A bioturbation classification of European marine infaunal
invertebrates
s1, Silvana N. R. Birchenough2, Julie Bremner2, Jasmin A. Godbold3, Ruth E. Parker2,
Ana M. Queiro
Alicia Romero-Ramirez4, Henning Reiss5,6, Martin Solan3, Paul J. Somerfield1, Carl Van Colen7,
Gert Van Hoey8 & Stephen Widdicombe1
1
Plymouth Marine Laboratory, Prospect Place, The Hoe, Plymouth, PL1 3DH, U.K.
The Centre for Environment, Fisheries and Aquaculture Science, Pakefield Road, Lowestoft, NR33 OHT, U.K.
3
Department of Ocean and Earth Science, National Oceanography Centre, University of Southampton, Waterfront Campus, European Way,
Southampton SO14 3ZH, U.K.
4
EPOC – UMR5805, Universite Bordeaux 1- CNRS, Station Marine d’Arcachon, 2 Rue du Professeur Jolyet, Arcachon 33120, France
5
Faculty of Biosciences and Aquaculture, University of Nordland, Postboks 1490, Bodø 8049, Norway
6
Department for Marine Research, Senckenberg Gesellschaft für Naturforschung, Südstrand 40, Wilhelmshaven 26382, Germany
7
Marine Biology Research Group, Ghent University, Krijgslaan 281/S8, Ghent 9000, Belgium
8
Bio-Environmental Research Group, Institute for Agriculture and Fisheries Research (ILVO-Fisheries), Ankerstraat 1, Ostend 8400, Belgium
2
Keywords
Biodiversity, biogeochemical, ecosystem
function, functional group, good
environmental status, Marine Strategy
Framework Directive, process, trait.
Correspondence
s, Plymouth Marine
Ana M. Queiro
Laboratory, Prospect Place, The Hoe,
Plymouth PL1 3DH, U.K. Tel: +44 1752 633
476; Fax: +44 1752 633 101;
E-mail: anqu@pml.ac.uk
Funding Information
Funding was supported by the Western
Channel Observatory, part of the UK Natural
Environmental Research Council’s National
s, Somerfield and
Capability (Queiro
Widdicombe), the Department of the
Environment, Food and Rural Affairs under
Science-Level Agreement SLA31
(Birchenough), the Conseil Regional
d’Aquitaine and the University of Bordeaux 1
(Romero-Ramirez), and the Flemish Fund for
Scientific Research (Van Colen).
Abstract
Bioturbation, the biogenic modification of sediments through particle reworking and burrow ventilation, is a key mediator of many important geochemical
processes in marine systems. In situ quantification of bioturbation can be
achieved in a myriad of ways, requiring expert knowledge, technology, and
resources not always available, and not feasible in some settings. Where dedicated research programmes do not exist, a practical alternative is the adoption
of a trait-based approach to estimate community bioturbation potential (BPc).
This index can be calculated from inventories of species, abundance and
biomass data (routinely available for many systems), and a functional classification of organism traits associated with sediment mixing (less available). Presently, however, there is no agreed standard categorization for the reworking
mode and mobility of benthic species. Based on information from the literature
and expert opinion, we provide a functional classification for 1033 benthic
invertebrate species from the northwest European continental shelf, as a tool to
enable the standardized calculation of BPc in the region. Future uses of this
classification table will increase the comparability and utility of large-scale
assessments of ecosystem processes and functioning influenced by bioturbation
(e.g., to support legislation). The key strengths, assumptions, and limitations of
BPc as a metric are critically reviewed, offering guidelines for its calculation and
application.
Received: 1 May 2013; Revised: 6 August
2013; Accepted: 12 August 2013
Ecology and Evolution 2013; 3(11): 3958–
3985
doi: 10.1002/ece3.769
Introduction
Marine soft-sediment habitats represent some of the most
functionally important ecosystems on Earth, being charac3958
terized by a high biomass and diversity of invertebrate
organisms that are fundamental to the mediation of a
wealth of goods and services (Lotze et al. 2006; White et al.
2010; Widdicombe and Somerfield 2012). Infaunal inverte-
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
This is an open access article under the terms of the Creative Commons Attribution License, which permits use,
distribution and reproduction in any medium, provided the original work is properly cited.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
brates exhibit significant influence over benthic sedimentary geochemical environments in soft sediments through
bioturbation, that is, the mixing of sediment and particulate materials carried out during foraging, feeding and burrow maintenance activities, and the enhancement of pore
water and solute advection during burrow ventilation
(Richter 1936; Rhoads 1974; Volkenborn et al. 2010). These
processes influence oxygen, pH and redox gradients (Stahl
et al. 2006; Pischedda et al. 2008; Queir
os et al. 2011), metal
cycling (Teal et al. 2009), sediment granulometry (Montserrat et al. 2009), pollutant release (Gilbert et al. 1994),
macrofauna diversity (Volkenborn et al. 2007), bacterial
activity and composition (Mermillod-Blondin and Rosenberg 2006; Gilbertson et al. 2012), and ultimately carbon
(Kristensen 2001) and nitrogen cycling (Bertics et al. 2010).
Hence, in light of anticipated changes to marine systems
associated with human activity (Halpern et al. 2008; Hoegh-Guldberg and Bruno 2010), large-scale assessments of
bioturbation can contribute to a better understanding of how
of ecosystem functioning is mediated by biological activity.
