosporous reproductive organs. However, this character, seen in Hete-
roporella and in other genera (Cylindroporella Johnson, Neomeris Lamouroux
and Cymopolia Lamouroux), only gains importance in the Cretaceous.
In the lagoon facies, more or less close to the outer lagoon margin,
Griphoporella curvata (Giimbel; PI. III, 1), Heteroporella zankli Ott and Hete-
roporella crosi Ott dominate; in shelf margin facies Diplopora tubispora Ott and
Diplopora phanerospora Pia flourished; in the shoal facies it is possible to find
Placklesia multipora Bilgutay, probably redeposited (Abate et al. 1977; Fliigel
1979; Senowbari-Daryan and Schafer 1979; Di Stefano and Senowbari-Daryan
1985).
Decline and Renewal During the Lower-Middle Jurassic. Compared with the
Upper Triassic, the Lower-Middle Jurassic represents a period of decrease of
dasycladaleans. The number of genera (about 17 in the Lias and 18 in the
Dogger), when compared with the Upper Triassic (about 19 genera), remains
more or less the same; similar behaviour is seen by the species/genera ratio (1.4
in the Upper Trias, 1.5 and 1.4 respectively in the Lias and in the Dogger).
However, the number of species tends to decrease even compared with the
duration of the time interval (species/Ma.).
Lower-Middle Jurassic dasycladaleans are almost exclusively euspondyle
and the number of genera with only primary branches is approximately the
same as that with more orders of branches. Reproduction was prevalently of
cladosporate-type (e.g. Dissocladella Pia, Fanesella Cros and Lemoine, Sar[a-
520 F . Barattolo
Plate III
tiella Conrad and Peybernes, Stichoporella Pia etc.), and maybe of choris-
tosporate-type as well (Eodasycladus Cros and Lemoine). The branches are
trichophorous (e.g. Selliporella Sartoni and Crescenti and Neoteutloporella
Bassoullet et al.), akrophorous (e.g. Fanesella Cros and Lemoine, Dobuniella
Elliott etc.), and phloiophorous (e.g. Neomizzia Levy, Dinarella Sokiie and
Nikler, Sar[atiella Conrad and Peybernes and Stichoporella Pia).
Almost half of all Lower Jurassic genera appear during this time interval;
the other half had already mainly appeared in the Upper Triassic. The more
frequent Liassic genera are Palaedasycladus Pia and Sestrosphaera Pia; besides
these two genera, Dinarella Sokiie and Nikler, Tersella Morellet and Eodasy-
cladus appear and are also restricted to the Lias; whereas Neomizzia, Disso-
cladella and Fanesella are also known in younger rocks. Palaedasycladus
mediterraneus Pia (PI. IV, 1) and P. barrabei Lebouche and Lemoine seem to be
present both in the lagoon facies and in those close to the shelf margin. For
further detailed information see mainly Sartoni and Crescenti (1959), De Castro
(1962), Chiocchini (1977), Chiocchini and Mancinelli (1977, 1978) and
Peybernes (1979).
The genera Sar[atiella Conrad and Peybernes, Coniporella Fischer and
Thierry, Dobunniella Elliott, Stichoporella Pia, Selliporella Sartoni and Cres-
centi appear in the Dogger; the last three are restricted to the Dogger. Sel-
liporella has been recently redescribed by Barattolo, De Castro and RadoiCie
(1988).
In the lagoon facies Selliporella donzellii Sartoni and Crescenti (PI. IV, 5) is
dominant; Sar[atiella dubari Conrad and Peybernes dominates in the lagoon
facies next to the shelf margin (De Castro 1962, 1987; Sartoni and Crescenti
1962; Chiocchini and Mancinelli 1977; Sokiie and Velie 1978a,b; Peybernes
1979).
According to Elliott (1977b, 1982, 1984) dasycladaleans gradually spread
out, starting in the Triassic, from tropical towards temperate zones; in the
Middle Jurassic in particular it is possible to find a differentiation due to climatic
causes between the dasycladaleans of northern and southern Europe.
Upper Jurassic-Lower Cretaceous Acme. The Upper Jurassic-Lower Cre-
taceous represents a time interval of considerable increase of dasycladaleans; in
fact we pass from 11 species in the Dogger to 73 species in the Maim and to 131
species in the Lower Cretaceous. The same goes for the behaviour of the number
Plate III. Chlorophyta (Dasycladales). 1 Griphoporella curvata (Gumbel). Longitudinal-oblique

section. Thin section A.7776.4, Upper Triassic (Rhetian), Erchie (Campania, Italy), ca. x15.
2 Diplopora annulata Schafhautl. Oblique section. Thin section A.6884.1, Middle Triassic, vicinity
of Gracac (Velebit, Yugoslavia), ca. xIO. 3-4 Terquemella sp. Respectively equatorial and axial
sections. These small lenticular bodies represent calcified reproductive organs of some dasy-
cladaleans (e.g. Bornetelleae). Thin section A.2959.2. Lower Cretaceous (Aptian-Albian) of Piano
di Pecore near Laviano (Campania. Italy), ca. x 120.5 Diplopora nodosa Schafhautl. Longitudinal
section. Thin section A.30IO.24, Middle Triassic of II Castello near Marsico Nuovo (Basilicata,
Italy), ca. x 10
522 F. Barattolo
Plate IV
of genera, passing from 18 genera in the Dogger to 37 genera in the MaIm and
to 43 genera in the Lower Cretaceous; this last value represents the generic
maximum during the Mesozoic and Cenozoic.
Particularly in the MaIm there is a considerable generic renewal (about
60%), while the number of species per genus remains rather low; in the Lower
Cretaceous the opposite happens: the percentage of new genera decreases while
the number of species per genus increases considerably (Fig. 5). A similar
phenomenon happens in the Palaeocene-Eocene.
As regards morphology, thalli have mainly euspondyle branches;
aspondyle-branched forms (e.g. Megaporella Deloffre and Gyroporella GUm-
bel) only occasionally occur. The reproductive organs are mainly cladosporous
(Neogyroporella Yabe and Toyama; Sar[atiella Conrad and Peybernes; Pra-
turlonella Barattolo, PI. VII, 1; Triploporella Steinmann; Suppiluliumaella
Elliott; Pseudotriploporella Jaffrezo and Poisson; Megaporella Deloffre; Gy-
roporella GUmbel, etc.) even though the choristosporate-type is not uncommon
(Cylindroporella Johnson, Heteroporella Praturlon, Neomeris Lamouroux, PI
VII, 2; Actinoporella GUmbel, etc.); doubtful endosporous forms also occur
(Linoporella Steinmann, Campbelliella RadoiCic, etc.).
About two-thirds of the genera included in this time interval have twofold
branches.
In the Upper Jurassic-Lower Cretaceous Salpingoporella Pia, Clypeina
Michelin, Cylindroporella Johnson, Heteroporella Praturlon, and Triploporella
Steinmann (PI. IV, 2) are dominant.
Isolated calcified reproductive organs, probably assignable to dasy-
cladaceans (Russoella Barattolo and Terquemella Morellet and Morellet; PI. III,
3-4 and PI. IV, 3-4) and acetabulariaceans (Acicularia D'Archiac), also occur.
In the lagoon and/or inner margin facies Salpingoporella grudii RadoiCic,
S. annulata Carozzi (PI. VI, 2), Clypeina jurassica Favre (PI. V, 1) and Camp-
belliella striata (Carozzi) (PI. V, 2) dominate. In the margin facies Salpin-
goporella pygmaea (GUmbel), Neoteutloporella socialis (Praturlon) (PI. VI, 1)
and Petrascula bursi[ormis (Etallon) (Praturlon 1963, 1966; Chiocchini and
Mancinelli 1978; FlUgel 1979; Peybernes 1979; Bernier 1984) dominate. In
the Lower Cretaceous lagoon facies Triploporella neocomiensis RadoiCic,
Plate IV. Chlorophyta (Dasycladales). 1 Palaeodasycladus mediterraneus Pia. Oblique section.

Primary, secondary and, less clearly, tertiary branches can be seen. Thin section 420.1, Lower
Jurassic ofL'Ascensione near Nocera lnferiore (Campania, italy), ca. xiS. 2 Triploporella marsicana
Praturlon. Oblique section. Notice the large, stout primary branches and the tufts of secondaries.
Thin section A.5874.45, Lower Cretaceous of Serra Campo dell'Acero near Matese Lake
(Campania, italy), ca. xiS. 3-4 Russoella radoiciciae Barattolo. Axial (3) and equatorial (4) sections.
Russoella represents calcified reproductive organs of some dasycladaleans (e.g. Triploporelleae and
Bornetelleae). For both figures: thin section A.2l68.4, Lower Cretaceous (Albian) ofMt. Calvi near
Caserta (Campania, italy), ca. x120. 5 Selliporella donzellii Sartoni and Crescenti. Oblique section
by De Castro (1962). Short and stout primary branches and tufts of two trichophorous secondary
branches can be seen (see also Barattolo, De Castro and RadoiCic 1988, Fig. I). Thin section 269.1,
Jurassic (Middle Dogger), Case Ciaramello near Nocera Inferiore (Campania, italy), ca. xiS
524 F. Barattolo
0
1.8
0 2 3 4 5 6
6
S2
0 ----
species / genera
---
N
23
33 0
Z >
UJ
51 ...-
.- .-
65 U Pa
K
U
130
0
N
0
en
UJ
204 ~
Tr
245
P
290
C
U
360 0
N
0 D
400 UJ
418
-'
«
S ,
~ "-
"-
0 ,/
)
/
,/
/
49 5 ,/
Ca ,/ - percentage of new genera

530
20 40 60 80 100
Fig. 5. Dasycladales. Species-genera ratio and percentage of new genera (first appeared genera-
total genera ratio per time interval) during the Phanerozoic. The species-genera ratio in the Lower
Triassic is merely specu lat ive. Only strongly calcified living species are taken into consideration for
the Quaternary values. Time sca le is from Odin (1982)
Plate V. Chlorophyta (DasycJadales). J Clypeina jurassica Favre. Variously sectioned verticils of

disarticulated thalli. Thin section 421.1, Uppermost Jurassic of Colle di S, Pietro near Nocera
Inferiore (Campania, Italy), ca. x 10.2 Campbelliella striata (Carozzi). Variously sectioned verticils
of disarticulated thalli. Thin section A.6834, Uppermost Jurassic-Lowermost Cretaceous, Tuoro di
Sasso near Nola (Campania, Italy), ca. xlO
Mesozoic and Ce nozoic Marine Ben thic Calcareous Algae 525
Plate V
526 F. Barattolo
Plate VI
Macroporella embergeri Boroullec and Deloffre, Likanella campanensis Azema

and Jaffrezo, Clypeina radici Sokiie and Velie and Salpingoporella dinarica
RadoiCie (PI. VI, 3) dominate.
In the margin facies Macroporella praturloni Dragastan, "Griphoporella"
piae Dragastan and Salpingoporella genevensis Conrad, A crop orella radoiciciae
Pra turlon etc. (e.g. Praturlon 1964, 1966; Chiocchini and Mancinelli 1977, 1978;
Conrad 1977; Velie 1977; Sokiie and Velie 1978b) dominate.
Upper Cretaceous Collapse. In the Upper Cretaceous (duration about 30 Ma)

