Complex behavioural patterns and ethological analysis of the

trace fossil Zoophycos: evidence from the Lower Devonian of

South China
LI-JUN ZHANG AND ZHAO ZHAO

Zhang, L.-J., & Zhao, Z. 2015: Complex behavioural patterns and ethological analysis
of the trace fossil Zoophycos: evidence from the Lower Devonian of South China.
Lethaia, DOI: 10.1111/let.12146.
The trace fossil Zoophycos is abundant in the shallow-marine deposits (tempestites) of
the Lower Devonian (Emsian) Yangmaba Formation in Ganxi of Sichuan, South
China. It often occurs as part of complex trace fossils that comprise different integrated elements: scratch traces, simple to complex spreiten structures with marginal
tubes (Zoophycos) and vertical tunnels. The complex Zoophycos burrows consist of
spreiten with a marginal tube, preserved as convex hyporeliefs on the sole of an erosion surface. The exquisite, complex spreiten are interpreted to have been formed by
deposit-feeding behaviour, where the animal constructed the trace upwards without
leaving faeces in the spreiten. The width of the marginal tube in different whorls is
almost constant. The scratches are observed on the wall of the marginal tubes. The
Zoophycos intergrades with Spongeliomorpha and Chondrites and was later cut by vertical shafts. All these features together indicate that the Zoophycos-maker might have
been a vermiform polychaete instead of a predator such as a decapod crustacean
(Spongeliomorpha producer). Based on stratigraphical and ichnological features, the
complex trace fossils resulted from the complex activity of different opportunistic
organisms (r-strategist) that quickly occupied and thrived within the quiet, nutrientrich environment after storm events. Complex trace fossil, Lower Devonian, Polychaeta, Yangmaba Formation, Zoophycos.
Li-Jun Zhang [ljzhanghpu@gmail.com] and Zhao Zhao [zzhaohpu@126.com], Key Laboratory of Biogenic Traces and Sedimentary Minerals of Henan Province, Henan
Polytechnic University, Jiaozuo, Henan 454003, China; manuscript received on 24/08/
2014; manuscript accepted on 12/03/2015.

Zoophycos is one of the most famous but enigmatic

trace fossils known from the Cambrian to the Holocene (Webby 1969; Alpert 1977; Bottjer et al. 1988;
Olivero 2003; L
owemark et al. 2006; Knaust 2009a;
L
owemark 2012). Given over 190 years of studies on
Zoophycos (Massalongo 1855; Olivero 2007) and the
proposition of various three-dimensional models
(Wetzel & Werner 1981; Kotake 1989; Bromley
1991; Olivero & Gaillard 1996; L
owemark & Schafer
2003; Knaust 2009a; L
owemark 2012), the taxonomy
and morphological/ethological evolution of this
ichnogenus remains unsolved (Chamberlain 2000).
These uncertainties are primarily due to the varied
morphology and ethology of Zoophycos in different
environments and to the fact that no living organisms have been observed creating Zoophycos-type
traces today. For the high morphological variability,
many ichnologists now recognize and use grouping
instead of species names (Uchman 1995). An example is the Amuri Zoophycos (Ekdale & Lewis 1991).
Another enigmatic feature is that Palaeozoic
Zoophycos mainly occurs in shallow-marine deposits,
whereas Mesozoic and Cenozoic Zoophycos are

predominantly in shelf and deep-sea deposits (Bottjer et al. 1988; Neto de Carvalho & Rodriguez 2003;
Olivero 2003; Knaust 2004; Seilacher 2007). An
interesting hypothesis proposed by Olivero (2003)
suggests that the bathymetric shift of Zoophycos
through geological time reflects the same trend of its
producer, probably to survive possible mass extinctions.
In particular, Palaeozoic examples can contribute to a better understanding of the early evolution of Zoophycos and its producers. In this
article, a new occurrence of Zoophycos is reported
from the Lower Devonian mixed siliciclastic carbonates of South China. The new findings provide
insights into the morphological complexity of this
trace fossil and the related ethological pattern of
its producer. This study provides the opportunity
to examine burrowing behaviour of the Zoophycos-organism when the trace was first developed
in high abundance and to compare it with other
Late Palaeozoic examples from the Early and Middle Permian (Gong et al. 2008, 2010; Knaust
2009a).