Community bioturbation potential (BPc) is a metric
first described by Solan et al. (2004a), which combines
abundance and biomass data with information about the
life traits of individual species or taxonomic groups. This
information describes modes of sediment reworking (Ri)
and mobility (Mi) of taxa in a dataset, two traits known
to regulate biological sediment mixing, a key component
of bioturbation (Solan 2000; and refereces therein; Solan
et al. 2004b). BPc is thus not a direct measure of the
process of bioturbation. Rather, BPc provides an estimate
of the potential of a community to bioturbate. Hence,
where macrofauna abundance and biomass data are
available, BPc provides a means to estimate the extent to
which benthic communities are likely to affect important
ecosystem properties that underpin ecosystem functioning. The consequences of environmentally driven changes
in biodiversity to BPc, and its relation to ecosystem functioning, have been explored in this way in terrestrial (Bunker et al. 2005) and marine habitats (Solan et al. 2004a,b);
at the local (Lohrer et al. 2010; Teal et al. 2013) and regional scales (Queir
os et al. 2011; Birchenough et al. 2012;
Solan et al. 2012); for different contexts (e.g., habitat
structure and hypoxia, Queir
os et al. 2011; Van Colen
et al. 2012; Villn€as et al. 2012); and for a variety of ecosystem functions including productivity (Solan et al. 2012),
nutrient cycling (Solan et al. 2004a), carbon storage (Bunker et al. 2005; Solan et al. 2012), and decomposition (Josefson et al. 2012). By calculating BPc over time, or for
different locations or scenarios, changes in the efficiency
of the organism-sediment couple can be monitored for
compliance in support of management and policy objectives (Painting et al. 2012; Van Hoey et al. 2013). For
example, the effects of simulated changes in benthic com-
Bi and Ai are the biomass and abundance of species/taxon i
in a sample. Trait scores were derived from an extensive
review of published material and expert knowledge (consensus of 12 authors), and details of the scoring system are
provided below. Species for which no published information was available were scored based on descriptions of species behavior and information on closely related species at
the nearest taxonomic level. As BPc captures information
about sediment particle reworking, pelagic species and
those living on hard substrates were not included. Sediment reworking functional types were also defined (Fti),
according to Francßois et al. (1997), Solan (2000), and
Gerino et al. (2003). Taxonomic information and Aphia
ID (a unique species database identifier) were extracted
from the World Register of Marine Species (2012).
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3959
munity structure have previously been used to explore
possible changes in ecosystem properties like sediment
organic carbon at the North Sea scale, based on empirically derived relationships between BPc and sediment
organic carbon (Fig. 1). Similar uses of BPc could invaluably contribute to an increased understanding of the role
of ecosystem structure in the sustenance of marine functioning and its resilience to human activities, an urgent
need under current European legislation (Marine Strategy
Framework Directive, 2008/56/EC).
A significant obstacle in the widespread application of
BPc, however, is the need for a standard classification
scheme that is supported by the benthic research community. As a first step in fulfilling this research gap, we present the findings of the Study Group on Climate Related
Benthic Processes in the North Sea, an expert group
appointed by the International Council for the Exploration of the Sea (ICES SGCBNS). We present the conclusions of a series of dedicated workshops tasked with
deriving a functional classification of northwest European
marine invertebrate species to facilitate the calculation of
BPc in different regions of the North Atlantic.
Methods
The classification of marine invertebrate infauna into bioturbation groups was carried out using 18 datasets compiled from northwest European waters (n = 1033 species).
Following Swift (1993) and Solan et al. (2004a), each taxon
(1) was scored on categorical scales that reflect increasing
mobility (Mi) from 1 (living in a fixed tube) to 4 (free three
dimensional movement via burrow system), and increasing
sediment reworking (Ri) from 1 (epifauna that bioturbate
at the sediment–water interface) to 5 (regenerators that excavate holes, transferring sediment at depth to the surface).
BPc ¼
n pffiffiffiffiffiffiffiffiffiffiffi
X
Bi=Ai Ai Mi Ri
i¼1
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Results
Table 1 provides the classifications for mobility (Mi) and
sediment particle reworking (Ri) assigned to the 1033
marine invertebrate species (and other taxa) from northwest European waters, and the associated sediment
reworking functional types (Fti). Please refer to the table
for details of the scoring criteria.
Discussion
Figure 1. Percentage of sediment organic carbon (circles, diameter
scaled to %) at each of 109 sites sampled during the North sea
benthos Survey conducted by the Benthos Ecology Working Group of
the International Council for the Exploration of the Sea in 1986 (left
column, “100%”), and following the implementation of simulated
trait based extinction scenarios (right column, “50%”). By combining
measurements of ecosystem functions with information on relevant
species traits (abundance, biomass, functional group and behaviour),
empirically derived relationships between specific ecosystem functions
and BPc can be derived. In this case, the relationship between BPc and
sediment % of organic carbon. Species extinction scenarios can be
simulated and the consequential changes in ecosystem functioning
recalculated based on changes in community composition and/or
structure following implementation of each scenario. In this example,
the predicted levels of sediment organic carbon content are presented
for a 50% reduction in species richness (right column, “50%”)
following sequential local expiration of species ordered by the most
abundant species within a site (top row), those with the largest
biomass (second row from top), or by the most abundant species
across the region (bottom row). The consequences for ecosystem
functioning (here, sediment organic carbon content) following these
ordered extinction scenarios contrast to a scenario where species are
extirpated in a random order (third row from top). Implementing
various trait-based extinction scenarios in this way provides insights
on possible outcomes following, for example, changes in
management or as a result of anthropogenic forcing. Modified from
Solan et al. (2012).
As with any functional classification that on which BPc is
calculated relies on three main assumptions. Understanding these assumptions, and the need to correct for them
where information is available, is key steps in the
adequate use of BPc as a metric:
1 If body size is constant, the BPc of a species/taxon
(BPi) is transferable across space and time. BPc
accounts for two “fixed” traits (Ri and Mi) that are
assumed to be directly related to life-history traits and
activity levels of each species, that are not altered by
context or spatiotemporal variation. Where information to the contrary is available about the alteration of
species behavior in response to external stimuli, context-specific adjustments to reworking and mobility
trait scores should be made accordingly: for example,
thermal stress (Ouellette et al. 2004; Przeslawski et al.