we witness a rapid decrease in the number of both genera and species of
dasyclada1eans; the former decreases from 44 (Lower Cretaceous) to 23; the
calculation takes into account the Permian genus Salopekiella Mi1anovic, the
presence ofwhich in the Maastrichtian (S. inopinata GUS-ie) is perplexing (Herak
et aI. 1979). The number of species decreases from 131 (Lower Cretaceous) to45.
The number of species per Ma is only 1.5.
All the Upper Cretaceous genera a part from H arlanjohnsonella Elliott were
present earlier and mainly in the Lower Cretaceous; Likanella Mi1anovie is
known from the Permian too. The percentage of new genera is very low (4%);
ever lower values can also be found in the Upper Devonian, Lower Triassic and
Neogene.
As regards morphology, the thalli are exclusively euspondyle; the re-
productive organs are mainly either cladosporous (Triploporella Steinmann,
Harlanjohnsonella Elliott, Suppiluliumaella Elliott, Dissocladella Pia, etc.) or
choristosporous (Neomeris Lamouroux, Cymopolia Lamouroux, Cylindropo-
rella Johnson, etc.), even if the presence of endosporous species cannot be
excluded (Acroporella Praturlon).
Most genera show twofold branches. The most common genera are Trino-
cladus Raineri, Cymopolia Lamouroux (PI. VII, 4), Neomeris Lamouroux,
Salpingoporella Pia, Clypeina Michelin, Cylindroporella Johnson (PI. VII, 3) and
Heteroporella Praturlon.
Cenozoic Dasycladaleans
Cenozoic dasycladaleans belong mainly to the Dasycladaceae and Ace-
tabulariaceae. Other families are either not known, or their presence is
questionable.
The beginning of the Cenozoic marks a renewal in the development of
dasycladaJeans. In the Palaeocene (duration about 14 Ma) the number ofgenera
...
Plate VI. Chlorophyta (Dasycladales). 1 Neoteutloporella socialis (Praturlon). Longitudinal-ob-
liq ue sections of closely packed thalli. Thin section 697. U ppermostJ urassic of Mt. Como (Abruzzi,
Italy), ca. xiO. 2 Salpingoporella annulata Carozzi. Variously sectioned specimens. Thin section
A.4498.1. Lowermost Cretaceous (Neocomian) ofMt. Monaco di Gioia near Faicchio (Campania,
Italy), ca. x25. 3 Salpingoporella dinarica (RadoiCic). Variously sectional specimens. Thin section
A.5940.4, Lower Cretaceous (Aptian), Suio Terme (Latium, Italy), ca. x25
528 F . Bara ttolo
Plate VII
rises from 23 (Upper Cretaceous) to 39; the number of species rises from 45 (Upper
Cretaceous) to lO6. The number of species per Ma is 7.5; when compared to
those of other geologic intervals this value represents the absolute maximum
(Fig. 4). The percentage of new genera and the number of species per genus (Fig.
5) show a similar behaviour to the Upper Jurassic-Lower Cretaceous interval.
After the Eocene the group gradually decreases in importance up to present.
In Figs. 4 and 5, in order to make the Recent data comparable with the fossil
ones, only extant genera and species with significant calcification have been
taken into consideration.
In the Recent the order is represented only by the two families, Ace-
tabulariaceae and Dasycladaceae, both living in environments comparable to
those of the past, i.e. mainly in the shelf lagoon or inner platform and subor-
dinately in shelf margin habitats (Valet 1979). They mainly populate the tropical
and subtropical zones, with few species in the Mediterranean: Acetabularia
mediterranea Lamouroux, A. calyculus Quoy and Gaimard, A. parvula
Solms-Laubach and Dasycladus vermicularis (Scopoli). Only the first of these
species calcifies significantly.
The family Acetabulariaceae is at present represented by two living genera
and 20 species. The family Dasycladaceae is represented by 6 genera and 19
species. The forms that live in reef environments belong mainly to the genera
Bornetella Munier-Chalmas, Neomeris Lamouroux and Acetabularia Lamou-
roux (section Polyphysa).
For the stratigraphical distribution of Mesozoic and Cenozoic dasy-
cladaleans see Fig. 6.
5 Incertae Sedis
Among the benthic algae a certain number of taxa is present which, although
assignable to the algae on the basis of their vegetative simplicity, are not closely
comparable with known taxa of thallophytes. They are not taken into con-
sideration in Fig. I. These incertae sedis forms are numerous; some ofthem have
already been dealt with and assigned to higher taxa as proposed by some
authors.
Plate VII. Chlorophyta (OasycIadales). 1 Praturlonella salernitana Barattolo. Longitudinal sec-

tion. Thin section A.3519.a.76, Lower-Middle Eocene of Mt. Vesole near Trentinara (Campania,
Italy), ca. x30 (Barattolo 1978).2 Neomeris cretacea Steinmann. Oblique section. Note the ellip-
soidal ampullae and the slender secondary branches, primary branches are uncalcified. Thin section
n.9, Lower Cretaceous (Albian) of Cerro Escamela near Orizaba (Varacruz, Mexico), ca. x20.
3 Cylindroporella kochanskyae RadoiCic. Oblique-tangential section. Note the sterile (secondary)
branches and reproductive organs (ampullae) alternated in the same verticil. Thin section A.4l56.1,
Upper Cretaceous (Early Turonian) of Milanovac plantae (Okovan, Yugoslavia), ca. x40.
4 Cymopolia cf. paronai Raineri. Oblique section. Primary branches, ampullae and tufts of
phloiophorous secondary branches can be seen. Thin section A.6750.1, Upper Palaeocene
(Thanetian) of Colle di Medea near Manzano (Friuli, Italy), ca. x40
530 F. Barattolo
Pseudocymopolia
Pseudoc 1y peina
-
Radoiciciella
Neoqyroporella
-
Apinella
Megaporella
Conodictyum
Thyrsoporella
-
.... -
--
Cylindroporella
Goniohna
Myrmeltioporella
Acicularia
-
Coniporella
--
Se llipore lla
Stichoporella
Dobunniella
Sarfatiella
--
Fanesella
Sestrosphaera
Eodasyc1adus
Dissocladella
--
Neomizzia
Tersella
-
Palaeodasyc1adus
Dinarella
Heteropore Ila
Clypeina
Uragiella
Salpingoporella
---
Petrascula
Plack lesia
Pentaporella
-
Holospore\la
Aciculopora
Griphopore lla
---
N eoteutioporella
Kantia
Anisoporel\a
Clavapora
Salopekiella
.. ............ .
Likanella
...
Gyroporel\a
Pseudoepimastopora ........
I
.......
I
Macroporella
Epimastopora
Teu tiopore lla ..
Poikiloporella
,
Aciculella
Diplopora
Ph ysopore lla --
-
Poncetella
Oligoporella
I I I I I
A
Tr J K Pa E pi M
Fig.6A,B. Dasycladales. Stratigraphic distribution of Mesozoic and Cenozoic genera .• Genus
known also in the Palaeozoic;. extant genus; ----: genus with uncertain stratigraphical range; •••• :
uncertain occurrence of the genus. Time scale is from Odin (1982)
Acetabularia
Dactylopora ~
Sarosiella ~
Carpenterella -~
Belzungia -~
--
Tibetipora -~
Uglasiella
Microsporangiella
Parkerella 1-
Montiella
Ollaria
Ferganella
---_.. ......
1-
1--
Catellaria ----
Zittelina
Frederica l - I--
Uteria l - I--
---- ·
Jodotella l - I--
Furcoporella I--
Sedalanella ~
-·
Rostropore Iia
-- ·
Pagodaporella
-- ---
Orioporella
Morelletpora
Indopolia
-- ·
Hamulusella
Broeckella
-
Morelletina
Sandalia
-
Piania
Harlanjohnsonella
...- ...
Cymopolia
Russoella
...
Halicoryne
Pseudotriplopore lIa
Neomeris
-
---
Praturlonella
Pseudoactinoporella
Angioporella
--
Korkyrella
Helioporella
Crinella
Suppiluliumaella
Balkhanella
Acroporella
-
Terquemella
Triploporella
Linoporella
,
-I--
--I--
Humiella
Kopetgaria
Campbelliella ~
Trinocladus
Actinoporella
I I I I I
B
Tr J K Pa E PI M
532 F. Barattolo
Some other incertae sedis taxa mentioned mainly in the Mesozoic are
Paronipora Capeder, Bacinella RadoiCie, Lithocodium Elliott, Tubiphytes
Maslov, and Baccanella Pantie.
The taxonomic and stratigraphic problems of Paronipora Capeder 1904 (=
Microcodium Gluck 1912) (PI. 1,3) have been reviewed recently by Cherchi and
Schroeder (1988). This fossil shows similarities with the mineral productions
caused by the action offilamentous boring bacteria; these dissolve the carbonate
of the substrate that then precipitates around the bacterial filaments (Lucas and
Montenat 1967).
The genus, which occurs from the Carboniferous to the Pliocene, is mainly
present in the Palaeogene. According to Lucas and Montenat (1967) and Smit
(1979) many literature reports should be checked and Paronipora, which occurs
prior to the end of the Cretaceous, would probably not be actually recognized
up to the Recent, nor would the colonies have developed in a marine envi-
ronment. The capacity that it has for penetrating underground, through joints,
cracks and diaclases, could explain the occurrences in pre-Cenozoic rocks (Smit
1979) and also reflects its terrestrial origin.
Palaeomicrocodium Mamet and Roux, from the Devonian of Australia and
Canada is, according to Mamet and Roux (1983) and Mamet et a1. (1987), a
marine genus similar to Paronipora. According to Cherchi and Schroeder (1988)
Palaeomicrocodium is a junior synonym of Baccanella Pantie ofthe Alpine Trias
reef facies.
Bacinella RadoiCie and Lithocodium Elliott, according to some authors
(Segonzac and Marin 1972), represent two different parts of the same thallus;
respectively the older and the younger.
The two genera range from Upper Trias to the Cretaceous and flourished
mainly in the external parts ofthe lagoons and in the shelf margin facies (Conrad
1977; Fltige11979; Peybernes 1979; Senowbari-Daryan and Schafer 1979). The
majority of authors consider these two genera as incertae sedis; but Elliott (1955)
and some other authors (e.g. Johnson 1969 and Dragastan 1980) have assigned
Lithocodium to the Codiaceae; Bacinella has been referred to cyanophyceans by
Johnson (1969).
The genus Tubiphytes Maslov is known from Carboniferous to Lower
Cretaceous margin facies: see Maslov (1956a), Crescenti (1969), Dragastan
(1969, Micro-oncholithes), FlUgel (1981), Bernier and Gaillard (1974) and
Barattolo and Pugliese (1987). According to Maslov (1956a) Tubiphytes could be
attributed to the incrustations offoreign bodies by thin calcified cyanophyceans.
Elliott (1962) also noted that Tubiphytes develops around other bodies. In the
Mesozoic of the Apennines, as Crescenti (1969) has pointed out, Tubiphytes (PI.
L 4-5) does not develop around foreign material, but contains a tubular cavity
with periodical narrowings. Similar observations had been made by Dragastan
(1969) for the "Micro-oncholithes sous type Nubecularioidal"; according to this
author it is possible that these structures originate from the interrelations among
Nubecularia-like foraminifers, cyanophyceans and sediment. According to
Bernier (1984), Tuhiphytes morronensis Crescenti, and probably also the genus
Tubiphytes, is a foraminifer (Nubeculariidae) in which the thick outer wall was
an integral part of the porcellanaceous test and therefore was not due to an
oncoidal envelope.
The genus Tubiphytes, as Nigriporella Rigby, was considered a hydrozoan by
Rigby (1958). For further information see Crescenti (1969), Bernier (1984) and
Roux (1985).
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Whybrow PJ, Mc Clure HA, Elliott GF (1987) Miocene stratigraphy, geology and flora (Algae) of
eastern Saudi Arabia and the Ad Dabtiyah vertebrate locality. Bull Brit Mus Nat Hist Geol Ser
4,41 :371-382
Wray JL (I 977a) Late Paleozoic calcareous red algae. In: Fliigel E (ed) Fossil algae. Springer, Berlin
Wray JL (l977b) Calcareous algae. Elsevier, Amsterdam
Zorn H (1972) Bau und Funktion der Wirteliiste einiger triassischer Dasycladaceen (Kalkalgen). N
Jahrb Geol Palaontol Mouatsh Ser 12, 1972:727-740
Zorn H (1974) Variationstatistische Untersuchungen an Diplopora annulata (Schafh.) und D.
philosophi (Pia), Kalkagen der alpinen Mitteltrias. Eclog Geol Helv 67: 1-30
Zorn H (1976) Uber den Lebensraum fossiler Wirtelalgen in der Trias der Alpen. Naturwissens-
chaften 63:426-429
Zorn H (l977a) Statistical analysis of fossil algae. In: Fliigel E (ed) Fossil algae. Springer, Berlin
Zorn H (l977b) Der Einfluss der Umwelt auf die Variation fossiler Arten. Nat Mus Ser 1,107: 1-5
Chapter 23
Modern Stromatolites: A Review
S. GOLUBIC 1
Abstract
Modern stromatolites comprise a wide spectrum of various, laminated and non laminated organo-
sedimentary structures that form in various marine and freshwater habitats. Some stromatolites
form under extremely harsh ecological conditions, such as those found in hypersaline and
ephemeral ponds, or in thermal springs, others form in environments with normal marine salinity
or in freshwater. Microbial communities of different species composition participate in their
formation, lithification and destruction, often leaving marks of their particular activities.
1 Introduction
Historically, the main driving force for studies of modern stromatolites was the
need for interpretation of ancient stromatolites, particularly those found
abundantly in pre-Phanerozoic strata. The discovery of Precambrian microbial
fossils, and their association with stromatolitic structures stimulated the interest
in modern stromatolitic microbiota.
Understandably, there was no shortage of pertinent questions. Are
stromatolites shaped by environmental forces, or by the genetically determined
properties of the organisms that built them? What is the significance of their
lamination; does it record solar or lunar cycles, or perhaps less regularly
recurring events such as storms? Can they serve as a measure of ancient
astronomic events, or palaeolatitudes? Can they be used in biostratigraphy?
How do stromatolites trap and bind sediment, and how are they lithified and
preserved? Do they form in intertidal or subtidal environments? What limits
their growth and accretion? Which forces or organisms contribute to stroma-
tolite construction, and which to their destruction?
The hope was that many of these questions could be answered by studying
modern stromatolites, but some disappointment about the slowness of the
progress in this research area has been recently expressed (Burne 1986). An
important, often overlooked reason, that the research on modern stromatolites
did not fulfil expectations is that simple answers were expected regarding
exceedingly complex situations. The euphoria accompanying early discoveries
sometimes led to overgeneralizations and simplistic models that often persisted
and hindered further analytical work.
I Department of Biology. Boston University, 2 Cummington St., Boston, MA 02215, USA

