BATALLERIA 19 2013 41- 46 (Barcelona, noviembre 2013) ISSN0214-7831

Chirality in the Late Palaeozoic fenestrate

bryozoan Archimedes
Paul D. Taylor & Consuelo Sendino
Department of Earth Sciences Natural History Museum, Cromwell Road,
London SW7 5BD, UK

ABSTRACT – The Carboniferous–Permian bryozoan Archimedes (Order Fenestrata) has characteristic

screw-like axes from which a typical fenestellid meshwork radiated during life. Thickly calcified Archimedes
screws broken into length of a few centimetres are commonly at outcrops collected minus the meshwork.
Screws can be either sinistral or dextral, respectively coiling in an anticlockwise or clockwise direction with
distal growth. Here we investigate Archimedes chirality in order to establish whether there is any preference
for sinistral or dextral screws in samples from particular localities and also in composite collections from
multiple localities. While the composite collections revealed no predominance of one coiling direction over
the other, two of the seven locality samples showed a statistically significant dominance of sinistral over
dextral screws. The two samples with a dominance of sinistral screws may be either statistical artefacts or
due to the fragmentation of long sinistral screws. There is no evidence for a biological preference for one
coiling direction over the other in Archimedes and it is known that individual colonies may contain both
sinistral and dextral screws.

KEY WORDS – Bryozoa, Fenestrata, Carboniferous, Morphology, Handedness

RESUMEN – El briozoo del Carbonífero-Pérmico Archimedes (Orden Fenestrata) tiene una forma
característica de ejes tipo tornillo desde los cuales una retícula típica fenestėlida se radiaba en vida. Los
“tornillos” densamente calcificados de Archimedes aparecen comúnmente sin la retícula en fragmentos de
unos pocos centímetros de longitud en los afloramientos en los que se han recogido. Los tornillos pueden ser
o bien sinistrales o dextrales, teniendo en cuenta el crecimiento distal en el sentido de las agujas de reloj o en
sentido contrario de éstas. Aquí investigamos la quiralidad de Archimedes con el fin de establecer si existe
alguna preferencia por la forma sinistral o dextral en los ejemplares de determinadas localidades y también
en colecciones con muestras de varias localidades. Mientras que las colecciones compuestas no mostraron
un predominio en una u otra dirección de enrollamiento, dos de las siete localidades estudiadas mostraron
una dominancia estadísticamente significativa de las formas sinistrales sobre las dextrales. Las localidades
con preponderancia de sinistrales puede ser debido por artefactos estadísticos o por la fragmentación de los
ejemplares sinistrales más largos. No hay evidencia de una preferencia biológica en un sentido de
enrollamiento sobre otro en Archimedes y se sabe que las colonias individuales pueden contener ambos
tornillos, sinistral y dextral.

KEY WORDS – Bryozoa, Fenestrata, Carbonífero, Morfología, Lateralidad

INTRODUCCTION seen to form a continuous spiral growing upwards

Named after the water pump invented by the
Greek philosopher, Archimedes Owen, 1838 is
probably the most iconic of all fossil bryozoans.
Screw-like axes (Fig. 1) of this fenestellid (Order
Fenestrata) consisting of dense calcite frequently
weather from outcrops after loss of the more
delicate meshwork of branches and dissepiments.
In more complete colonies the meshwork can be
and outwards from the helical axis. The axis itself
is actually formed by thickening of the skeleton of
axial parts of the meshwork; it is not an
independent structure.
Fossils of Archimedes are particularly common
in the Carboniferous of North America from
where McKinney (1993) recognized the presence
of 39 species. However, the genus is also recorded
42 PAUL D. TAYLOR & CONSUELO SENDINO