DOI 10.1111/let.12146 2015 Lethaia Foundation. Published by John Wiley & Sons Ltd

L.-J. Zhang & Z. Zhao

Geological setting
The Ganxi of Sichuan in South China provides
excellent outcrops of Upper Palaeozoic strata, especially from the Devonian Period (Hou et al. 1988).
The study area is located in the northwestern part of
the Upper Yangtze Plate and belongs to the part of
the Early Palaeozoic Longmenshan depositional
basin (Fig. 1). The Devonian strata in this area were
deposited on a continental margin where no significant tectonic events occurred throughout the Devonian (Xian et al. 1995). The outcrops of Devonian
strata are exposed along the Pingtong River on a ca.
15-km-long section. The trace fossils presented in
this article were collected from the Lower Devonian
Yangmaba Formation of the Ganxi Section (Fig. 1)
exposed at 31550 47 N, 104400 16 E.
The trace-bearing interval (Bed 90) belongs to the
Lower Devonian Yangmaba Formation (Fig. 1) and

LETHAIA 10.1111/let.12146

consists of sandstone with mudstone interbeds in

the upper part and bioclastic limestone in the lower
part (Fig. 1). The Yangmaba Formation conformably overlies the Lower Devonian Ertaizi Formation and underlies the Middle Devonian
Jinbaoshi Formation (Fig. 1). The Ertaizi Formation
consists of dark-grey medium- and thick-bedded
bioclastic limestones and shales, most likely deposited in open platform environments (Hou et al.
1988). The Jinbaoshi Formation consists of thickbedded limestones intercalated with thin-bedded
siltstone and thick-bedded fine-grained sandstone.
The trace fossils described in this article occur in
the upper part (Bed 90) of the Yangmaba Formation, which records a transition from open carbonate platform (beds 8589) to storm-affected neritic
environments (beds 9092) (Zhang 2014). The
trace-bearing interval starts with 1-m-thick
medium-bedded grey bioclastic limestone, overlain
by 2-m-thick grey medium-bedded sandstone with

Fig. 1. Location maps and lithostratigraphy of the Ganxi Section in Sichuan Province of South China. A, B, location of the studied section. C, geological map of the Devonian Ganxi Section (modified after Hou et al. 1988). D, stratigraphical column of the LowerMiddle
Devonian and the detailed lithostratigraphy where the complex trace fossils of the Yangmaba Formation in the Ganxi Section occur.

LETHAIA 10.1111/let.12146

intercalated mudstones. The trace fossils were found

above an erosion surface and on the sole of a bed of
medium-bedded sandstone intercalated with mudstone (Fig. 2). Above this bed, sandstone dominates
and is accompanied by minor muddy siltstone and
siltstone (Bed 91) interpreted as storm deposits
(tempestites) (Zhang 2014). On the basis of the sedimentological evidence, the depositional environment of the interval with Zoophycos is interpreted as
being in water depths between fair-weather wave

Zoophycos from the Lower Devonian of South China

base and storm-wave base (Hou et al. 1988; Zhang

2014).

Material and methods

The Ganxi Section was studied during several field
seasons between 2009 and 2013. The specimens in
Figs 46 were collected and stored in the collection
of the Earth Science Museum of Henan Polytechnic

Fig. 2. Studied section displaying the upper part of the Yangmaba Formation (beds 8991). A, macroscopic field photograph of studied
Zoophycos in Bed 90 of the Yangmaba Formation. Red asterisk marks the location of complex trace fossils. B, close-up view of the position of the studied complex Zoophycos. Dashed line represents an erosional surface. s and m refer to sandstone and mudstone, respectively. C, field photograph showing the complete U-shaped Zoophycos and the scratch traces on the upper part before collection.