542 S.Golubic
Modern stromatolite studies concentrated on extreme environments in

which microbial ecosystems dominate today, more or less unimpaired and
unmasked by the presence of macroscopic plants and animals. They provided
valuable information on microbial ecology. However, modern counterparts for
most of the ancient stromatolitic situations have not been found. Conversely, a
number of modern stromatolite settings lack fossil counterparts. Are these
stromatolites relatively new, or are they old ones that do not preserve? How
relevant are they then for the interpretation of fossil stromatolites?
Over the past three decades, modern stromatolites and microbial mats were
discovered in various environments, ranging from hypersaline to freshwater,
and from tropical to polar. They attracted the attention of researchers from
different disciplines, such as sedimentology, geochemistry, limnology and
microbial ecology, and became the focal point of an interdisciplinary research
effort. Awramik (1976) and co-workers (Awramik et al. 1976, 1979) compiled an
extensive list of references (which, incidentally, needs another update).
As a result of these studies, many overgeneralizations could be corrected,
definitions refined, and the scope of stromatolite-related phenomena and
environments within which they occur significantly widened. The insight into
structural, and particularly, functional complexity of modern stromatolites
highlighted the importance of the role of microbes in promotion and regulation
of biogeochemical cycling of elements in modern ecosystems. The principles
learned in the process are probably the best contribution which can be applied
to our understanding of fossil stromatolites and their environments.
When different scientific disciplines meet to form an interdisciplinary
approach to a problem, each discipline carries its own separate traditions,
concerns, and historic biases. Thus, misunderstandings and differences in the
use of terminology are to be expected. It is surprising, however, how deeply
rooted some traditional practices have become in the course of establishment
and development of various schools of thought, and how difficult it is to find
enough open minded ness and cooperative spirit to reconcile these differences
and arrive to reasonable new conventions. The field of stromatolite research is
no exception to this experience (see Monty 1982 vs Krumbein 1982, 1983).
2 Definitions and Terminology
The term 'stromatolites' entered the literature with full recognition of their
organo-sedimentary nature, together with an interpretation of their genesis
(Kalkowsky 1908). This view is shared by most researchers today, although
some prefer to use the term in a strictly descriptive sense, meaning a laminated
rock. The latter approach encounters problems when dealing with unrelated
laminated structures that are only superficially similar to stromatolites (e.g.
geyserite, calcrete, varved lake sediments, Liesegang rings). To study a modern
or living stromatolite means to study a stromatolite in the process of its
formation. Stromatolite lithification occurs as a part of stromatolite genesis or
diagenesis. As with other fossilized structures, lithification may help preser-
Modern Stromatolites: A Review 543
vation, but it is neither a requirement nor a warranty for it. Therefore, the term
'potential stromatolites' when used for mats and non-lithified stromatolites
(Krumbein 1983), may be misleading, because it raises expectations that may
not be fulfilled. Preservation potential (see, for example, Park 1977), is a
probabilistic measure analogous to life expectancy calculated by insurance
companies. Early lithification increases the expectancy of preservation.
The portion of the stromatolite in which the living community of stroma-
tolite-building microorganisms operates, is often spread as a thin, soft mat over
the structure's surface. Microbial activity in this mat may result in accretion of
the underlying sediment or rock. The stromatolitic structure then becomes a
cumulative record of an upwardly displaced microbial mat over time (Hofmann
1969). The lower portion of the mat may be 'dead', relating to the living
community as the wood tissue relates to the cambium (or the foliage). Thus the
term 'Recent stromatolites' includes both living and fossil stromatolites of the
Holocene age (see Monty 1982).
When first found, fossil stromatolites were treated as organisms and named
according to biosystematic practice using Linnean binomial nomenclature: e.g.
Cryptozoon proli[erum Hall 1884 and Collenia undosa Walcott 1914. This
traditional classification practice provides reference points, which can be hie-
rarchically organized into a system, or used in stratigraphic and facies cor-
relations. Since there are no skeletal fossils in pre-Phanerozoic rocks, a strati-
graphic subdivision of these strata based on stromatolites has been introduced
(Krylov 1963; Raaben 1969; Semikhatov 1976). For this purpose, Russian
geologists developed an elaborate methodology to study three-dimensional
morphology of fossil stromatolites, described a number of form-genera and
form-species, and named them according to Linnean nomenclature. This
approach was followed by Australian, French and, with some reluctance, also
by American researchers.
There was no problem in using Linnean nomenclature for shapes, as long
as modern and fossil stromatolites were treated separately. Krylov (1976)
argued convincingly that the Linnean binomial nomenclature has been histo-
rically applied to different unrelated items such as parts of the human body,
chemical compounds and drugs, plant extractions, and entire communities, so
why not for stromatolites? However, once modern stromatolites were ap-
proached by a combined team of palaeontologists and microbiologists, there
was difficulty in reconciling the usage of Linnean nomenclature for both the
structure and the organism that built it (Walter et al. 1976). In this particular
example from the Yellowstone thermal springs, the cyanobacterium Phor-
midium tenue var. granuli[erum was found to be the main constructive element
of the stromatolite structure Conophyton weedii, whereas Phormidium trunca-
tum var. thermale built the fiat-topped stromatolite Vacerrilla walcottii. Such
confusion in dual naming could be resolved by new conventions, e.g. by
restricting the use of italics to organismal names only.
544 s. Golubic
3 What Controls Stromatolite Shape?
The use of stromatolites in biostratigraphy is based on the assumption that

biological, genetically controlled properties of (mostly unknown) ancient
stromatolite-builders determined the morphological features recognized in
fossil stromatolites. Consequently, evolutionary changes and/or extinctions of
taxa would be reflected in the record of changing stromatolite morphologies.
The theoretical basis of this approach was explained by Maslov (1960).
This view was challenged by Logan et al. (1964). According to their view,
various stromatolite shapes are produced by a 'generic' microbial mat, which is
shaped differently by environmental forces, such as desiccation, waves and
currents. Their assumption was that stromatolites are built by entire microbial
communities, thus properties of individual organisms could not be expressed in
the shape of the resulting structure.
Successful application of stromatolites in stratigraphic and interbasinal
correlations (Walter 1972) supported the 'biological' view, while determination
of palaeocurrents in Precambrian shallow water facies (Hoffman 1967) sup-
ported the 'environmental' view.
A conclusive answer has been sought in studying modern stromatolites. The
evidence that has accumulated from these studies over several decades, shows
that a combination of biological and environmental factors is responsible for
stromatolite shaping. The relative importance of these factors for the resultant
shape, however, is proportioned differently from case to case.
Distribution of organisms in a community, microbial communities
notwithstanding, is not even, but rather log-normal. This observation has been
formulated by MacArthur (1960) as the 'broken stick' theorem. Accordingly,
every community is dominated by only few species which leave also the highest
impact in the community's structure and function. As one approaches extreme
environments, the total number of taxa decreases, and the discrepancy between
the dominant and sub dominant species increases (further breaking the 'stick').
Lowering the number of species, and discrepancy in their distribution, both
express a decrease in diversity. Thus, it is explained why a microbial community
may be shaped by the 'biological' properties of its dominant members. More-
over, some modern stromatolitic structures are shaped by virtually monospecific
populations. Good examples are Phormidium hendersonii (Golubic and Focke
1978), Entophysalis major (Golubic 1976a) and Rivularia haematites (Golubic
1976b; Schneider 1977).
On the other hand, in high energy environments it becomes evident that
wave scouring and currents contribute directly to the trimming and shaping of
stromatolites (Plate II, 11; Logan 1961). Less apparent is the fact that each taxon
has different, genetically determined environmental requirements, and re-
sponds accordingly to environmental changes. Under extreme conditions ofthe
intertidal zone, for example, this response is reflected in zonal distribution of
taxa. In the case ofthe Yellowstone stromatolite study (Walter et al. 1976) it was
the environmental change that caused the shift in microbial composition and
dominance from one to the other species of Phormidium, whereas the inherently
different growth habits of these two taxa resulted in the change of the
stromatolite shape.
Gebelein (1974) expressed the fact that stromatolites are mostly built by
microscopic organisms by formulating the generalization, that the stromatolites
micromorphology is largely 'biological', while their macro morphology is
'environmental' in origin. The studies of the microenvironments within which
the stromatolite growth takes place (e.g. Cohen et al. 1977; Krumbein et al. 1977)
provided further refinement, indicating that complex interactions between
organisms and environmental determinants take place at the microscopic as
well as macroscopic scale.
We may conclude (1) that genetically determined properties of dominant
micro-organisms, including their specific growth habits and behaviour (e.g
phototropism and phototaxis) are often revealed in stromatolite morphology. A
good example is provided in the genesis of conical stromatolite shapes, where
analogous relationships between micro- and macromorphology was found in
modern and ancient stroma tolites (Awramik and Vanyo 1986); (2) that beca use
of specifically different environmental requirements of microorganisms, their
distribution and dominance depends on environmental conditions and re-
straints, and (3) within these restraints, the environmental forces can influence
stromatolite morphology directly, by trimming and shaping form analogous to
the way trees and shrubs are trimmed by the wind in the mountains and along
the coasts.
4 What Is Lamination?
The word stromatolite implies layering. Yet, as we learn more about modern and
fossil stromatolites, we find (1) that lamination is a property which is differently
expressed, ranging from fine and regular to coarse, to hardly noticeable or
non-existent, and (2) that following the genesis of a stromatolite we need to
distinguish between community stratification and structure lamination, both
resulting in distinct layers.
The living stromatolite-building microbial community is often stratified.
Layers dominated by different organisms that arrange themselves in strata of
optimal growth, follow a vertical gradient oflight, O2 , Eh, pH and concentra-
tions of dissolved minerals. Such a gradient, orchemocline, becomes established
within the interstitial waters as a result of microbial metabolic activities in a
homeostatic fashion. Thus, the mat is microenvirorunentally and biologically
differentiated into strata, analogous to lake stratification, but with each stratum
only a fraction of a millimetre in thickness (e.g. Guerrero and Mas 1989).
The primary producers ofthis community, usually cyanobacteria and algae,
form the top layer. Analogous to the plankton in the epilimnion oflakes, they
may show an inhibition of photosynthetic output at the very top of the mat due
to excessive light. The organisms may respond either by elevated intracellular
carotenoid production, as protection against photo-oxidation, or by production
of extracellular shielding pigments within their polysaccharide sheaths, or both.
546 S. Golubic
The highest output of organic matter production by oxygenic photosynthesis is