2009); habitat structure (Godbold et al. 2011); ocean
acidification (Godbold and Solan 2013); or presence of
a predator (Maire et al. 2010). For instance, sediment
type has been observed to be influential when determining the classification of a particular species into
one of two specific functional groups (Needham et al.
2011), but this has not been documented for the vast
majority of bioturbators. Incorporation of this type of
information could be achieved using more sophisticated routines, such as fuzzy coding, to capture the
influence of intraspecific variability in reworking and
mobility traits (Maire et al. 2007; Bremner 2008; Godbold et al. 2011) across known sources of variation
(habitat, season, food availability, etc.). The paucity of
such information present for the majority of marine
species (Tyler et al. 2012) is a source of concern and
will be needed to project potential changes in BPc
under future policy or environmental scenarios. Typical
body size is a “flexible” trait in the metric, which will
vary in response to environmental variation, seasonality, stress, and disturbance (Queir
os et al. 2006;
Macdonald et al. 2012). BPc captures this information
through changes in the biomass/abundance ratio on
which typical body size is calculated.
2 An organism’s reworking (Ri) and mobility (Mi) modes
remain the same across the life span of each individual.
3960
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Bioturbation potential allocations for 1033 macrofaunal species. Mi and Ri are the reworking and mobility traits, and Fti is the
corresponding sediment reworking functional types.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Grania
Tubificoides amplivasatus
Tubificoides insularis
Tubificoides pseudogaster
Oligochaeta
Cossura longocirrata
Chirimia biceps
Clymenura lankesteri
Euclymene
Lumbriclymene
Nicomache
Rhodine
Ophelia borealis
Ophelina cylindricaudata
Ophelina modesta
Ophelina norvegica
Travisia forbesii
Orbinia norvegica
Aricidea (Acmira) laubieri
Aricidea (Allia) roberti
Aricidea (Aricidea) wassi
Aricidea simonae
Aricidea suecica
Cirrophorus armatus
Paraonis fulgens
Polygordius
Protodriloides chaetifer
Polyphysia crassa
Paramphinome jeffreysii
Pareurythoe borealis
Euphrosine foliosa
Capitellida
Ophryotrocha
Parougia caeca
Parougia eliasoni
Parougia nigridentata
Protodorvillea kefersteini
Schistomeringos
Schistomeringos rudolphii
Eunice
Eunice dubitata
Eunice pennata
Eunicidae
Lysidice ninetta
Marphysa
Marphysa bellii
Nematonereis unicornis
Abyssoninoe
Abyssoninoe hibernica
Lumbrineriopsis paradoxa
Lumbrineris
Lumbrineris agastos
Lumbrineris aniara
Lumbrineris fragilis
Lumbrineris gracilis
369702
137570
137578
137582
2036
129984
130277
130284
129347
129350
129357
129363
130491
465714
130507
130508
130512
156278
326587
326598
326605
130570
130572
152339
146932
129472
130837
130977
129837
129838
130083
890
129266
130036
130037
130039
130041
129274
154127
129278
130055
130060
966
130071
129281
130072
594957
129331
146469
130235
129337
130236
130238
152285
130244
4
4
4
4
4
2
3
3
3
3
3
3
4
4
4
4
4
4
2
2
2
2
2
2
2
2
2
4
4
4
2
3
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
3
3
3
3
3
3
2
1
1
1
1
1
3
3
3
3
3
3
3
3
3
3
3
3
3
2
2
4
3
3
3
2
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
B
B
B
B
B
S
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC
B
B
B
B
B
B
S
S
S
S
S
S
S
S
S
B
B
B
S
UC
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Clitellata
Clitellata
Clitellata
Clitellata
Clitellata
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Enchytraeida
Haplotaxida
Haplotaxida
Haplotaxida
Caenogastropoda
Tubificidae
Tubificidae
Tubificidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Amphinomida
Amphinomida
Amphinomida
Capitellida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Cossuridae
Maldanidae
Maldanidae
Maldanidae
Maldanidae
Maldanidae
Maldanidae
Opheliidae
Opheliidae
Opheliidae
Opheliidae
Opheliidae
Orbiniidae
Paraonidae
Paraonidae
Paraonidae
Paraonidae
Paraonidae
Paraonidae
Paraonidae
Polygordiidae
Protodriloididae
Scalibregmatidae
Amphinomidae
Amphinomidae
Euphrosinidae
Dorvilleidae
Dorvilleidae
Dorvilleidae
Dorvilleidae
Dorvilleidae
Dorvilleidae
Dorvilleidae
Eunicidae
Eunicidae
Eunicidae
Eunicidae
Eunicidae
Eunicidae
Eunicidae
Eunicidae
Lumbrineridae
Lumbrineridae
Lumbrineridae
Lumbrineridae
Lumbrineridae
Lumbrineridae
Lumbrineridae
Lumbrineridae
3961
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Lumbrineris latreilli
Lumbrineris magnidentata
Ninoe armoricana
Scoletoma
Scoletoma impatiens
Arabella
Arabella iricolor
Drilonereis
Drilonereis filum
Hyalinoecia tubicola
Nothria conchylega
Nothria hyperborea
Onuphis
Paradiopatra \quadricuspis