L.-J. Zhang & Z. Zhao

University, China. The specimens in the field were

weathered and mossy as shown in Fig. 3. To examine their detailed morphology, the specimens were
cleaned with a brush and distilled water in the
Laboratory of Pretreatment of the Key Laboratory of
Biogenic Traces and Sedimentary Minerals of Henan
Province. After that, the specimens were photographed in the Lab of Palaeontology.

Results
Complex trace fossils are large and elaborate structurally: probably representing a long period of occuA

LETHAIA 10.1111/let.12146

pation and operation, and variable behaviours

instead of one single type of simple or dominant
behaviour (Bromley & Hanken 2003; Miller 2003).
They fall largely into two groups: compound trace
fossils, resulting from the changing behaviour of a
single producer, and composite trace fossils, resulting from the interaction of discrete ichnotaxa (Pickerill 1994).
The complex (composite) trace fossils within the
studied interval are mostly preserved as hyporelief
on an erosion surface (Fig. 2). They consist of different types of trace fossils that commonly merge and
cross-cut each other (Figs 3, 4). The first type are
scratch traces in the galleries on the outer surface of
B

Fig. 3. Field photographs showing the details of the complex trace fossils. A, complex trace fossils on the sole of the sandstone unit. B,
close-up view of A, showing the detailed U-shaped Zoophycos (Z) and scratch traces (Sp). C, close-up view of A, showing the gently
curved scratch traces (Sp) and J-shaped Zoophycos (Z) and some vertical shafts (Sk).

LETHAIA 10.1111/let.12146

tunnel systems (Figs 3, 5F), the second are simple

planar to more complex spreiten with a marginal
tube (Fig. 5A), the third are vertical shafts cross-cutting the scratch traces and Zoophycos (Figs 5C, 6C),
and the last are small Chondrites attached to the surface of Zoophycos spreiten (Fig. 5C, E).

Scratch traces (Spongeliomorpha)

The straight to gently curved scratch traces are preserved in convex hyporeliefs within the sand-filled
burrows at the bases of the sandstone units (Figs 2B,
3, 5F). The scratches form rhomboidal or net-like
patterns (Fig. 5F). The tunnel systems with scratches
are horizontal or sub-horizontal, although some
other small oblique and planar tunnels (Fig. 5F) and
vertical shafts (Fig. 3C) can be seen cross-cutting the
scratched tunnels. The scratched tunnels are commonly curved and may display an irregular sinuous
pattern (Fig. 3C). Sharp tapering terminations are
seen on one side of tunnel. The cross-section of the
tunnel is sub-circular to ovate, with tunnel width
between 2 and 4 cm. The tunnels are commonly
infilled with fine sand. The tunnels co-occur with

Zoophycos from the Lower Devonian of South China

Zoophycos and are located adjacent to the marginal

tubes (Fig. 3B).

Zoophycos
The studied interval is locally crowded with Zoophycos that vary in dimensions and complexity. Zoophycos is also preserved as convex hyporeliefs on the
soles of the sandstone units (Figs 2, 3). The density
of burrows is very high, and the J-shaped and Ushaped Zoophycos spreiten were only occasionally
identified (Fig. 3). The width of the spreiten usually
ranges from 5 to 21 cm and the diameter of the marginal tube varies from 1.5 to 2.5 cm. The cross-sectional shape of the marginal tube is ovate in vertical
view. The Zoophycos spreiten observed are protrusive. The spreiten occur in the deepest tier of the ichnofabric and are cross-cut by vertical shafts (Figs 4,
5). They intergrade with Palaeophycus and Chondrites on the burrow walls (Figs 5C, E, 6A, C).
A well preserved U-shaped Zoophycos spreite,
which originated by gradual extension of the simple
spreiten and constructed upwards (Fig. 4), was carefully identified. It can be up to 28 cm in width and