observed in the subsurface layer, which is also supersaturated with respect to
oxygen, and usually blue-green or green in colour. The light inhibition is
reduced, or eliminated, in deeper water or in shaded places (caves).
The density of the surface layers acts as a light filter, so that self-shading
becomes effective deeper within the mat. The initial reaction of cyanobacteria
to light attenuation is a compensative increase in pigment production, par-
ticularly ofthe red light-gathering pigment phycoerythrin (Jorgensen 1989). An
abrupt drop in primary production follows, accompanied by intensive aerobic
decomposition of organic matter (primary product) which, in turn, removes the
ambient oxygen, forming an anaerobic layer beneath the green portion of the
mat. Release ofH 2 S by bacterial protein decomposition and, more importantly,
by S04 reduction through the activity of anaerobic respirers, fuels the non-
oxygenic bacterial photosynthesis. This photosynthesis uses light of longer
wavelengths (700-900 nm) that filters through the cyanobacterial layers. Some
cyanobacteria show the ability to switch between oxygenic and non-oxygenic
mode of photosynthesis (see Cohen 1989). A layer of purple sulfur bacteria
usually follows the blue-green one. Filamentous chlorobacteria of the
Chloroflexus-type are often intermingled, providing mechanical support and
modifying the colouration of this zone. The layers beneath are in darkness,
dominated by fermenting decomposers, and possibly methanogens. Excessive
H 2 S is bound in a black FeS precipitate, which may diagenetically recrystallize
into pyrite.
The conditions inside microbial mats vary diurnally and seasonally, in fact
the chemocline often shifts up and down in the same rhythm. Microorganisms
may respond either by vertical migration (Golubic 1973a) or by physiological
adjustments (de Wit 1989).
The described stratification is typical of many water-logged intertidal mats.
Such mats line the channels, ponds and mud fiats of Abu-Dhabi, (UAE Plate
11,9,10), of protected embayments in Shark Bay, Australia, and of lagoons of
Baja California, Mexico, and can be considered cosmopolitan in distribution.
Intertidal sediments with better drainage are covered with a thinner mat in
which oxygen depletion occurs transiently, restricting the development of
anaerobic microorganisms. Decomposition by the more efficient aerobic re-
spiration is usually able to recycle all of the organic matter produced, which
keeps the mat thin. Change in drainage regime may trigger a microorganismal
succession, promoting colonization by heliotropic cyanobacteria (Awramik and
Vanyo 1986). In Abu-Dhabi fiats this change results in formation of upright
pinnacles formed by the cyanobacterium Schizothrix splendida (Plate II, 10;
Golubic 1973a,b).
In perpendicular sections through a mat, the microbial stratification is
conspicuous as coloured bands, however, this layering is not the predecessor of
lithified stromatolite laminae. As the mat accretes due to sediment trapment
(Plate I, I), or mineral precipitation (Plate 1,2), members of the community escape
by either growing or moving upward. An upward displacement usually starts
with the primary producers of the community. and is then followed by a cohort
of consumers, degraders, and anaerobic photobacteria. Their activities, in turn,

causes an upward displacement of the entire chemocline. If this progression
proceeds at an even rate, the resulting stromatolitic structure, i.e. the cumulative
record, will also appear uniform and smooth. Only recurring disturbances in the
process of stromatolite accretion are recorded as laminae. The underlying
causes behind a sequence of alternating laminae, typical of most stromatolites
may be quite different. In the case of Phormidium hendersonii, diurnal alter-
nation in light was sufficient to change filament orientation and cause lami-
nation (Monty 1965; Golubic and Focke 1978).
The thickness of organic-rich laminae may depend on the timing of organic
production within the cycle, but may also reflect the relationship between
organic production and decomposition. Observations on intertidal mats of Abu
Dhabi show that decomposition rates are more sensitive to drainage patterns
than production rates are. Water shortage, on the other hand, is likely to affect
production rates directly, while decomposition continues for a while unim-
paired protected by the overlying mat. The thickness of organic-poor stroma-
tolitic laminae may depend on sediment supply. Stromatolites that trap
sediment particles carried by tidal currents (Gebelein and Hoffman 1968), may
record lunar cyclicity. Storm-associated sediment transport is often reflected in
organic-poor laminae of varying thickness. If storms cluster in a seasonal
pattern, such laminae may record a solar cyclicity.
With so many interacting sources of lamination, is it reasonable to expect
that palaeoastronomic interpretations could be reliably made from stromatolite
lamination patterns (Pannella et al. 1968; Awramik and Vanyo 1986)? This may
be so, providing there is some limitation in the number of irregular disturbances
in the palaeoenvironment. Subtidal stromatolites growing continuously in
relatively protected and calm environments should qualify as the best potential
models for such studies. However, this hypothesis is difficult to test, as no such
model has been found in modern environments (Phormidium hendersonii has a
short season of about 2 months).
5 Preservation and Lithification of Modern Stromatolites
Living microbial mats are metabolically active ecosystems which effectively

recycle nutrients. Aerobic decomposition of organic matter is almost complete,
lea ving very little contribution for the fossil record. Under anaerobic conditions
(e.g. in water-logged mats) complete decomposition of organic matter is possible
(Doe mel and Brock 1977), but it involves a succession of several metabolically
different groups of microbes. Organic matter is more likely to be preserved and
fossilized anaerobically, because the required multimembered, slower-growing
set of anaerobic degraders is, statistically speaking, less likely to congregate,
diminishing the potential for complete removal. Sedimentary components of
the mat, unless consolidated, may easily fall victim to erosion or pressure
distortion. The best chance for stromatolite preservation (by no means a
requirement) is achieved through early lithification. Most commonly, this
548 S. Golubic
Plate I
involves cementation by calcium carbonate. However, early lithification is not

conducive to preservation of microbial fossils, which are best preserved in
silicified sediments (see Knoll1989). (Embedding in silica, iron or manganese
occurs under special conditions and will not be discussed here.)
Biogenic destruction of lithified stromatolite surfaces by endolithic
microorganisms that leave specific traces (microborings) has been demon-
strated in marine (Golubic 1983) and freshwater (Anagnostidis and Pantazidou
1988; Winsborough et al. 1990) environments (Plate I, 7, 8). Microbial en-
doliths are known from pre-Phanerozoic oolites (Campbell 1982; Knoll et al.
1986), as well as stromatolites (Zhang and Golubic 1987).
6 Abiotic and Biogenic Factors in Calcium Carbonate Deposition
Carbonate deposition is often an integral part of stromatolite formation. The

role of microbial activity in this process is not easily determined and remains
controversial. In the process of calcification the carbonate ion passes through
several phases, starting from an aqueous solution and ending as calcium
carbonate within a limestone rock. The contribution of microorganisms in each
of these phases needs to be assessed separately.
In its initial phase, the process of carbonate precipitation belongs to the
realm of solution chemistry. The solubility of different chemical species of the
calcium carbonate buffer system in natural waters is under constraints of
temperature, pH and overall ionic composition. The solution can reach states of
saturation and supersaturation, which may lead to precipitation ofCaC0 3 from
solution. This process can be influenced by abiotic factors such as evaporation.
It may also be influenced by biotic factors such as photosynthetic removal of
CO 2 and HCO:i, or heterotrophic ammonia release and sulfate reduction, all of
these being associated with pH increase.
Following chemical precipitation from solution, carbonate molecules or-
ganize, following a crystal lattice. This process belongs to the realm of mineral
Plate I. 1 Aragonite particles trapped on the surface of the polysaccharide sheath of the marine
cyanobacterium Microcoleus chthonoplastes. Florida Keys. Nomarski Interference contrast; scale
bar is 10 fJ.m. 2 Calcite precipitated on the surface of the polysaccharide sheath of the freshwater
cyanobacterium Scytonema julianum. Nomarski Interference contrast; same scale as in 1.
3 Species-specific calcite skeleton formed by the desmid Oocardium stratum. The entire cell
population formed a single calcite monocrystal. The cells are removed in this preparation using
sodium hypochlorite (Clorox). SEM; scale bar is 100 fJ.m. 4 Detail from 3; scale bar is 20 fJ.m.
5 Calcitic tubules formed within the sheath of the cyanobacterium Geitleria calcarea. SEM; scale bar
is 20 fJ.m. 6 Detail from Fig. 5 showing the monocrystalline structure and orientation of calcite; scale
bar is I fJ.m. 7 Endolithic filaments ofthe calcium carbonate penetrating freshwater cyanobacterium
in stromatolites of the EI Mojarral, Cuatro Cienegas, Mexico, 3 m depth. Arrow indicates a smaller
endolith Plectonema sp. Nomarski interference contrast; scale same as in 8. 8 Resin casts of endoliths
microborings produced by the cyanobacterium Plectonema sp. Same sample as in 7. SEM; scale bar
is 10fJ.m
550 S. Golubic
formation and is commonly referred to as 'nucleation'. It may depend on

inorganic factors such as temperature or the presence of other ions or colloids,
that serve as potential nucleation sites that compete for carbonate molecules.
Biogenic influences on the nucleation process may involve structural
macromolecules (e.g. polysaccharides and polypeptides) or metabolic products
that come into contact with the carbonate ions. They may promote or retard
nucleation. Nucleation and crystal growth occurs either in the water column,
forming a suspension (sometimes referred to as a 'whiting'), or at the
sediment-water interface.
Ultimately, all the precipitated carbonate settles and contributes to the
formation of sediments. The rules that are followed thereafter belong to the
realm of sedimentology. This phase includes depositional and diagenetic
processes in which prominent a biotic factors include current and wave energies,
and pore water chemistry, respectively. Biogenic modifications include trapping
and binding of particles, bioturbation, as well as release of metabolites.
The abiotic and bIOgenic influences may be qualitatively different in each
of the sequential phases of carbonate deposition, and their relative importance
may shift from phase to phase. Keeping them conceptually separate as
'chemical', 'mineralogical' and 'sedimentological', may help to resolve some
controversies that persist in the literature merely because the disagreeing parties
are referring to different parts of a multifaceted, complex process.
7 Degree of Biological Control
One can view the cell as a special type of microenvironment in which the
processes are subject to tight biological control, principally through biocatalytic
action of enzymes. By extension, in multicellular organisms, tissues and inter-
cellular spaces comprise microenvironments where the combined activity of
many cells exercises its influence. A decreased degree of biological control is
expected outside the organism, within its immediate surrounding that comprises
its microenvironment. The degree of biotic influence here will depend on the size
of this microenvironment in relation to the size and activity of the organisms
within it, as well as on the degree of seclusion of this microenvironment from the
environment at large - the macroenvironment. It is increasingly becoming
recognized that there is no environment on Earth that has not been modified by
biotic processes throughout the geological past (e.g. Westbroek 1983). We have
just grown accustomed to taking the present state of the environment as a given
background, against which we measure the influences of organisms that are
presently active.
Mineral deposition under close biological control also follows the organ-
ization of biological structures (templates) resulting in species-specific skele-
tons. A comprehensive comparative treatment of biomineralization processes
has been recently published by Lowenstam and Weiner (1989). Eukaryotes have
produced. and left in the fossil record. a wide variety of skeletons and skeletal
fragments. Mineral deposits associated with prokaryotic organisms are usually
not considered to be skeletal. However, clear distinction between skeletal and