in the Carboniferous and Permian of Russia. In were collected from the Carboniferous of the
eastern North America Archimedes mainly USA.
inhabited shallow-water environments behind
migrating calcarenite dunes (McKinney, 1979). Specimens of Archimedes are most commonly
Most of the colonies in this environment were preserved as broken lengths of screw axes. These
formed by fragmentation of pre-existing colonies, tend to measure 2–3 cm in length, with the longest
i.e., by cloning rather than the more conventional example in the NHMUK collection measuring
mode of bryozoan recruitment via larvae produced 18.5 cm (D2084, Warsaw Group, Warsaw, Iowa).
by sexual reproduction (McKinney, 1983). However, Archimedes screws exceeding one
metre in length are known to occur (F.K.
Interest in Archimedes during the mid-19th McKinney, historical comment to PDT). In order
century was stimulated by the theory of Condra & to determine handedness, screws were oriented
Elias (1944) that the fossil was a consortium with their proximal ends at the bottom and distal
between a conventional planar fenestellid ends at the top (see Figure 1), i.e. in growth
bryozoan and an alga which was responsible for orientation. Determining the polarity of screws is
the helical axis, a hypothesis that did not win straightforward because the undersides of the
support and which has been discarded. More whorls are long and slope at a low angle to the
recently, McKinney and coauthors (see McKinney axis of the screw, whereas the uppersides are
& McGhee, 2003) focused on the geometry of the shorter and steeper. When oriented in this way
helical colonies, using computer simulations to sinistral screws have whorls running from lower
investigate the morphospace occupied by left to upper right, dextral screws from upper left
Archimedes as well as inferring the feeding to lower right. Note that this is the opposite of
currents that were generated by living colonies gastropod molluscs, the reason being that
(McKinney et al., 1986). conventionally oriented gastropods with the
aperture at the bottom grow from top to bottom
The helical coiling of the Archimedes axis can rather than bottom to top.
be either left-handed (sinistral) or right-handed
(dextral) which produce enantiomorphic axes that
are asymmetrical but not superimposable on each
other (chiral). Twisting or translation with distal
extension of the axis was anticlockwise in sinistral
axes, but clockwise in dextral axes. Surprisingly,
little or no attention has been paid to the
handedness of Archimedes axes, and handedness
does not feature in descriptions of species of
Archimedes. Does Archimedes as a whole or do
particular species of the genus show a
preponderance of sinistral or dextral axes, or do
these two enantiomorphs occur in equal numbers?
Here we investigate chirality in museum samples
of Archimedes to address this question potentially
of importance in understanding the developmental
palaeobiology of this distinctive bryozoan.

MATERIAL AND METHODS

A ‘population’ of 912 specimens of

Archimedes in the following collections was
studied: Natural History Museum, London, UK
(NHMUK); Trinity College, Dublin, Ireland
(TCD); National Museum of Natural History
(United States National Museum), Smithsonian
Institution, Washington DC, USA (USNM). All
Fig. 1. Typical example of a fragment of an
Archimedes screw. Oriented with distal at the top,
this is a sinistral screw. NHMUK PI D34127,
Lower Carboniferous, Golconda Formation,
Marigold, Illinois, USA. Scale bar = 5 mm.
 CHIRALITY IN THE LATE PALAEOZOIC FENESTRATE
BRYOZOAN ARCHIMEDES 43

An initial survey was undertaken of chirality in
a total of 322 Archimedes screws in the NHMUK
collection. In most cases, sample size from
individual localities numbered fewer than 10
specimens. Only three samples comprising
more than 30 screws were judged large
enough to warrant separate analysis. In order
to supplement these large samples, additional
sizeable samples from single localities were
solicited from M. McKinney (Appalachian
State University, Boone, North Carolina) and
P. N. Wyse Jackson (Trinity College, Dublin),
and were studied at the USNM. It should be
noted that species identification of
Archimedes screws is difficult in the absence
of thin sections of the meshworks and
therefore samples from each locality may
potentially comprise screws from more than
one species. Finally, 77 images of Archimedes
screws published on the web were surveyed
for handedness, as were the 64 screws
illustrated by Condra & Elias (1944) for
which handedness could be determined
unequivocally.
Counts were analysed using simple chi-
squared (χ2) tests. The null hypothesis is that an
equal number of sinistral and dextral screws will
be present in each sample. Two-tailed P values of
less than 0.09 were taken to indicate a statistically
significant difference in the number of sinistral vs.
dextral screws.
RESULTS
Table 1 summarises the numbers of sinistral
and dextral screws counted in the various samples
analysed. Of the 10 samples analysed, only two
showed a statistically significant dominance of
one coiling direction over the other. These
samples came from the Bangor Limestone of
Foxtrap, Alabama, and the Fayetteville Formation
of Uinita, Oklahoma. The Foxtrap sample was
obtained by one of us (PDT) during two separate
collecting trips separated by 16 years, whereas the
Uinita sample (Fig. 2) was made by Harrell
Strimple and presented to the USNM. In both of
these cases, sinistral screws.