Eunicida
Eupanthalis kinbergi
Aphrodita aculeata
Aphroditidae
Glycera
Glycera alba
Glycera celtica
Glycera fallax
Glycera gigantea
Glycera lapidum
Glycera oxycephala
Glycera rouxii
Glycera tesselata
Glycera tridactyla
Glycera unicornis
Glyceridae
Glycinde nordmanni
Goniada
Goniada maculata
Goniadella
Goniadella gracilis
Gyptis
Hesionidae
Kefersteinia cirrata
Microphthalmus
Microphthalmus similis
Nereimyra punctata
Oxydromus agilis
Oxydromus flexuosus
Podarkeopsis capensis
Psamathe fusca
Syllidia armata
Aglaophamus agilis
Nephtyidae
Nephtys
Nephtys assimilis
Nephtys caeca
Nephtys cirrosa
Nephtys hombergii
Nephtys hystricis
Nephtys incisa
Nephtys kersivalensis
130248
155360
130254
129340
130263
129199
129854
129200
129856
130464
130467
181500
129404
130480
895
129735
129840
938
129296
130116
130119
336908
130122
130123
130126
130127
130128
130130
130131
952
130136
129300
130140
129301
130145
129307
946
130164
129313
130176
130185
710683
710680
130195
152249
130198
130343
956
129370
130353
130355
130357
130359
130360
130362
130363
4
4
4
4
4
4
4
4
4
2
2
2
2
2
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
2
2
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
B
B
B
B
B
B
B
B
B
S
S
S
S
S
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
S
S
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Eunicida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Lumbrineridae
Lumbrineridae
Lumbrineridae
Lumbrineridae
Lumbrineridae
Oenonidae
Oenonidae
Oenonidae
Oenonidae
Onuphidae
Onuphidae
Onuphidae
Onuphidae
Onuphidae
3962
Acoetidae
Aphroditidae
Aphroditidae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Glyceridae
Goniadidae
Goniadidae
Goniadidae
Goniadidae
Goniadidae
Hesionidae
Hesionidae
Hesionidae
Hesionidae
Hesionidae
Hesionidae
Hesionidae
Hesionidae
Hesionidae
Hesionidae
Hesionidae
Nephtyidae
Nephtyidae
Nephtyidae
Nephtyidae
Nephtyidae
Nephtyidae
Nephtyidae
Nephtyidae
Nephtyidae
Nephtyidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Nephtys longosetosa
Nephtys paradoxa
Nereididae
Alitta succinea
Ceratocephale loveni
Eunereis elittoralis
Eunereis longissima
Hediste diversicolor
Micronereis variegata
Neanthes fucata
Neanthes irrorata
Nereis
Nereis pelagica
Nereis zonata
Websterinereis glauca
Paralacydonia paradoxa
Pholoe
Pholoe assimilis
Pholoe baltica
Pholoe inornata
Pholoe minuta
Pholoe pallida
Chaetoparia nilssoni
Eteone
Eteone barbata
Eteone flava
Eteone foliosa
Eteone longa
Eulalia
Eulalia mustela
Eulalia viridis
Eumida
Eumida bahusiensis
Eumida sanguinea
Hesionura elongata
Hypereteone foliosa
Mysta picta
Nereiphylla rubiginosa
Paranaitis kosteriensis
Phyllodoce
Phyllodoce (Anaitides)
groenlandica
Phyllodoce groenlandica
Phyllodoce laminosa
Phyllodoce lineata
Phyllodoce longipes
Phyllodoce maculata
Phyllodoce mucosa
Phyllodoce rosea
Phyllodocidae
Pirakia punctifera
Pseudomystides limbata
Sige fusigera
Ancistargis hamata
Ancistrosyllis groenlandica
Glyphohesione klatti
130364
130365
22496
234850
130367
130374
130375
152302
130380
130387
130389
129379
130404
130407
130426
130545
129439
130598
130599
130601
130603
130604
130610
129443
231870
130613
130614
130616
129445
130631
130639
129446
130641
130644
130649
152250
147026
130659
130662
129455
130668
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
2
2
2
2
2
2
2
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
3
3
4
4
3
4
4
4
4
4
4
4
4
4
4
2
2
2
2
2
2
2
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
S
S
S
S
S
S
S
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Nephtyidae
Nephtyidae
Nereidae
Nereididae
Nereididae
Nereididae
Nereididae
Nereididae
Nereididae
Nereididae
Nereididae
Nereididae
Nereididae
Nereididae
Nereididae
Paralacydoniidae
Pholoidae
Pholoidae
Pholoidae
Pholoidae
Pholoidae
Pholoidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
334506
130670
334508
130673
334510
334512
334514
931
147104
130683
130690
130692
130695
130696
4
4
4
4
4
4
4
4
4
4
4
4
4
4
3
3
3
3
3
3
3
3
3
3
3
3
3
3
B
B
B
B
B
B
B
B
B
B
B
B
B
B
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Phyllodocidae
Pilargidae
Pilargidae
Pilargidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3963
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Pilargis verrucosa
Sigambra parva
Pisione remota
Acholoe squamosa
Alentia gelatinosa
Enipo elisabethae
Enipo kinbergi
Enipo kinbergi
Eunoe nodosa
Gattyana amondseni
Gattyana cirrhosa
Harmothoe
Harmothoe extenuata
Harmothoe
fraserthomsoni
Harmothoe glabra
Harmothoe impar
Lepidonotus
squamatus
Malmgrenia
Malmgrenia andreapolis
Malmgreniella
Malmgreniella arenicolae
Malmgreniella
castanea
Malmgreniella
darbouxi
Malmgreniella lilianae
Malmgreniella
ljungmani
Malmgreniella
lunulata