non-skeletal mineral deposits is not always possible, because the degree of
biological control varies from phase to phase of the process, and from taxon to
taxon in eukaryotes and prokaryotes alike. Similarly, the distinction between
biologically induced and biologically controlled mineralization processes is a
conceptual simplification, whereas in the real world there is a wide spectrum of
possibilities with different degrees of biological vs environmental control
(Lowenstam and Weiner 1989).
A common characteristic of living systems is their ability to act against
chemical gradients. This is exemplified at the cellular level by the active
transport of nutrients across the plasma membrane, and at the environmental
level by the biogenic polarization of reduced and oxidized microenvironments
in a mat. Thus, in assessing the biogenicity of mineral precipitation in a complex
natural situation, it is good strategy to study situations with known environ-
mental 'resistance' to the process, against which the biological impact is well
contrasted. The opposite is true for situations where both abiotic and biotic
conditions favour a process, and it is difficult to separate their respective
contributions. This may explain some controversies accompanying the assess-
ment of the organismal impact on exergonic oxidation processes (e.g. oxidation
of iron and manganese) along redox interfaces or, for that matter, on the
precipitation of calcium carbonate from heavily supersaturated waters.
Marine waters of normal salinity, although often chemically supersaturated
with respect to calcium carbonate, show a strong resistance to spontaneous
precipitation. Calcification is therefore largely under direct organismal control
resulting in production of various skeletons. The prevalent crystallization form
ofCaC03 in sea water is aragonite, biogenic calcification, however, frequently
takes the mineral form of calcite. (There is a possibility, however, that at
different times of the Earth's history the overall ocean chemistry may have been
different, and more conducive to spontaneous precipitation of calcite rather
than aragonite). Low Mg-calcite precipitation is today prevalent in nonmarine
waters of normal temperatures, however, rapid precipitation from hot springs is
often aragonitic. The following examples illustrate different degrees of
biological impact on the calcification process.
In eukaryotes, calcitic coccoliths of species-specific morphology are ex-
amples of marine intracellular calcification (Westbroek et al. 1984). Mg-calcite
is deposited in the cell walls of crustose rhodophytes in a microenvironment that
is technically extracellular (outside plasma membrane), but in intimate contact
with cellular tissues. Open intracellular spaces in marine siphonaceous chlo-
rophytes are filled with aragonitic encrustations, and show correspondingly less
cellular control over the calcification process, although the initial precipitation
seems to be guided by biogenic templates (Borowitzka 1984).
In prokaryotes, biogenic carbonate encrustations of species-specific mor-
phology is found in two marine cyanobacteria Rivularia mesenterica and R.
polyotis. It takes place in an environment in which spontaneous carbonate
precipitation is not known to occur (Adriatic Sea). Calcium carbonate is
deposited outside cells, but within an extracellular polysaccharide gel. The
552 S. Golubic
shapes of the concretion in the two species differ markedly, indicating

specifically different modes of nucleation templating. The mineral form of the
precipitate is aragonite in both cases, although the initial deposition appears
amorphous (Golubic and Campbelll981).
Spontaneous precipitation of calcium carbonate from supersaturated
freshwater occurs readily, making determination of biogenic control of the
process more difficult. Critical differences between calcifying and non-cal-
cifying taxa, however, can be observed in waters close to equilibrium or slightly
undersaturated with respect to calcium carbonate. Under such conditions,
within the same microenvironment, the cyanobacterium Phormidium incrus-
tatum is heavily calcified while Ph. autumnale is not (Golubic and Fischer 1975).
The fact that both species are found calcified in highly supersaturated waters of
the Plitvice waterfalls (Yugoslavia) masks this distinction. Similarly the cyano-
bacterium Scytonema julianum is consistently covered with a coat of charac-
teristic CaCOa crystals (Plate I, 2; see Coute 1985), while S. myochrous remains
completely uncalcified. Species-specific templating and imprinting of charac-
teristic biogenic patterns over those determined by inorganic mineral lattices is
evident in many freshwater stromatolite-building organisms, both prokaryotic
and eukaryotic. Biologically modified crystals occur in the cyanobacteria
Rivularia haematites (Schneider 1977), Schizothrix fasciculata, Hydrocoleum
homoeotrichum, and Geitleria calcarea (Plate I, 5, 6; Friedmann 1955; Coute
1985), in the filamentous chlorophyte Gongrosira incrustans (Golubic and
Fischer 1975), and desmid Oocardium stratum (Plate I, 3, 4). Carbonate
deposition on gelatinous stalks of benthic diatoms also carries biologically
specific marks. It takes place at a certain distance from the cells indicating that
it is the nucleation of calcium carbonate crystals on the polysaccharide cell
product, rather than metabolic activity (photosynthesis) of the cell, that directly
affects calcification (Winsborough and Golubic 1987).
The crystals in all these taxa conform to some degree with the miner-
alogically expected rhombohedral form. However, they are deposited tightly
around templating cyanobacterial or algal products and bear their species-
specific imprints. In this sense they can be considered skeletal (Riding 1977).
Experimental evidence from cultures shows that heterotrophic bacteria
engage in activities that result in an increase in pH (e.g. sulfate reduction or
ammonia release), and can cause precipitation of carbonates in their vicinity.
Bacterial precipitation of calcium carbonate in nature has also been reported
(Krumbein 1974). The evidence is based on findings of carbonate-encased
bacteria in the hypersaline environment of Solar Lake, Sinai. Under the
circumstances of critical supersaturation, the actual role of these bacteria in the
calcifica tion process is not clear. Their role may ha ve been meta bolic, promoting
selective nucleation of chemically precipitated minerals, or entirely passive in
the sence of serving as incidental surfaces.
The experimental evidence that particular bacteria in pure culture have the
capacity to initiate mineral precipitation does not necessarily explain the origin
of the minerals found in natural environments, where processes may act
synergistically or in opposition to each other. For example, under natural
conditions, organic matter is decomposed via fermentative and respirative

processes in which H 2 , CO 2 , and organic acids are released, generally lowering
the environmental pH. Some processes, such as deamination and sulfate
reduction act in the opposite direction, elevating pH. The balance between these
processes and their microenvironmental distribution need to be known before
the ultimate effect of bacteria on carbonate dissolution or precipitation in
particular cases can be assessed.
8 Are Stromatolites Intertidal, Subtidal or Both?
Important information about ancient stromatolites, which is expected from the

study of their modern counterparts, entails assessment of the conditions under
which they form, in order to facilitate their use in reconstruction of palaeoen-
vironments. Again, we need to resort to particular case histories. The study of
modern stromatolites of Shark Bay, Australia has provided most of our current
knowledge, but has also spawned serious misconceptions, which now need to be
sorted out. Logan (1961) described several important features of one type of
intertidal stromatolite (which we now know to be formed by the cyanobacterium
Entophysalis major, see Golubic 1973a, 1983; Golubic and Hofmann 1976).
Logan established the wave exposure-dependence of their domal morphology,
assessing the role of erosion and wave scouring on their shape and directional
orientation. By extrapolation from these observations he concluded that all
stromatolites, modern and fossil, must be intertidal in origin and are shaped by
environmental forces prevalent there. The presence of modern subtidal
stromatolites in Hamelin Pool were dismissed by Logan as relics of intertidal
stromatolites of earlier times when water levels were lower than today.
On the other extreme, some 20 years later, Burne and James (1986) claim
that subtidal ranges of the Hamelin Pool are the real home of stromatolitic
structures. They explain today's intertidal stromatolites as being relics of earlier
higher water levels, stranded by the relative fall in sea level, and recolonized by
intertidal microbial mats. Which stromatolites are actively accreting today and
which can be considered fossil? Marking of subtidal stromatolites with stainless
steel nails and following the subsequent increments showed that their accretion
is patchy and slow (Playford and Cockbain 1976). In the intertidal zone, the
co-existence of growing Entophysalis mat and hard lithified surfaces, implies
similar irregularity of the process.
In order to test the validity of either of these postulates, the following
question needs to be answered for both subtidal and intertidal stromatolites:
Are microbial mats that currently colonize stromatolite surface contributing to
the accretion of the stromatolite upon which they grow, are they neutral to it, or
perhaps even contributing to stromatolite destruction?
Subtidal stromatolites are colonized by a distinct (colloform) mat com-
munity composed of cyanobacteria and algae, and dominated by a coccoid
cyanobacterium, new to science (Golubic 1985). The role of the eukaryotic
component of this community is discussed by Awramik and Riding (1988).
554 S.Golubic
Sediment particles are trapped and cemented by in situ carbonate precipitation.

The same mineral elements that occur within the active mat are also found
consolidated in the interior of the stromatolite. Thus, there is evidence for the
contribution of the current microbiota to stromatolite accretion, and no
evidence of any discontinuity in this process, that would separate the past,
vestigial structures and processes from those occurring today.
Intertidal stromatolites are colonized by the mamillate (pustular) mat, an
entirely different, much less diverse microbial community, which is dominated
by the coccoid cyanobacterium Entophysalis major. For the most part of the
year, this mat colonizes loose sediment and is shaped by wa ve action and erosion
(Plate II, 11). At critical times at the peak of the austral summer, precipitous
lithification occurs within the Entophysalis mat, destroying the microorganisms
in the process. Thus, there is evidence that the soft Entophysalis mat turns
periodically into hard stromatolite (Plate II, II, arrow), contributing
significantly to its accretion. In the intermittent times between two colonizations
by Entophysalis the stromatolites are attacked by carbonate-boring endolithic
microflora (Golubic 1983).
In addition to the two stromatolite types under discussion, several others,
which are built by different microorganisms, show different degrees of
lithification and occur in distinct zones. The shapes, densities and orientation of
subtidal, lower intertidal, and upper intertidal stromatolites are all different
(compare illustrations in Golubic 1985) and cannot be explained as 'inherited'
from one another as Logan (1961) or Burne and James (1986) suggested. As
compared with the predominantly large, round-based subtidal stromatolites
(Golubic 1985, PI. 1.1, Fig. 3), the lower intertidal ones are smaller and denser
(Golubic 1985, PI. 1.1, Fig. 6), whereas the upper intertidal ones are often
elongated, following the prevailing wind direction and wave scouring (Golubic
1985, PI. l.2, Fig. 2), an activity that the subtidal stromatolites are hardly
exposed to. We deal here with separate, but active ranges in which stromatolite
formation and lithification takes place, each formed by different micro biota and
lithified differently.
Among modem stromatolites some are intertidal, and others subtidal. In all
probability, the same should be true for their fossil counterparts. The case
Plate II. 9 Heliotropic pinnacles built by the cyanobacterium Schizothrix splendida occupying
slightly elevated points on top of a periodically waterlogged, Microcoleus chthonoplastes dominated
mat. Intertidal flats east of Abu Dhabi, Arabian Gulf(Photo courtesy ofR. Park). 10 Vertical section
through one of the heliotropic pinnacles in Fig. 9 showing continuity of pinnacle differentiation in
the process of mat accumulation (Photo courtesy of R. Park). 11 Domal stromatolites 'carved' by
wave erosion from a contiguous sediment platform (in the background). Sediments are stabilized by
the mamillate mat dominated by the coccoid cyanobacterium Entophysalis major. Arrow marks the
lithified portion of the stromatolite. Hamelin Pool, Shark Bay, Western Australia. 12 Large subtidal
stromatolites built by trapping and binding of oolitic sands, Exuma Island, Bahamas (Photo
courtesy of S.M. Awramik). 13 Freshwater domal stromatolites in Pozo Azul. Cuatro Cienegas,
Mexico
Modern Stroma tolites: A Review 555
Plate II
556 S. Golubic
history of Shark Bay teaches us, among other things, that simple sweeping
generalizations, no matter how attractive, are unlikely to be usefuJ as inter-
pretation models.
9 Are Modern Stromatolites Restricted to Extreme Environments?
Diversity of forms, vast distribution ranges and absolute dominance of