Fig. 4. Specimen exhibiting the relationship between the upper and bottom surfaces of Zoophycos spreiten (KBS 09/2013). The diagrammatic illustration demonstrates the contrasting components within the complex trace fossil and their relationship. The spreiten were constructed upwards and formed about three whorls. Each whorl has a marginal tube. The relationships of the complex trace fossils are best
observed in the bottom view. Sk, Ch and Pa represent vertical shafts, Chondrites and Palaeophycus, respectively. Sp and Ps represent
scratch traces and some planar and vertical shafts, respectively. W1, W2 and W3 refer to the first, second and third whorls following the
construction upwards.

L.-J. Zhang & Z. Zhao

LETHAIA 10.1111/let.12146

Fig. 5. Photographs showing the detailed features on the bottom surface of Zoophycos (KBS 09/2013). A, Zoophycos associated with adjacent scratch traces. B, scratches on the wall of the lamellae. C, close-up the middle part of A, spreiten consist of primary lamellae (Pl)
cross-cut by vertical shafts (e.g. Skolithos (Sk), Bifungites (Bi)) and associated with small dark Chondrites (Ch). D, close-up of A, first tube
(marginal tube) (Ft) and subsequent marginal tube (Mt). S represents the vertical shafts Skolithos. E, Bifungites (Bi) and Chondrites (Ch)
on the bottom surface of Zoophycos spreiten. F, detailed photograph showing the rhomboidal or net-like scratch traces (Sp) on the bottom surface. Some planar and vertical tunnels (Ps) also associated with scratch traces.

43 cm in length and is circular to elongate in outline

(Fig. 4). The complex Zoophycos spreiten were constructed along the bedding plane to a greater extent
than the simple ones. Branching of spreiten is
absent, but parts tend to be abandoned to build new
spreiten. According to the planar and cross-sectional
views, three whorls were identified (Figs 3, 6A).
They are displayed as thin flat spreiten. Each whorl
had an individual marginal tube (Fig. 5A). In a
cross-sectional view, Zoophycos was composed of
closely spaced concave spreite laminations terminating with a marginal tube (Fig. 6D). The filling materials within the spreiten are similar to the matrix
immediately adjacent to them, although texture and
cementation are different. No faeces are found
within the spreiten.

Scratch features on the walls of marginal tubes are

common (Figs 5B, 6D), but the scratches are strialike, not similar to the adjacent rhomboidal scratch
traces of Spongliomorpha (Fig. 5F).

Vertical shafts and other associated traces

Small (ca 1 mm burrow diameter) bifurcated Chondrites burrows were also found on the bottom surface
of Zoophycos spreiten (Fig. 5C, E). Dumbbell-like
burrows assigned to Bifungites also occur as convex
hyporeliefs on the soles of the Zoophycos spreiten
(Fig. 5C, E). In general, Zoophycos is frequently cut by
the associated vertical shafts which might be assigned
to Skolithos (Figs 3, 5C, D, 6B, C, D), which displays
cylindrical or sub-cylindrical, vertical, unbranched

LETHAIA 10.1111/let.12146

Zoophycos from the Lower Devonian of South China

Fig. 6. Photographs showing the detailed features on the cross-section and upper surface of Zoophycos spreiten (KBS 09/2013). A, three
whorls can be observed. B, Palaeophycus (P) and Skolithos (S) on the surface of the second whorl (W2). C, close-up view of the left part
of A, Palaeophycus (Pa) and Skolithos (Sk) occur on the surface of the spreite and marginal tube (Mt). D, two whorls can be observed on
the cross-sectional view. The concave lamellae can be found in the second whorl. Mt represents the marginal tube, and W1 and W2 are
the first and second whorls, respectively.

and unlined burrows with a diameter between 7 and

12 mm. Small Palaeophycus burrows occur on the
upper surface of Zoophycos spreiten (Fig. 6C).