Malmgreniella
mcintoshi
Malmgreniella
polypapillata
Polynoidae
Polynoinae
Labioleanira yhleni
Neoleanira tetragona
Sigalion mathildae
Sigalionidae
Sthenelais
Sthenelais boa
Sthenelais limicola
Ephesiella abyssorum
Sphaerodorum gracilis
Erinaceusyllis
erinaceus
Eurysyllis tuberculata
Eusyllis assimilis
Eusyllis blomstrandi
Exogone (Exogone)
naidina
Exogone (Exogone)
verugera
130700
130702
130707
146474
130722
130737
130738
130738
130745
130748
130749
129491
130762
130764
4
4
4
4
4
4
4
4
4
4
4
4
4
4
3
3
3
3
3
3
3
3
3
3
3
3
3
3
B
B
B
B
B
B
B
B
B
B
B
B
B
B
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Pilargidae
Pilargidae
Pisionidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
571832
130770
130801
4
4
1
3
3
1
B
B
E
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Polynoidae
Polynoidae
Polynoidae
147006
147008
129499
130810
130811
4
4
4
4
4
3
3
3
3
3
B
B
B
B
B
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Polynoidae
Polynoidae
Polynoidae
Polynoidae
Polynoidae
130812
4
3
B
Annelida
Polychaeta
Phyllodocida
Polynoidae
130814
130815
4
4
3
3
B
B
Annelida
Annelida
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Polynoidae
Polynoidae
130816
4
3
B
Annelida
Polychaeta
Phyllodocida
Polynoidae
130818
4
3
B
Annelida
Polychaeta
Phyllodocida
Polynoidae
236712
4
3
B
Annelida
Polychaeta
Phyllodocida
Polynoidae
939
155091
131067
131069
131072
943
129595
131074
131077
131081
131100
195953
4
4
4
4
4
4
4
4
4
4
4
2
3
3
3
3
3
3
3
3
3
3
3
3
B
B
B
B
B
B
B
B
B
B
B
S
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Polynoidae
Polynoidae
Sigalionidae
Sigalionidae
Sigalionidae
Sigalionidae
Sigalionidae
Sigalionidae
Sigalionidae
Sphaerodoridae
Sphaerodoridae
Syllidae
131288
131289
131290
131304
4
4
4
4
3
3
3
3
B
B
B
B
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Syllidae
Syllidae
Syllidae
Syllidae
131307
4
3
B
Annelida
Polychaeta
Phyllodocida
Syllidae
3964
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Exogone (Parexogone)
hebes
Myrianida
Myrianida prolifera
Odontosyllis
ctenostoma
Odontosyllis gibba
Pionosyllis
Proceraea
Prosphaerosyllis
tetralix
Sphaerosyllis bulbosa
Sphaerosyllis hystrix
Sphaerosyllis taylori
Streptodonta
pterochaeta
Streptosyllis websteri
Syllidae
Syllides benedicti
Syllis
Syllis armillaris
Syllis cornuta
Syllis gracilis
Syllis hyalina
Syllis prolifera
Trypanosyllis
(Trypanosyllis)
coeliaca
Trypanosyllis zebra
Tomopteris
(Johnstonella)
helgolandica
Galathowenia
oculata
Myriochele
Myriochele heeri
Owenia fusiformis
Oweniidae
Sabellaria spinulosa
Sabellariidae
Branchiomma
bombyx
Chone
Chone acustica
Chone duneri
Euchone
Euchone rubrocincta
Euchone southerni
Jasmineira candela
Jasmineira caudata
Jasmineira elegans
Paradialychone
filicaudata
Potamethus murrayi
Potamilla reniformis
Pseudopotamilla reniformis
131302
4
3
B
Annelida
Polychaeta
Phyllodocida
Syllidae
129659
238200
131325
2
2
4
2
2
3
S
S
B
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Syllidae
Syllidae
Syllidae
131328
129669
129671
195980
4
4
2
2
3
3
2
3
B
B
S
S
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Syllidae
Syllidae
Syllidae
Syllidae
131379
131388
131394
238207
2
2
2
4
3
3
3
3
S
S
S
B
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Syllidae
Syllidae
Syllidae
Syllidae
131402
948
131405
129680
131415
157583
131435
131436
131452
335151
2
2
2
2
2
4
2
2
2
2
3
3
3
3
3
3
3
3
3
3
S
S
S
S
S
B
S
S
S
S
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Phyllodocida
Syllidae
Syllidae
Syllidae
Syllidae
Syllidae
Syllidae
Syllidae
Syllidae
Syllidae
Syllidae
131467
334946
2
1
3
2
S
E
Annelida
Annelida
Polychaeta
Polychaeta
Phyllodocida
Phyllodocida
Syllidae
Tomopteridae
146950
2
1
S
Annelida
Polychaeta
Sabellida
Oweniidae
129426
130542
130544
975
130867
979
130878
2
2
2
2
1
1
2
1
1
1
1
1
1
2
S
S
S
S
E
E
S
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Oweniidae
Oweniidae
Oweniidae
Oweniidae
Sabellariidae
Sabellariidae
Sabellidae
129525
130885
130888
129528
130909
130910
130919
130920
130921
558743
2
2
2
2
2
2
2
2
2
2
1
1
1
1
1
1
1
1
1
1
S
S
S
S
S
S
S
S
S
S
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Sabellida
Sabellidae
Sabellidae
Sabellidae
Sabellidae
Sabellidae
Sabellidae
Sabellidae
Sabellidae
Sabellidae
Sabellidae
416508
155389
130963
2
2
2
1
1
1
S
S
S
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Sabellida
Sabellida
Sabellida
Sabellidae
Sabellidae
Sabellidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3965
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Sabella pavonina
Sabellidae
Apistobranchus
Apistobranchus tenuis
Apistobranchus
tullbergi
Chaetopteridae
Chaetopterus
variopedatus
Spiochaetopterus
costarum
Spiochaetopterus
typicus
Magelona
Magelona alleni