stromatolites in the pre-Phanerozoic have always intrigued researchers, par-
ticularly when contrasted with their relative paucity in the Phanerozoic. Aw-
ramik (1971) has shown the decline of stromatolites in the fossil record at the end
of the Proterozoic to coincide with the appearance of skeleton bearing organ-
isms. Garrett (1970) offered an explanation for this decline showing that in
modern marine environments the formation of microbial mats, as precursors of
stromatolites is effectively prevented by the grazing pressure of invertebrates.
Monty (1973) later extended this view, by including competition with newly
evolving reef-forming organisms. These may have caused displacement of
stromatolites from normal marine environments into the less favorable inter-
tidal zones and into freshwaters.
Intertidal ranges and ephemeral pools within them, with their fluctuating
water supply and chemistry, particularly in arid climates certainly qualify to be
called extreme environments. The studies of such environments (e.g. Kinsman
and Park 1976; Park 1976,1977; Horodyski et al. 1977; Krumbein et al. 1977;
Campbell and Golubic 1985) show organismal distribution patterns that are
consistent with Garrett's model, and confirm an almost absolute dominance of
prokaryotes. Similarly, extreme conditions in the non-marine realm are found
in thermal springs (Brock 1978; Castenholz 1973; Ward et al. 1989), and
somewhat less extreme in the Antarctic lakes (Parker et al. 1981; Parker and
Wharton 1985). However, some modern stromatolites persist in normal marine
environments as well as in nonmarine environments with less extreme condi-
tions (Moore et al. 1984; Dill et al. 1986). In addition, fossil Phanerozoic
stromatolites are often found surrounded by normal marine floras and faunas
(see Monty 1981). These findings testify that in many environments stroma-
tolites did survive competition and accommodated animal grazing, in an
apparent contradiction to the Garrett's dictum, and require a different
explanation.
Although the research of the Exuma stromatolites (Plate II, 12; Dill et al.
1986) is still in its initial stages, several peculiar properties of this environment
are known. They grow under a regime of strong tidal currents that carry loads
of ooidal sands. The key to their accretion is the ability of stromatolite building
microorganisms to trap the sediments and escape burial. The key to their
persistence lies probably in rapid cementation and induration of these struc-
tures. Whether, and to what extent this process is facilitated by organisms
remains to be established.
Carbonate deposition in non-marine setting occurs predominantly as tufa
(cool water travertine), forming porous biohermal structures which show certain
stromatolitic properties such as lamination. They are usually associated with

intensive in situ precipitation of carbonate from springs and running waters that
are supersaturated with respect to CaC0 3 , although they also involve par-
ticipation of micro- and macroorganisms. Their early lithification seems re-
sponsible for their preservation and accretion (Golubic 1967, 1973a). Benthic
carbonate deposition in lakes has been known to result in formation of oncoids
(Irion and Milller 1968), and calcareous crusts, which usually do not exceed a
thickness of about 1 cm. Their accretion is counteracted by bacterial de-
gradation and corrosion (Golubic 1962), destruction by endoliths (Schneider
1977), probably combined with animal grazing and wave and, in cooler regions,
ice erosion (Kann 1941).
Lacustrine stromatolites in lakes with normal freshwater chemistry, are rare
and known only from warmer climates. In habitats where such lacustrine
stromatolites occur, they accommodate rich and diverse animals populations,
which include grazers, such as snails and fish (Winsborough et al. 1990).
Well-developed lacustrine stromatolites are found on the margins of lim-
nocrene springs, locally called 'pozos' in Cuatro Cienegas basin, Coahuila,
Mexico (Plate II, 13). They are domal in outline, with corrugated brain-like
surface configuration. The surface of these stromatolites is soft, and loaded with
trapped faecal pellets. Toward the interior, these stromatolites become
gradually harder due to cementation. They grow best in shallow water,
frequently coalescing into reef-like bioherms, that prograde from the shore
toward the centre of the 'pozo' creating platforms with overhangs. With in-
creasing depth, they show evidence of decreased accretion rate, accompanied by
increase in hardness, and in deepest ranges they show signs of corrosion (Plate
I, 7, 8).
10 Conclusions
Modern stromatolites offer an opportunity to study relationships between

microbiota and sedimentary structures within their ecological context, and to
follow their genesis. Actualistic models can be postulated and experimentally
tested. As such they can provide useful interpretational tools for fossil
stromatolites. However, a number of modern stromatolite settings are not
known from the fossil record and, conversely, most of the known ancient
stromatolites do not have close modern counterparts. Thus, the studies of
modern stromatolites rarely allow direct comparisons. They provide, however,
information of more general nature, about the principles and ecological con-
straints involved. We learn that the shapes of modern stromatolites may be
biologically controlled by growth habits of dominant stromatolite-building
microorganisms. Hence, there is a need for accurate taxonomic identification of
these builders. The environment may act on two levels: by selecting for or
against microorganisms with particular growth requirements, and by physically
trimming the shape of the resulting structure. These interactions determine the
distribution patterns of stromatolites. Like a coral reef, a stromatolite is a habitat
558 S. Golubic
as well as a growing structure. The role of organisms associated with it can

be constructive, neutral, or destructive. The balance of combined constructive
and destructive forces determines the stromatolite accretion rates and
preservability.
Acknowledgements. Barbara and Philip Winsborough are thanked for field guidance in Cuatro
Cienegas, and Jacob Seeler, Susan E. Campbell, and Stanley M. Awramik for critically reading the
manuscript and offering valuable suggestions. The work has been partially supported by the NSF
grants EAR 8306179, INT-8509365, and Boston University grant GRS 8021-BI.
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Index
Abacella 354 Bacillariophyta, see diatoms

Abatea 158 Bacinella 73-74,77,467,532
Acetabularia 10,128,131,138-139,141, bacteria 30, 351, 546, 552
142, 515, 529 calcification 25
Acicularia 141, 142, 523 Baculella 384
Aclistochara 197, 198 Baicalia 298
Acroporella 141 Batinevia 325
Aeolisaccus 506 Batophora 126, 134, 141, 142
Aethesolithon 509 Beltina 30
Agardhiellopsis 510 Belzungia 134
Alasphaera 272 Beresella 128, 379, 381, 384,461, 515
A lbertaporella 518 Berestovia 385
Amia 118 Bevocastria 76, 349, 352, 354, 382, 385,
Amgaella 306, 327 505, 514
Amorfia 375 Bianolithus 240
Ampullipora 356 Bicarinellum 274, 281
ancestral corallines 458 Bija 306, 307, 313, 325
Anchicodium 460, 511 biomineralization 3,22, 25, 41, 46, 213,
Angulocellularia 26, 35, 37, 60, 62, 64-65, 220, 305, 550- 553 (see also calcification)
76, 77, 78, 307, 313, 317, 319, 321, 323 Biscutum 231- 232, 233 - 234
Anthracoporella 376,436,461-462, 517 Biumbella 386
Anthracoporellopsis 376, 381 Borisovella 386
Aoujgalia 379 Bornetella 115, 120, 131, 132, 134, 529
Aphralysia 72, 381, 511 Botomaella 67, 76, 78, 310, 317,321,
Aphroditicodium 149, 464 325
Aphrostoma 382 Botominella 306, 310, 325
Apidium 115, 351 Botryella 356
Arabicodium 149, 150, 153, 492, 496, 511 Boueina 147,149,151,153,492,496,511,
Aranea 382 513
Archaeobatophora 126, 515 Braarudosphaera 219,233-234,249
Archaeolithophyllum 103, 381, 382, 458, Brazhnikovia 386
460 Broeckella 138, 142
Archaeolithoporella 452, 453, 459, 464, Broinsonia 233
470,471
Archaeolithothamnium 94-95, 110, 509 Cabrieropora 387
Archaeozygodiscus 231-232 Calcidiscus 219, 248
Arcticella 350 calcification 99, 278, 490, 549-550, 551
Ardengostella 383 (see also biomineralization)
Artophycus 383 algae 3, 10
Asphaltina 72, 365, 383 bacterial 25
Asphaltinella 383 Charophyta 11,204,209,213-215
Atopochara 193, 194 Chlorophyta 10-11
Atractyliopsis 381, 384, 461-462 Chrysophyta 14-15
coccolithophorid 11-13, 16, 220
Baccanella 532 Corallinaceae 99-103
564 Index.
calcification (cont.) oosporangium 189,204

cyanobacteria 4-10,42,56-58,74-78, reproduction 205
79,80-81 taxonomy 198-201, 204, 215
cyclocrinitid 118 Chiasmolithus 236, 242, 255
Dasycladales 141 Chondrostoma 388
diagenetic 79 Chrysophyta 14-15
diatoms 15 Cladophora 11
dinoflagellate 277 Claracrusta 388
Gymnocodiaceae 157 Clavapora 518
Halimedaceae 170 Clavaporella 379, 389
microbial 22, 45 Clavatoraceae 193
Phaeophyta 13 Clibeca 356
processes 4, 15, 22, 57 Clypeina 138, 141, 496, 523, 524
Rhodophyta 13 -14 Coactilum 389
and seawater chemistry 80-81 coccolith 12,217,221,226,232,252,258,
stromatolite 296, 547 - 549 551 (see also nannofossil)
Udoteaceae 150 diagenesis 247 - 248
Calciodinellaceae 268 dissolution 244-248, 252
Calciodinellum 272, 280 heterococcoliths 222
Calcifolium 379, 387, 437 holococcoliths 222, 245, 248
Calcisphaera 387 morphology 221- 223, 237
calcispheres 267,274-280,281, 365-366, placoliths 222, 232, 243, 248
373, 387, 425 sedimentation 226,229,244-248,
Calothrix 9, 62, 63 251-252
Cambrian 60, 75, 349 coccolithophorids 217-266 (see also
calcareous cyanobacteria and nannofossil)
algae 305 - 334 bathymetry 225, 248-249
ranges 329 biology 220-221
systematics 314-328 biostratigraphy 239-244
Cambroporella 306, 312, 327 calcification 11-13, 16, 220, 221,
Campbelliella 523, 524 237 - 239, 258
Capitan Reef Complex 452, 458, 461, 470, and climate 229, 253-260
473 ecology 225-229,236,248-260
Carboniferous 76, 335, 365, 437, 516 evolution 229-239
calcareous algae 370-451 freshwater 228
ranges 436-442 ingestion of 245
Carinellum 272 Coccolithus 11, 12, 16, 219, 227, 228, 237,
Carpenterella 134 247,248
Cayeuxia 60, 67, 77, 505 Codiaceae 147, 151, 152, 153, 154, 160,
Cenozoic 131-145,191,195,235-239, 349,351,352,354,371,511-514
249, 254-258, 267, 504-540 Codium 70, 151, 152
Ceratolithus 223 Codonophycus 389
Chabakovia 310, 322, 366, 388 Coelosphaeridium 118, 121, 351
Chaetophorales 514 Coelosporella 389
Chantonia 388 Collen ella 467
Chara 190, 204, 205, 206, 209-213, Collenia 543
214-215 Columbiapora 376, 390
Charophyta 189-203,204-216, 353, 365 Confervites 309
calcification 11,204,209,213-215 Coni/alia 390
classification 191, 201-202 Connexia 390
compound oosporangial wall Conophyton 299, 543
(COW) 205,206-213 Contortoporidium 390
evolution 191-193, 199 coralligene 109
gyrogonite 189, 191 Corallina 99
morphology 191-193 Corallinaceae 13, 95, 98-113, 155, 454,
oosphere 205 458, 506, 509-510
Index 565
calcification 99-103 morphology 131-132,134,138,139,