Discussion
Zoophycos ethology and its
palaeoenvironmental significance
Zoophycos normally occurs from nearshore to offshore in the Devonian, which was the first bloom of
this trace fossil in the Palaeozoic (Miller 1991; Neto
de Carvalho & Rodriguez 2003; Gaillard & Racheboeuf 2006; Seilacher 2007; Zhang 2014). The
Zoophycos producer was able to exhibit various
behaviours (Bromley 1991; Knaust 2009a). The
Zoophycos studied here mainly belongs to U-shaped
spreiten (Fig. 3), which were formed in the mixed
layer (soft to firm substrate) together with scratch
traces (Spongeliomorpha), vertical shafts (Skolithos),
Chondrites, Bifungites and Palaeophycus. The morphology of the scratch traces (Figs 3, 5F) is similar
to the Spanish Miocene Spongeliomorpha burrows
described by de Gibert & Ekdale (2010), where the
sharp ridges on the walls form a rhomboidal pattern

and the tunnels have a very characteristic tapering

termination. Spongeliomorpha is typically produced
by decapod crustaceans (Ekdale & de Gibert 2010;
de Gibert & Ekdale 2010), where the trace maker
with rigid appendages is capable of scratching the
firm substrate. The average width of the marginal
tube is about 2 cm. The scratches are also found on
the wall of the marginal tube, which indicates that
the Zoophycos might have been partly reburrowed by
the decapod crustacean in the firm substrate. The
scratch traces probably served for feeding (fodinichnia) and/or moving (repichnia), while the Zoophycos
represents deposit feeding.
A curved tube was observed (Fig. 5D) and might
be the first position of the marginal tube following
the constructional model proposed by Olivero &
Gaillard (2007). The Zoophycos here display at most
three whorls, unlike the Zoophycos observed on the
continental slopes off Europe and West Africa,
which often consist of a large number of whorls
(Wetzel & Werner 1981; L
owemark & Schafer 2003).
The Zoophycos spreite is U-shaped in each whorl,
and the width of marginal tube is almost un-changed, which indicates that the spreite is protrusive
and produced in the bottom of the sediment, and
the subsequent U-shaped spreiten were added in an

L.-J. Zhang & Z. Zhao

upward spiral. In other words, the Zoophycos burrow

system enlarged by upward spiralling.
The Zoophycos here is also cut by the vertical
shafts (Skolithos), which represent open tunnel systems created for dwelling (domichnia). Small treelike Chondrites are found on the wall of marginal
tubes which are filled with dark-coloured muddy
materials. Small black Chondrites represent the last
trace fossil in a bioturbation sequence (Bromley &
Ekdale 1984), which indicate that after the Zoophycos-maker finished the complex burrow system, the
Chondrites-maker only colonized the filled burrows
in a later stage and reburrowed them, while tolerating dysoxic conditions.
All of the traces fossils occur at the bottom of an
erosion surface, the whole sequence of which consists of medium-bedded sandstone intercalated with
mudstone (Fig. 2). The substrate and trace fossil
analyses indicate a change from oxic to dysoxic environments after a storm or large wave surge and also
demonstrate that the trace maker of Zoophycos and
the scratch traces were opportunistic organisms (rstrategists) that thrived in a low energy nutrient garden after storm events. The Zoophycos producer
dominated the first oxic, nutritious substrate in the
bottom and burrowed upwards for feeding on the
organic matter that slowly settled on the substrate
after the storm or wave event. Subsequently, due to
the reduced oxygen content in the softground of the
sediment, Chondrites was developed in a later stage
of the storm-influenced depositional sequence. After
that, different scratch trace producers dominated the
firm substrate and partly reburrowed the Zoophycos
spreiten. In the last stage, vertical traces cross-cut the
former Zoophycos and scratch traces in a normal
nearshore environment (Fig. 7).
No faeces were founded in the spreiten, which
might indicate that their inhabitants either did not
excrete in the sediment or moved onto the seafloor
for defecation (unlike the Holocene Zoophycos
reported by Wetzel & Werner (1981) and Kotake
(1992)). The sediment-filled spreiten imply that the
animal defecated without depositing faecal pellets
within the burrow as in the Oman Zoophycos
(Knaust 2009a). Furthermore, the producer of Quaternary deep-sea Zoophycos can alternated between
pelleting behaviour and non-pelleting behaviour as
documented by Wetzel & Werner (1981), suggesting
the pelleting behaviour is not essential for the
Zoophycos producer. The U-shaped marginal tube of
the spreiten structure can provide well-aerated
microenvironment within the burrow as proposed
by Knaust (2009a).
The behaviour of Zoophycos has been debated for
a long time and still remains unresolved, resulting in