Magelona filiformis
Magelona johnstoni
Magelona minuta
Magelona mirabilis
Magelonidae
Poecilochaetidae
Poecilochaetus
serpens
Aonides
Aonides oxycephala
Aonides
paucibranchiata
Aurospio banyulensis
Dipolydora caulleryi
Dipolydora coeca
Dipolydora socialis
Laonice
Laonice bahusiensis
Laonice cirrata
Laonice sarsi
Malacoceros
fuliginosus
Microspio atlantica
Minuspio cirrifera
Paraprionospio
pinnata
Paraspio decorata
Polydora
Polydora ciliata
Polydora cornuta
Prionospio
Prionospio caspersi
Prionospio cirrifera
Prionospio dubia
Prionospio ehlersi
Prionospio fallax
Prionospio malmgreni
Prionospio
multibranchiata
Prionospio saldanha
Prionospio sexoculata
130967
985
129198
129850
129851
2
2
2
2
2
1
1
3
3
3
S
S
S
S
S
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Sabellida
Sabellida
Spionida
Spionida
Spionida
Sabellidae
Sabellidae
Apistobranchidae
Apistobranchidae
Apistobranchidae
918
129914
3
3
1
1
UC/DC
UC/DC
Annelida
Annelida
Polychaeta
Polychaeta
Spionida
Spionida
Chaetopteridae
Chaetopteridae
129922
3
1
UC/DC
Annelida
Polychaeta
Spionida
Chaetopteridae
129924
3
1
UC/DC
Annelida
Polychaeta
Spionida
Chaetopteridae
129341
130266
130268
130269
130270
130271
914
916
130711
2
2
2
2
2
2
2
2
2
2
1
2
2
2
2
2
2
2
S
S
S
S
S
S
S
S
S
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Magelonidae
Magelonidae
Magelonidae
Magelonidae
Magelonidae
Magelonidae
Magelonidae
Poecilochaetidae
Poecilochaetidae
129605
131106
131107
3
3
3
2
2
2
UC/DC
UC/DC
UC/DC
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Spionida
Spionida
Spionida
Spionidae
Spionidae
Spionidae
146532
131116
131117
131124
129613
131127
131128
131129
131131
3
4
3
4
3
3
3
3
3
2
3
1
3
1
1
1
1
2
UC/DC
B
UC/DC
B
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
131137
152392
131140
3
3
3
2
2
2
UC/DC
UC/DC
UC/DC
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Spionida
Spionida
Spionida
Spionidae
Spionidae
Spionidae
334397
129619
131141
131143
129620
131152
131153
131155
131156
131157
131159
131160
3
3
3
3
3
3
3
3
3
3
3
3
2
1
1
1
2
2
2
2
2
2
2
2
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
338540
131163
3
3
2
2
UC/DC
UC/DC
Annelida
Annelida
Polychaeta
Polychaeta
Spionida
Spionida
Spionidae
Spionidae
3966
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Prionospio steenstrupi
Pseudopolydora
antennata
Pseudopolydora
paucibranchiata
Pseudopolydora
pulchra
Pygospio elegans
Scolelepis
Scolelepis (Scolelepis)
foliosa
Scolelepis (Scolelepis)
squamata
Scolelepis bonnieri
Scolelepis korsuni
Scolelepis tridentata
Spio
Spio armata
Spio filicornis
Spio goniocephala
Spio martinensis
Spio multioculata
Spionidae
Spiophanes
Spiophanes bombyx
Spiophanes kroyeri
Spiophanes wigleyi
Streblospio benedicti
Macrochaeta
clavicornis
Macrochaeta
helgolandica
Ampharete
Ampharete acutifrons
Ampharete baltica
Ampharete falcata
Ampharete
finmarchica
Ampharete grubei
Ampharetidae
Amphicteis gunneri
Amphicteis midas
Amythasides macroglossus
Anobothrus gracilis
Melinna cristata
Melinna elisabethae
Melinna palmata
Mugga wahrbergi
Pterolysippe vanelli
Samytha sexcirrata
Sosane sulcata
Aphelochaeta
Aphelochaeta
filiformis
Aphelochaeta marioni
Caulleriella
131164
131166
3
3
2
1
UC/DC
UC/DC
Annelida
Annelida
Polychaeta
Polychaeta
Spionida
Spionida
Spionidae
Spionidae
131168
3
1
UC/DC
Annelida
Polychaeta
Spionida
Spionidae
131169
3
1
UC/DC
Annelida
Polychaeta
Spionida
Spionidae
131170
129623
334741
3
3
3
1
2
2
UC/DC
UC/DC
UC/DC
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Spionida
Spionida
Spionida
Spionidae
Spionidae
Spionidae
157566
3
2
UC/DC
Annelida
Polychaeta
Spionida
Spionidae
131171
131174
131178
129625
131180
131183
131184
131185
131186
913
129626
131187
131188
131190
131191
129745
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
2
2
2
2
1
1
1
1
2
2
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
S
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Spionida
Terebellida
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Spionidae
Acrocirridae
129746
2
2
S
Annelida
Polychaeta
Terebellida
Acrocirridae
129155
129775
129776
129777
129778
3
3
3
3
3
2
2
2
2
2
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
152272
981
129784
129785
129788
129789
129804
129805
129808
129813
334692
129819
129821
129240
129937
3
3
3
3
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
1
1
1
1
2
1
2
2
2
2
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
UC/DC
S
S
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Ampharetidae
Cirratulidae
Cirratulidae
129938
129241
2
2
2
2
S
S
Annelida
Annelida
Polychaeta
Polychaeta
Terebellida
Terebellida
Cirratulidae
Cirratulidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3967
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Caulleriella alata