sample total sinistral dextral χ2 P

NHMUK total collection 322 173 149 1.789 0.1811
NHMUK Bangor Limestone, Foxtrap, Alabama 39 28 11 7.410 0.0065*
NHMUK Sloans Valley Member, Burnside, Kentucky 66 34 32 0.061 0.8055
NHMUK Bangor Limestone, Reid Gap, Alabama 68 38 30 0.941 0.3320
NHMUK Pride Mountain Fm., Huntsville, Alabama 63 27 36 1.286 0.2568
TCD Big Clifty Formation, Sulphur, Indiana 72 35 37 0.056 0.8137
USNM Chesterian, Addison, Kentucky 72 43 29 2.722 0.0990
USNM Fayetteville Fm., Uinita, Oklahama 69 45 24 6.391 0.0115*
Internet pages 77 37 40 0.117 0.7324
Condra & Elias (1944) figures 64 35 29 0.563 0.4533

Table 1. Handedness of Archimedes screws. Data comes from: (1) the complete NHMUK collection of
specimens; (2) material from individual localities containing more than 30 screws represented in the
NHMUK, TCD and USNM collections; (3) screws illustrated on web pages; and (4) screws figured in
the monograph on Archimedes by Condra & Elia (1944).  values (1 degree of freedom) test the
null hypothesis that equal numbers of sinistral and dextral screws are present in each sample, with
two-tailed P values (* = statistically significant differences). Note that the screws from Foxtrap were
collected during two different field trips, one in 1982 and the second in 1998. Samples from Reid Gap
and Huntsville, Alabama now in the NHMUK collection were acquired after the count of total
NHMUK specimens
44 PAUL D. TAYLOR & CONSUELO SENDINO

were more numerous than dextral screws: 28
sinistral vs. 11 dextral from Foxtrap, and 45
sinistral vs. 24 dextral from Uinita.
DISCUSSION
Unlike gastropod molluscs where the
overwhelming majority of species have shells of
one handedness, usually dextral (e.g. Hendricks,
2009), there is no strong pattern of handedness in
screws of the bryozoan Archimedes. Samples
comprising screws from multiple localities in the
collections of the NHMUK, illustrated on the
Internet and figured by Condra & Elias (1944) all
show statistically equal numbers of sinistral and
dextral screws. Equal proportions of the two
enantiomorphs also characterize most samples
(‘fossil populations’) from single localities.
However, samples from two localities – Fox Trap,
Alabama (NHMUK), and Uinita, Oklahoma
(USNM) – each show a significant predominance
of sinistral over dextral screws. Several
explanations may be proposed for these two
anomalous samples.
and on the seemingly rare occasions that screws
bifurcate the two daughter screws have the same
chirality as each other and as the parental screw
(Condra & Elias 1944, p. 65). This means that all
of the screw fragments resulting from the
breakage of a single screw will have the same
handedness. Such breakage could occur, for
example, by weathering at outcrop. Given that
screws can be up to about a metre in length, and
fragments in museum collections are typically
only 2–3 cm long, an individual screw could
break into 30–50 screw fragments.
Collector selectivity – Interestingly, a sample
of 16 Archimedes screws, presumably of the same
species, bought from F. Braün in the 19th century
were entirely dextral. However, these specimens
came from the same locality (Huntsville,
Alabama) as a more recent collection in which
there was no statistically significant difference in
handedness. It seems possible that dextral screws
may have been chosen preferentially over sinistral
screws in the Braün material, either when they
were collected in the field or prior to their sale as
the vendor may have wanted to sell specimens of