LETHAIA 10.1111/let.12146
A

Fig. 7. Schematic reconstruction of the complex trace fossil from

Ganxi, South China. A, initially, the Zoophycos producer burrowed in the softground of the sediment after a storm event and
exploited the oxic environment by constructing the trace
upwards without any scratches on the wall. In a later stage, under
nearly dysoxic conditions, Chondrites burrows originated from
the bottom of the sediment and attached to the Zoophycos spreiten. B, during the subsequent firmground phase of the sediment,
rhomboidal scratch traces were created and the spreiten became
partly reworked by the similar scratches. Subsequently, the
scratches and spreiten were partly reburrowed by vertical shafts.
The complex trace fossils are interpreted as forming from different producers, the spreiten represent the deposit-feeding behaviour of polychaetes, and the larger scratches are interpreted as
being feeding and moving structures formed by crustaceans.

a number of ethological models, such as the depositfeeding model (Wetzel & Werner 1981; Ekdale &
Lewis 1991; Kotake 1992; Olivero & Gaillard 1996;
Knaust 2009a), detritus-feeding model (Kotake
1989, 1991, 1992), refuse dump model (Bromley
1991; L
owemark & Schafer 2003; Knaust 2009b),
cache model (Fu & Werner 1995; Bromley et al.
1999; Miller & DAlberto 2001; L
owemark 2012) and
the gardening model (Bromley 1991; Bromley &
Hanken 2003; Gong et al. 2008). The Zoophycos
described here is most consistent with the depositfeeding model in which the Zoophycos-maker
exploits and farms the sediment simultaneously. The
Zoophycos-maker reworked an individual sand bed
and formed a single protrusive spreiten, after that, it
moved upwards for food access, where a nutrientrich level may have been located (Fig. 3), and the
trace maker created the spreiten but no faecal pellets
accumulated within the spreiten (Fig. 5A).

Zoophycos probable producer

Zoophycos as described in the Lower Devonian of
South China belongs to a morphological group that
is common in Palaeozoic strata (Vanuxem 1842; Seilacher 1983; Bhargava et al. 1985; Miller 1991; Gaillard & Racheboeuf 2006; Gong et al. 2008, 2010;

LETHAIA 10.1111/let.12146

Knaust 2009a; Sappenfield et al. 2012; Zhang 2014).