Caulleriella bioculata
Caulleriella killariensis
Caulleriella zetlandica
Caulleriella zetlandica
Chaetozone
Chaetozone christiei
Chaetozone gibber
Chaetozone setosa
Cirratulidae
Cirratulus
Cirratulus caudatus
Cirriformia
tentaculata
Monticellina
dorsobranchialis
Monticellina
heterochaeta
Tharyx
Fauveliopsis
Brada villosa
Diplocirrus glaucus
Flabelligeridae
Pherusa monilifera
Pherusa plumosa
Amphictene auricoma
Lagis koreni
Pectinaria
Pectinaria
(Amphictene)
auricoma
Pectinariidae
Sternaspis scutata
Amaeana trilobata
Axionice maculata
Eupolymnia nesidensis
Hauchiella tribullata
Lanice
Lanice conchilega
Lysilla loveni
Neoamphitrite affinis
Nicolea zostericola
Paramphitrite birulai
Phisidia aurea
Pista cristata
Pista lornensis
Pistella lornensis
Polycirrus
Polycirrus medusa
Streblosoma bairdi
Streblosoma
intestinale
Terebella
Terebellidae
Thelepus cincinnatus
Thelepus setosus
129943
129944
129945
129948
129948
129242
152217
129953
129955
919
129243
129957
129964
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
S
S
S
S
S
S
S
S
S
S
S
S
S
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
Cirratulidae
129972
2
2
S
Annelida
Polychaeta
Terebellida
Cirratulidae
129973
2
2
S
Annelida
Polychaeta
Terebellida
Cirratulidae
129249
129288
130099
130100
976
130112
130113
152448
152367
129437
130590
2
2
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
1
1
1
1
S
S
UC/DC
UC
UC
UC
UC
UC
UC
UC
UC
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Cirratulidae
Fauveliopsidae
Flabelligeridae
Flabelligeridae
Flabelligeridae
Flabelligeridae
Flabelligeridae
Pectinariidae
Pectinariidae
Pectinariidae
Pectinariidae
980
131242
131471
131484
131490
152389
129697
131495
131500
131502
131508
152454
131513
131516
154972
131522
129710
131531
131538
131540
3
4
3
3
1
3
3
3
3
3
3
2
3
3
3
3
3
3
3
3
1
3
1
1
1
1
1
1
1
1
1
2
1
1
1
1
1
1
1
1
UC
B
UC/DC
DC
E
DC
DC
DC
DC
DC
DC
S
DC
DC
DC
DC
DC
DC
DC
DC
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Terebellida
Pectinariidae
Sternaspidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
Terebellidae
129713
982
131543
131544
3
3
3
3
1
1
1
1
DC
DC
DC
DC
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Terebellida
Terebellida
Terebellida
Terebellida
Terebellidae
Terebellidae
Terebellidae
Terebellidae
3968
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Terebellides stroemii
Trichobranchidae
Trichobranchus
glacialis
Trichobranchus roseus
Arenicolides
Capitella
Capitella capitata
Capitella minima
Capitellidae
Heteromastus
filiformis
Mediomastus fragilis
Notomastus
Notomastus latericeus
Peresiella clymenoides
Pseudoleiocapitella
Cossuridae
Asychis biceps
Clymenella torquata
Clymenura
Euclymene
droebachiensis
Euclymene
lombricoides
Euclymene oerstedi
Heteroclymene
robusta
Johnstonia
clymenoides
Macroclymene
santandarensis
Maldanidae
Notoproctus
Praxillella affinis
Praxillella gracilis
Rhodine gracilior
Euzonus flabelligerus
Ophelia limacina
Opheliidae
Ophelina acuminata
Polyophthalmus pictus
Orbinia (Orbinia)
sertulata
Orbiniidae
Scoloplos
Scoloplos (Scoloplos)
armiger
Aedicira mediterranea
Aricidea
Aricidea (Acmira)
catherinae
Aricidea (Acmira)
cerrutii
Aricidea (Acmira)
lopezi
131573
983
131574
3
3
3
1
1
1
DC
DC
DC
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Terebellida
Terebellida
Terebellida
Trichobranchidae
Trichobranchidae
Trichobranchidae
131575
129207
129211
129876
129879
921
129884
3
3
3
3
3
3
3
1
2
2
2
2
2
2
DC
UC
UC
UC
UC
UC
UC
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Terebellida
Trichobranchidae
Arenicolidae
Capitellidae
Capitellidae
Capitellidae
Capitellidae
Capitellidae
129892
129220
129898
129906
129226
908
146523
130279
129346
130291
3
3
3
3
3
2
3
3
3
3
2
2
2
2
2
3
1
1
1
1
UC
UC
UC
UC
UC
S
UC
UC
UC
UC
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Capitellidae
Capitellidae
Capitellidae
Capitellidae
Capitellidae
Cossuridae
Maldanidae
Maldanidae
Maldanidae
Maldanidae
209899
3
1
UC
Annelida
Polychaeta
Maldanidae
130294
146978
3
3
1
1
UC
UC
Annelida
Annelida
Polychaeta
Polychaeta
Maldanidae
Maldanidae
130298
3
1
UC
Annelida
Polychaeta
Maldanidae
130301
3
1
UC
Annelida
Polychaeta
Maldanidae
923
129358
130322
130324
130330
130487
130494
924
130500
130510
334310
3
3
3
3
3
4
4
4
4
4
4
1
1
1
1
1
3
3
3
3
3
3
UC
UC
UC
UC
UC
B
B
B
B
B
B
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Maldanidae
Maldanidae
Maldanidae
Maldanidae
Maldanidae
Opheliidae
Opheliidae
Opheliidae
Opheliidae
Opheliidae
Orbiniidae
902
129425
334772
4
4
4
3
3
3
B
B
B
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Orbiniidae
Orbiniidae
Orbiniidae
130546
129430
333034
2
2
2
3
3
3
S
S
S
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Paraonidae