Fig. 2. Sample of Archimedes screws from the Lower Carboniferous Fayetteville Formation of Chester,
Fragmentation
Uinita, Oklahoma,ofUSA,
singlecollected
screws –byIndividual
H. Strimple (USNM accession 216603). Sinistral screws (left) are
screws do not change chirality along
significantly more abundant than their lengths,
dextral screws (right) in this sample. Scale bar = 20 mm.
 CHIRALITY IN THE LATE PALAEOZOIC FENESTRATE
BRYOZOAN ARCHIMEDES
uniform appearance. Of the two large samples
showing a predominance of one coiling direction,
the sample from Foxtrap was collected by one of
us (PDT) in the field without any conscious
bias, allowing collector selectivity to be
largely discounted.
Statistical artefact – Only 2 of the 7 large
samples from single localities showed a
statistically significant preponderance of one
enantiomorph (both sinistral) over the other. It is
conceivable that these are purely chance
occurrences. Flipping coins a set number of times
if repeated often enough will eventually return
some repeats in which either heads or tails are
significantly over-represented.
Empirical proof of the occurrence of both
sinistral and dextral screws within the same
colony comes from some exceptionally preserved
specimens of Archimedes. McKinney & Burdick
(2001) described a rare example of the base of a
sexually recruited colony of Archimedes cf.
intermedius (Ulrich, 1890) from the Bangor
Limestone of Alabama. This colony, which grew
around a perished cylindrical substrate, has two
similarly sized screws growing in opposite
directions. The screw on the left-side is sinistral
whereas that on the right-side is dextral.
It is also known that screws may originate for
the outer edges of the meshworks of parent
colonies. These screws could subsequently break-
off living colonies to initiate new clonal colonies,
an important mode of asexual colony reproduction
in Archimedes (McKinney, 1983). Condra & Elias
(1944, pl. 9) depicted the edge of a meshwork of
A. distans Ulrich, 1890 from which two screws
are originating, one sinistral and the other dextral.
In summary, there is no conclusive evidence
for a predominance of left- vs. right-handed
screws in the Carboniferous–Permian bryozoan
Archimedes. Counts of the proportions of sinistral
and dextral screws in various types of samples of
Archimedes seldom showed a statistically
significant preference for one direction of coiling
over the other, and the few instances in which this
was not the case can be explained by non-
biological biases. Among these biasing factors the
most probable is considered to be that screw
fragments collected from some localities may be
dominated by or derived entirely from the
breakdown of a single long screw. This idea
would be worth investigating further as it may
45
furnish useful information about the relative
abundance of Archimedes colonies in the benthic
communities of which they were a component.
The equal proportion of sinistral and dextral
screws in Archimedes implies that there is no
developmental control over chirality in this
bryozoan. Similarly, no significant preference for
sinistral vs. dextral screws was noted in the
Eocene cyclostome Crisidmonea archimediformis
Taylor & McKinney, 1996, a species with screw
axes convergent with those of Archimedes. In
contrast, the Recent cheilostome Spiralaria florea
Busk, 1861 has helical branches that are always
sinistral (McKinney & Wass, 1981), implying a
greater degree of developmental control.
ACKNOWLEDGEMENTS
We are grateful to work experience students
Rebecca Scott and Oscar Lozada who undertook
some of the initial counts of Archimedes screws in
the NHMUK collections. Patrick Wyse Jackson
(TCD) loaned material from Sulphur and gave
constructive comments on the draft manuscript,
JoAnn Sanner (USNM) provided access to
collections under her care in Washington DC, and
Marjorie McKinney (Boone, North Carolina)
donated samples of Archimedes screws from
localities in Alabama. To all we are very grateful.
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