Until now, its ichnotaxonomic position remains
open and it shares some characteristics with Rhizocorallium commune, it is quite certain that a gradual
transition between Zoophycos and Rhizocorallium
exists (Knaust 2009a, 2013).
According to its general morphological features,
Zoophycos is usually considered to be produced by a
worm-like animal, of which several candidates have
been proposed: sipunculids (Wetzel & Werner 1981;
Olivero 2003; Olivero & Gaillard 2007), echiurids
(Kotake 1992) and polychaetes (Bischoff 1968;
Ekdale & Lewis 1991; Knaust 2009a).
The Zoophycos from South China exhibits key
characteristics that help in identifying its producer.
First, the Zoophycos spreite is not similar to the Late
Palaeozoic Zoophycos as described by Miller (1991),
Bhargava et al. (1985) and Knaust (2009a) but
resembles the French Jurassic Zoophycos (Olivero
2003) and the Libyan Devonian Zoophycos (Seilacher
1983). Miller (1991) described Zoophycos spreiten
from Devonian marine storm deposits that grade
into a horizontal meniscoid burrow whereas the
Middle Permian nearshore Zoophycos reported by
Knaust (2009a) is associated with a system of cylindrical burrows. Bhargava et al. (1985) described Rhizocorallium-like Zoophycos spreiten from Indian
Permian euxinic mud-shelf environment. The Lower
Devonian shallow-marine Zoophycos described from
South China, which shows a continuation from a
curved tube (first tube) to the marginal tube similar
to the French Zoophycos described by Olivero &
Gaillard (2007). The curved tube merges into the
marginal tubes, which suggests that the Zoophycos
spreiten were constructed upwards similar to the
French Zoophycos (Olivero 2003) and the Amuri
Zoophycos (Ekdale & Lewis 1991). From previous
studies (Miller 1991; Gong et al. 2008; Knaust
2009a; Zhang 2014), the occurrence of Zoophycos in
the Late Palaeozoic was largely reported from shallow-marine environments with storm influence.
They commonly occur in soft to firm substrates (e.g.
marlstone, mudstone and siltstone) within transgressive systems tracts (Knaust 2009a; Hu et al.
2010; Zhang 2014).
Second, scratch features are observed only on the
sole wall of marginal tubes and the Zoophycos was
later reworked producing scratch traces similar to
Spongeliomorpha, which suggests the Zoophycosmaker first dominated and thrived in the nutrientrich environment (soft substrate), and then, the
Zoophycos spreiten were reworked by crustacean
without any predation during the firm substrate episode. Third, the width of marginal tube and lamellae
is almost constant and the gradual expansion of the

Zoophycos from the Lower Devonian of South China

spreiten suggests that the Zoophycos-maker did not

grow in width during the formation of the structure,
just searched for the nutrient food step by step and
then formed the extension spreiten.
It seems, therefore, that worm-like animals, especially polychaetes, might have been the producers
for the Ganxi Zoophycos, as has been proposed for
the Amuri Zoophycos (Ekdale & Lewis 1991), the
Omani Zoophycos (Knaust 2009a) and the Greek
Zoophycos (Bischoff 1968). Sipunculans have been
repeatedly regarded as the producer of Zoophycos
(Wetzel & Werner 1981; Olivero 2003; Olivero &
Gaillard 2007), but they are much less motile, were
unable to produce complex burrow systems (Knaust
2009a) and are not consistent with the characteristics
of Zoophycos described here.

Conclusions
The trace fossil Zoophycos is described here from
shallow-marine storm deposits of the Lower Devonian Yangmaba Formation of Ganxi in South China.
It is part of a complex trace fossil and mainly consists of scratch traces, J-shaped and U-shaped spreiten with marginal tubes (Zoophycos) and some
additional trace fossils associated with Zoophycos,
including vertical shafts, Palaeophycus, Bifungites
and Chondrites.
The scratch traces mainly originated during feeding
and locomotion, whereas the Zoophycos spreiten
resulted from deposit feeding. The upward spiral Ushaped Zoophycos spreiten, the first tube (marginal
tube) and the lack of faecal pellets therein suggests an
ethology of deposit feeding by the same producers.
The burrow architectures herein suggest polychaetes might be the most likely Zoophycos producer.
A review of examples from the literature shows that
Zoophycos from the Devonian to Triassic demonstrates a similar behavioural pattern, and polychaetes
account for most Palaeozoic forms.
Acknowledgements. We gratefully acknowledge financial support for this study from the Natural Science Foundation of China
(41290260), the Specialized Research Fund for the Doctoral Program of Higher Education (20134116120002). We are indebted
to Ran Xu, Cheng Huang, Xuejiao Tan, Yuxuan Ma, Bin Liu,
Dequan Ma and Zhongxiang Li for assistance in the field. We
also gratefully thank Dr. Dirk Knaust, AE Alan Owen and an
anonymous reviewer for their insightful reviews and comments
on the manuscript.

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