ecology 108 - 110 516
morphology 103 - 107, 108 -11 0 ranges 144, 489, 530-531
palaeobathymetry 110 Dasycladophycus 393, 518
taxonomy 103 - 108 Dasycladus 10, 129, 529
Corydopodium 390 Dasyporella 343, 350, 393
Costatumbella 390 Demidella 361
Cretaceous 77, 125, 191, 233, 240, 249, 267, dendrolite 22, 26, 27, 29, 34, 39, 41-42,
278, 457, 484, 521 45, 59, 63, 306
calcareous algae 504-540 definition 46
Cretacicladus 513, 514 Dermatolithon 107
Cribrokamaena 391 Devonian 76, 191, 319, 349, 354-366
Cribroporidium 391 diatoms IS, 292, 552
Cribrosphaeroides 391 Dimorphosiphon 149, 153, 157, 340, 344,
Cricosphaera 229 350, 352, 353, 361
Cruciplacolifhus 242 Dimorphosiphonoides ISO, 340, 350, 352
Crucirhabdulus 231-232 dinoflagellates 267 - 286
cryptic microbial carbonates 29, 37 calcareous wall 277 - 280
definition 46 ecology 280-281
Crypfozoon 29, 543 phylogeny 281
Cummingsella 379, 391 taxonomy 268 - 277
Cuneiphycus 103, 392, 456 Dinophyta, see dinoflagellates
cyanobacteria 5, 30, 292, 295, 300, Diplopora 128, 357, 365, 461, 490, 518,
305-334, 351, 371, 466-467, 498, 519, 521
505-506, 545, 546,553 Discoaster 217,223,224,235,237-238,
calcification 4, 56-58, 74, 75-78, 79, 242, 243, 248, 249, 254, 256, 257
80,305,307 Discosphaera 225
calcified 55 - 87 Dissocladella 141, 519
classification 64 Distichoplax 509
coccoid 57, 61, 69, 313, 319, 323 Diversocallis 158
evolution 81, 82 Diversoporella 351
geological history 41-42, 74-78, 301 Dobunniella 521
sheath 5, 56, 57, 59, 65, 79, 300-301 Dokutchaevskella 393
cyanoid 46, 61-62 Donezella 379, 393, 461
Cyclocriniles 115-117,118-119,120,121 Dromastacheoides 394
cyclocrinitids 114 - 124 Dutroella 394
morphology 115 -119 Dvinella 379, 394
palaeoecology 120-121 Dzhuljanella 456, 510
Cyclocrinus 351
Cylindrojolia 392 ecology
Cylindroporella 519, 523, 529 coccolithophorid 225 - 229, 236,
Cymopolia 10, 127, 131, 132, 134, 141, 248-260
142, 519, 529 Corallinaceae 108 -110
cyrtoliths 225 cyclocrinitid 120-121
Dasycladales 128, 142
Dactylopora 138 dinoflagellate 280 - 281
Dasycladales 114,118,120,125-145,153, Gymnocodiaceae 159
317,327, 328, 340, 342-343, 349, 350, Halimedaceae 171-175
353, 357, 358, 371, 453, 455, 461, 464, nannofossil 236, 248 - 260
484,487,489,490,496, 513, 514-529 stromatolite 296,553-556, 556-557
calcification 141 Udoteaceae 153
classification 125 - 126, 128 - 129 Eflugelia 395
cysts 131, 136, 139, 141 Einorella 395
ecology 128, 142 Elenia 395
evolution 126, 141-142, 144,517,524 Elianella 509
extinctions 127, 518, 527 Ellipsagelosphaera 233
566 Index
Ellipsolithus 242 Girvanella 32, 55, 60, 66-67, 70, 76, 78,
Emiliania II, 12, 16, 220, 226, 228, 80, 151, 309, 310, 313, 315, 317, 325,
236-237, 243, 244, 245-247, 249-253, 381, 402, 403, 466
259 Givetianella 357
Ensicu/ijera 266, 274, 276, 282 Globator 194
Entophysa/is 79, 544, 552, 554 Globulijeroporella 403, 461
£ochara 193 Goksuella 403
£oclypeina 396 Goniolithon 99, 107
£oconusphaera 230 Gordonophyton 65, 309, 315, 321, 324
£ogoniolina 461 Gouldina 403
£okoninckopora 396 Griphoporella 519, 521, 527
£olithoporella 396 Grozdilovella 403
£osphaera 514 Guvencipora 509
£oumbella 390, 396 Gymnocodiaceae 146-166, 452, 455,
£oura/oporella 397 456-457, 463, 506, 510
£ove/ebitella 376, 379, 381, 397 calcification 157
£ovo/vox 514 classification 154, 157, 158,
Epimastopora 357, 379, 381, 397, 398, 159-160
461-462 ecology 159
Epimastoporella 379, 398, 461-462 morphology 154-155,159-160
Epiphyton 26, 35, 60, 62, 65-66, 69-70, reproduction 155-157
76, 306, 307, 309, 313, 315, 317, 319, silicification 157
323, 366, 398, 506 Gymnocodium 147, 151, 154, 155, 157,
Epistacheoides 377, 381, 399 158, 456
Ethelia 510 gyrogonite 189, 191
Eugonophyllum 399, 460, 470 Gyroporella 154, 379, 397, 403, 461, 463,
Evlania 400 517, 518, 523
extinction
Cretaceous/Tertiary 232-235, 527 Halenopora 404
discoaster 257 Halicoryne 138, 142
Jurassic/Cretaceous 233 Halimeda 10, 70, 127, 147, 148, 150, 152,
nannofossil 233 - 236, 240 153,154,160,167-188,494,496,511
Permian/niassic 126-127, 487, 518 Halimedaceae 167 - 188, 464
stromatolite 291, 302 bathymetry 174
niassic/Jurassic 492, 519 calcification 170 - 171
Exvotarisella 363, 381, 400 definition 167
Fanesella 519 ecology 171-175
Fasciculithus 242 morphology 170-171, 173
Fasciella 400 ranges 496-497
Foliophycus 401 and reefs 172-175,178-186
Flabellia 71, 351, 354, 365 sediments 175- 185, 494, 496
F/orisphaera 226, 249 Halysis 351, 354, 365
Fourstonella 381, 401 Harrisichara 195
Frustulata 401 Hayaster 226
Frutexites 65, 366 Hedstroemia 67-68,70,76,78, 151,325,
Furcoporella 518 350, 352, 354, 359, 404, 466, 467, 505
Heptasphaera 272
Galaxaura 148, 154, 155, 157, 456, 510 Herakella 405
Gatnerago 233 Heterococcolith 222
Garwoodella 402 Heteroporella 490, 519, 523
Garwoodia 70, 76, 350, 352, 359, 402, 505, Heterotrichella 514
514 Hikorocodium 405
Gemma 68-69, 322 Holococcolith 222, 245, 248
Gephyrocapsa 219, 226, 236, 244, 246, Homoeothrix 9
249-253, 259 Hydrolithon 110
geyserite 43 Hymenomonas 228
Index 567
Iapetus 336 Macroporella 409,461-462, 527

Iberiaella 405 maerl 109, 509
Intermurella 340-342, 350 Malacostroma 410
Intextulella 405 Malakhovella 410
Issinella 363, 376, 405 Mametella 410
Ivanovia 377, 381, 406, 460 Marinella 509
Izhella 68-69, 322, 365, 406, 467 Markalius 233-234
Masloviporella 410
Jania 99 Masloviporidium 379, 381, 390, 411
Jansaella 363 Mastopora 121, 351
Japhetella 376, 406 Mazaganella 231-232
Juraella 511 Mejerella 306, 312, 327
Jurassic 77, 88, 125, 191, 195, 232-233, Mellporella 411
240, 267, 281, 504, 519, Menselina 411
calcareous algae 481-503,504-540 Mesolithon 95
biostratigraphy 494 - 496 Mesophyllum 94-95, 99, 109, 110, 509
evolution 490-493 Metasolenopora 93
ranges 486 microbial carbonates 46, (see also cryptic
sedimentology 496 - 498 microbial carbonates)
composition and fabrics 38
Kamaena 363, 365, 407 definition 22
Kamaenella 363, 377, 381, 407 microbial community 292, 300-301, 544,
Karpinskya 198 545-547
Katavella 361 microbial crusts 497 - 498
Kazakhstanelia 340, 350 microbolite 22, 27, 44
Komia 381, 408, 459 definition 46
Koninckopora 381,408 microbush 59, 63
Koninckoporoides 409 Microcodium 366, 376, 412, 506, 532
Korilophyton 321, 323, 324 microrhabulids 223
Koskinobullina 511 mineralization 27, 28, 47, 98, 101
Kulikia 126, 409, 518 Mitcheldeania 70, 76, 151, 412, 511
Kymalithon 509 Mitrolithus 232
Mizzia 127, 461-462, 518
Lamprothamnium 197, 198,204, 205, 206, Moellerina 198
209-213,214-215 Moniliporella 352
Lancicula 356 Montiella 131
Leckhamptonella 149 Musacchiella 199
Lee Stocking Island 31, 42, 44, 556
Leptophyton 105 Nanjinophycus 157, 158, 159, 456-457,
Liagora 157 464,510
Likanella 518 Nannoconus 224, 240
Litanaia 150, 353, 354, 361 nannofossil 217 - 266, (see also coccolith
Lithocodium 73-74, 77, 459, 532 and coccolithophorids)
Litholepis 107 biostratigraphy 239-244
Lithophyllum 99, 107, 109, 110, 509 and climate 253 - 260
Lithoporella 90, 95 diagenesis 248
Lithothamnion 105, 107, 109, 110 dissolution 244-248
Lithothamnium 94-95, 99, 107, 509 ecology 236, 248 - 260
Litostroma 409 evolution 229 - 239
Lopsiella 515 extinction 233 - 236, 240
Lotharingius 232 isotopic composition 255, 258 - 260
Lowvillia 150, 153, 340, 350, 352 palaeobathyrnetry 248
Lulipora 515 sedimentation 244 - 248
Luteotubulus 409 Nanopora 381,412
Lychnothamnus 204 Nansenella 376, 413
Lysvaella 458 Neoanchicodium 377,413, 464
568 Index
Neogene 242 Parachaetetes 93-95, 351, 353, 361, 363,

Neogoniolithon 107, 109, 110 381, 417, 458, 509
Neogyroporella 523 Paradella 417
Neomeris 10, 115, 127, 131, 132, 134, 141, Paraepimastopora 379, 418, 461-462
142, 519, 523, 529 Paraepiphyton 366
Neosolenopora 92, 509 Paragarwoodia 151
Neoteutloporella 523, 527 Parakamaena 363, 418
Nephelostroma 309,312 Paralancicula 361
Nicholsonia 315, 325 Paralitanaia 361
Nidulites 116, 121 Parastacheia 419
Nigriporella 413, 466, 533 Parkerella 134
Nipponophycus 158,511 Paronipora 506, 532
Nitella 190, 204, 205 Pedobesia 167
Nitellites 190 Pekiskopora 419
Nitellopsis 195, 196, 199, 204 Penicillus 10,148,153,160,167,170-174,
Nostocites 354, 413 173, 181, 182-183
Novantiella 350 Pentadinellum 267, 283 - 284
Nuia 351, 352 Perimneste 193, 194
Peristacheia 419
Permian 76, 125, 159, 191, 232, 335, 437,
Obliquipithonella 268, 272, 274, 276, 279,
486, 489, 516
281
calcareous algae 452-480
Obruchevella 312,317, 326
classification 455 - 456
Oligoplagia 158, 510
environments 453-454
Oligoporella 490, 519
groups 454-467
oncoid 47, 59, 61, 62, 63, 76, 297, 466,
ranges 474, 488-489
496-497
Permocalculus 154, 155, 157, 158, 159,
Oocardium 11, 552
456-457, 510
Oolithotus 226, 245
Petrascula 523
Ordovician 76, 119, 120, 121, 335-348,
Petrophyton 93-95, 363
349-352
Petschoria 419
Orioporella 141
Peyssonelia 510
Orthopithonella 268, 269, 272, 277,
Phacotus 10
279-280
Phormidium 9,62,63,543, 544, 547, 552
Orthriosiphon 414
photosynthesis 4, 15, 81, 546
Orthriosiphonoides 150, 379, 381, 414, 464
phylloid algae 371, 375, 382-383, 387,
Ortonella 32, 55, 60, 67, 76, 151, 350, 352,
399-400, 406-461, 413, 437, 452, 453,
354, 359, 466, 505
454, 455, 458, 460-461, 464, 470, 473,
Osagia 415, 466
511
Ostreobium 79
Phymatolithon 99, lOS, 107, 109
Ottonosia 416, 466
Physoporella 490, 518
Ovulites 511
Pithonella 266
Placklesia 519
Pachysphaera 416 Placolith 222, 232, 243, 248
Pachysphaerina 416 Plectonema 67, 326
Padina 13 Pleistocene 243-244,253
Palaeoberesella 128, 363, 416 Pleurochrysis 220
Palaeocene 238, 240, 527 Poikiloporella 519
Palaeochara 193 Pokorninella 420
Palaeodasycladus 496, 526 Polymorphocodium 420
Palaeomicrocodium 365, 416, 532 Polystrata 510
Palaeoporella 147, 151, 340, 344, 349, 353, Poncetellina 361
361 Porochara 97, 199
Palaeosiphonocladales 363, 371 Porolithon 99, 109, 110
Paleothamnium 95 Porostromata 64, 151, 342,483-484,486,
Papponamonas 226 496
Index 569
Praedonezella 420 Rivularia 7, 41, 67, 78, 80, 307, 325, 326,
Praesycidium 191 327, 467, 506, 544, 551, 552
Praturlonella 141, 529 Roquesselsia 427
Precambrian 40, 41, 42, 75, 289 Rothia 197
Prediscosphaera 233 Rothpletzella 71-72, 76, 325, 350, 428
Principia 420 Russoella 523
Prinsiosphaera 230, 231
Proaulopora 314, 317, 326-327 Salpingoporella 495, 523, 527
Proninella 365, 421 Samarella 427
Protoumbella 421 Sandoella 427
Prymnesiophyta, see coccolithophorids Sarfatiella 519, 521
Pseudochaetetes 93-95, 353, 361, 421 Schizosphaerella 224, 230, 231, 233
Pseudocodium 151 Schizothrix 9, 60, 62, 64, 319, 323, 546
Pseudoemiliania 236, 259 Scrippsiella 266, 282
Pseudoepimastopora 421 Scytonema 67, 151
Pseudogyroporella 421 Seletonella 128, 306, 308, 315, 317,
Pseudohedstroemia 422 327-328, 349, 515
Pseudoissinella 363, 422 Selliporella 521, 523
Pseudokamaena 363, 422 Sgrossoella 513, 514
Pseudokomia 423 Shamovella 427, 466
Pseudolithophyllum 107 Shark Bay 31, 42, 43, 44, 78, 79, 292, 546,
Pseudolithothamnium 158, 510 553
Pseudonanopora 423 Shartymophycus 427
Pseudosolenopora 361, 423 Shermanophycus 427
Pseudostacheoides 423 Shuguria 322, 366, 427
Pseudoumbella 424 Siberia 310 - 311
Pseudovelebitella 424 Siberiella 327
Pseudovermiporella 461, 463 Silurian 76, 121, 349, 352-354
Pycnoporidium 93, 459, 509 Sinarella 428
Pycnostroma 424 sinter 43, 47
Pyrulites 154, 157 Sinustacheoides 428
Solar Lake 552
Quasiumbella 424 Solenomeris 509
Quasiumbelloides 425 Solenopora 76, 93, 99, 307, 312, 342, 350,
Queenslandella 376, 425, 518 353, 361, 363, 373, 428, 458-459, 468,
509
Radiosphaera 425 Solenoporaceae 88-97, 350, 353, 361, 455,
Razumovskia 310 458, 459, 509
receptaculitids 467 classification 92 - 95
Rectangulina 426 evolution 94-95
reefs 35, 59, 62, 63, 65, 77, 108 -109, 153, structure 89-92
172, 178, 185, 306, 325, 342, 351, 452, Solenoporella 93
453,454,458, 461, 470, 471, 473, 498, Solieriaceae 510
510 Sphaerinvia 428
Renalcis 26, 35, 63, 68 -70, 76, 307, 310, Sphaerochara 196, 197
313, 317, 319, 320, 321, 322, 365, 366, Sphaerocodium 71-72, 350, 352, 354, 358,
426,467 428
Retephycus 426 Sphaeroporella 72, 429
Reticulofenestra 218, 248 Sphaerospongia 119
Rhabdoceras 232 Sphenolithus 235, 242
Rhabdochara 190 Sphinctoporella 429
Rhabdoporella 73, 339-340, 343, 350, 357 Spinumbella 430
Rhabdosphaera 225 Spongiostroma 371, 382, 410, 424, 430,
Rhipocephalus 10,167,170-174 486,497
rhodolith 11 0 Spongites 108
Richella 377, 427 Squamariaceae 95, 101, 158, 460, 506, 510
.570 Index
Stacheia 430 Thorosphaera 226

Stacheoidella 430 thrombolite 27, 29, 35, 39, 41, 45, .59, 63
Stacheoides 431 definition 47
Stichoporella .519, .521 Thyrsoporella 138
Stipulella 431 Titanoderma 107-109, 110
stromatolite 24, 27, 29, 40, 43, 47, 61, 72, 7blypella 190, 196, 197, 204, 205
7.5, 76, 289-304, 307, 312, 467, 47.5, .50.5, 7bmilithon 432
.541-.561 travertine 27, 29, 37, 45, 47
~utinakd 22, 29, 30, 44, 61 lliassic 77, 125, 191, 230, 232, 269, 280,
definition 46 281
coarse-grained 31 calcareous algae 481-503, 504-540
classification 543 biostratigraphy 494-496
definition 30, 47, 293-294, 542 evolution 490-493
diversification 81, 297 - 302 ranges 486
ecology 296,553-556,556-557 sedimentology 496 - 498
extinction 291, 302 lrinocladus 141
geological history 39-42, 75, 289-292, Trinodella 433
294-302 Triploporella 129, 141, 523
history of research 42-44 Triquetrorhabdulus 235
lacustrine 557 1ilbiphytes 73, 77, 433, 452, 453, 464-466,
lamination 545-547 467, 470, 471, 495, 506, 532
lithification 75, 79, 296, 547-549 1ilbomorphophyton 65, 315, 324
morphology 290-291,297, 544 tufa 47, 63
nonskeletal 44, 47 tufa stromatolite 29, 30, 32, 45, 47, 63
potential 543 1ilrrisphaera 227
processes of formation 22-27 'Ijldemania 167, 170, 182
skeletal 29, 32, 41-42, 45, 47, 59, 72,
79-80 Udotea 10, 166, 170-174, 182, 184
subaerial 29, 33, 47 Udoteaceae 146-188, 340-341, 343, 350,
trapping 24, 75 353, 354, 355, 371, 492, 496, 511
tufa 29, 32, 45, 63 calcification 150
definition 30, 47 classification 150 -153
Subtifloria 321, 325 ecology 153
Sycidium 353 evolution 153-154
Syracosphaera 219 morphology 147 -148
Stylaella 431 reproduction 149 -150, 151
Stylocodium 431 Ulocladia 357
Stylophycus 432 Ulotrichales 514
Subkamaena 432 umbellids 365, 437
Subtifloria 60, 67, 76 Umbellosphaera 225, 245
Succodium 158, 159, 456, 464, 510 Umbilicosphaera 225, 246, 248
Sycidium 432 Unella 357
Ungdarella 381, 433, 459
Taldykites 432 Unjaella 434
Taninia 321 Uraloporella 379, 434
Tarthinia 36,63,68-70,321 Uteria 138
Tauridium 149, 456, 464 Uva 150
Tectochara 195, 196, 198
Tenarea 108
Terquemella 138, 521 Velebitella 434, 461, 517
Thtrasporales 513, 514 Vermiporella 128, 337, 344, 350, 365, 435,
Teutloporella 490, 518, 519 514
Tharama 366, 467 Volvocales 514
Thaumatoporellales 513, 514 Voycarella 356
Thoracosphaera 14, 224, 225, 230, 268, 282
Thoracosphaerales 268 Wallidinellum 267, 282-283
Index 571
Wetheredella 72-73, 76, 343, 351, 353, Yukonefla 436

365, 381, 435, 511
Wigwamma 226 Zaporella 436
Windsoporella 381, 435 Zidella 436
Zittelina 131, 134, 138
Yakutina 306, 312, 327 Zonotrichites 77, 454, 467, 506

Riding R., 1991. Calcareous Algae and Stromatolites

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Riding R., 1991. Calcareous Algae and Stromatolites

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Robert Riding (Ed.

With 16 Plates and 202 Figures

ISBN 978-3-642-52337-3 ISBN 978-3-642-52335-9 (eBook)

Library of Congress Cataloging in Publication Data. Calcareous algae and stromatolites / R.

"Calcareous algae and stromatolites" is shorthand for a wider array

Cardiff, August 1990 ROBERI' RIDING

II Major Groups .................................... 53

III Algae and StromatoUtes Through Time .............. 287

15 Archaean and Proterozoic Stromatolites

Index ................................................ 563

Awramik, S. M. 289 Hillis, L. 167

organically constructed template or framework, from 'biologically induced'

2 Calcification Processes in Cyanobacteria

Key for lettering

Fig. I. Pie charts illustrating the approx- Phaeophyta

cyanobacteria as "the nucleation of calcium carbonate upon or within the

Fig. 2a-d. Calcification and carbonate-binding by the cyanobacterium sheath. a Calcification

heterogeneous nucleation occurs at favourable sites in a sheath composed of

formation by ammonification or nitrate-reduction (Krumbein (979). The

......:..... - ..... .. .. ... _.

s·:·· ···_.... ·..;· .... ·

winter, the calcification is abiogenic, whilst in summer, it results from a com-

Lyngbya and Scytonema. Under the same conditions, Entophysalis and

3 Calcification Processes in Algae

Most calcified genera are marine macrophytes belonging to the Dasycladales

calcifies in vitro at high pH, possibly stimulated by photosynthesis (Devi-Prasad

Extracellular deposits of calcium carbonate occur on the internodal and some

analyzed. Within the Prymnesiophyta as a whole, the link between specific

3.6 Dinophyta, Chrysophyta, and Bacillariophyta

Calcification is rarely reported in these groups and deposition is varied, unusual,

species (Tappan 1980), a genus normally characteristic of aggressive, oligo-

4 Discussion and Conclusions

Most calcifying algal regimes are characterized by a water pH of7.8-9.5. Logic

protective coatings preventing dissolution of the mineral phase, which is often

the synthesis of intracellular coccoliths down to the random deposition of calcite

The principal categories of benthic microbial carbonate deposits are:

1 Department of Geology, University of Wales College of Cardiff, Cardiff CF I 3YE, UK

primarily by microbial organisms such as bacteria, cyanobacteria, and algae,

modern hot spring

The degree of control over mineralogy and crystal growth in biomineralization

3.shelter cavity &

Fig. 4. Composition of calcified microbial deposits exemplified by dendrolite, consisting of 1

2.3 External Surficial Precipitation of Minerals

This process, here termed mineralization, is, in effect, early cementation. In

3 Types of Benthic Microbial Carbonates

Microbial growth Prostrate Erect Coccoid

especially in the Early Palaeozoic, as probable cyanobacteria. Consequently,

The important distinction between stromatolites on the one hand and

I. Stromatolite: macroscopically laminated benthic microbial deposit.

3.2.1 Agglutinated Stromatolites

This term is introduced here for stromatolites produced by trapping of par-

Fine-Grained, Well-Laminated. This is the predominant Precambrian stroma-

1. Smooth lamination, of subaqueous origin:

Coarse-Grained, Crudely Laminated. Awramik and Riding (1988) recognized a

3.2.2 Tufa Stromatolites

This term is introduced here for stromatolites dominated by precipitation of

3.2.3 Skeletal stromatolites

Riding (1977) introduced this term to describe "stromatolites in which the

Fig. 6. Relationships between processes of carbonate deposition in the formation of nonskeletal

evasion is enhanced by water movement. As a result, it is possible for lacustrine

3.2.4 Subaerial Stromatolites

Research on highly calcareous soils and subaerially exposed limestone surfaces

Subaerial stromatolites form in environments dominated by desiccation

This term was introduced by Riding (1988) for non-laminated biomineralized