Paraonidae
Paraonidae
525497
2
3
S
Annelida
Polychaeta
Paraonidae
333036
2
3
S
Annelida
Polychaeta
Paraonidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3969
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Aricidea minuta
Cirrophorus
branchiatus
Levinsenia gracilis
Paradoneis lyra
Paraonidae
Polygordius
appendiculatus
Polygordius lacteus
Protodriloides
Asclerocheilus
intermedius
Scalibregma celticum
Scalibregma inflatum
Scalibregmatidae
Acidostoma
Ampelisca
Ampelisca armoricana
Ampelisca brevicornis
Ampelisca diadema
Ampelisca gibba
Ampelisca
macrocephala
Ampelisca sarsi
Ampelisca spinipes
Ampelisca tenuicornis
Ampelisca typica
Ampelisca vervecei
Byblis
Haploops tubicola
Amphilochoides
boecki
Amphilochoides
serratipes
Amphilochus
manudens
Amphilochus
neapolitanus
Gitana sarsi
Paramphilochoides
odontonyx
Ampithoe ramondi
Aora gracilis
Aora typica
Aoridae
Autonoe longipes
Microdeutopus
Argissa hamatipes
Atylus
Atylus guttatus
Atylus swammerdami
Atylus vedlomensis
Nototropis falcatus
Apherusa
Apherusa bispinosa
Apherusa clevei
130564
130576
2
2
3
3
S
S
Annelida
Annelida
Polychaeta
Polychaeta
Paraonidae
Paraonidae
130578
130585
903
130712
2
2
2
2
3
3
3
2
S
S
S
S
Annelida
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polychaeta
Paraonidae
Paraonidae
Paraonidae
Polygordiidae
130714
129513
130974
2
2
4
2
2
4
S
S
B
Annelida
Annelida
Annelida
Polychaeta
Polychaeta
Polychaeta
Polygordiidae
Protodriloididae
Scalibregmatidae
130979
130980
925
101586
101445
101888
101891
101896
101898
101908
4
4
4
2
2
2
2
2
2
2
4
4
4
3
1
1
1
1
1
1
B
B
B
S
S
S
S
S
S
S
Annelida
Annelida
Annelida
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Polychaeta
Polychaeta
Polychaeta
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Scalibregmatidae
Scalibregmatidae
Scalibregmatidae
Acidostomatidae
Ampeliscidae
Ampeliscidae
Ampeliscidae
Ampeliscidae
Ampeliscidae
Ampeliscidae
101923
101928
101930
101933
101937
101446
101958
101960
2
2
2
2
2
2
2
2
1
1
1
1
1
4
1
3
S
S
S
S
S
S
S
B
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Ampeliscidae
Ampeliscidae
Ampeliscidae
Ampeliscidae
Ampeliscidae
Ampeliscidae
Ampeliscidae
Amphilochidae
101963
2
3
B
Arthropoda
Malacostraca
Amphipoda
Amphilochidae
101967
2
3
B
Arthropoda
Malacostraca
Amphipoda
Amphilochidae
101968
2
3
B
Arthropoda
Malacostraca
Amphipoda
Amphilochidae
101977
101982
2
2
3
3
B
B
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphilochidae
Amphilochidae
102000
102012
146895
101368
102021
101471
102064
101497
102127
102131
102132
102139
101509
102160
102164
1
1
1
1
1
1
2
2
2
2
2
2
2
2
2
1
3
3
3
3
3
3
3
3
3
3
3
3
3
3
E
S
S
S
S
S
S
S
S
S
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Ampithoidae
Aoridae
Aoridae
Aoridae
Aoridae
Aoridae
Argissidae
Atylidae
Atylidae
Atylidae
Atylidae
Atylidae
Calliopiidae
Calliopiidae
Calliopiidae
3970
Order
Family
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Apherusa jurinei
Apherusa ovalipes
Caprella acanthifera
Caprella linearis
Pariambus typicus
Parvipalpus
capillaceus
Phtisica marina
Pseudoprotella
phasma
Cheirocratus
Cheirocratus
intermedius
Cheirocratus
sundevalli
Apocorophium
lacustre
Corophium
Crassicorophium
bonellii
Leptocheirus
Leptocheirus
hirsutimanus
Leptocheirus mariae
Leptocheirus
pectinatus
Leptocheirus
tricristatus
Medicorophium affine
Monocorophium
acherusicum
Monocorophium
insidiosum
Monocorophium
sextonae
Protomedeia
Peltocoxa brevirostris
Dexamine
Dexamine spinosa
Guernea (Guernea)
coalita
Tritaeta gibbosa
Dyopedos
Dyopedos
monacantha
Eusirus longipes
Haustorius arenarius
Hyperia galba
Iphimedia
Iphimedia minuta
Iphimedia obesa
Ericthonius punctatus
Jassa falcata
Siphonoecetes
(Centraloecetes)
kroyeranus
102168
102172
101822
101839
101857
101858
2
2
2
2
2
2
3
3
2
2
2
2
S
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Calliopiidae
Calliopiidae
Caprellidae
Caprellidae
Caprellidae
Caprellidae
101864
101871
2
2
2
2
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Caprellidae
Caprellidae
101669
102795
2
2
3
3
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Cheirocratidae
Cheirocratidae
102798
2
3
S
Arthropoda
Malacostraca
Amphipoda
Cheirocratidae
148594
2
4
S
Arthropoda
Malacostraca
Amphipoda
Corophiidae
101489
237004
2
2
4
4
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Corophiidae
Corophiidae
101470
102036
2
2
4
4
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Corophiidae
Corophiidae
102038
102039
2
2
4
4
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Corophiidae
Corophiidae
102041
2
4
S
Arthropoda
Malacostraca
Amphipoda
Corophiidae
423507
225814
2
2
4
4
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Corophiidae
Corophiidae
148592
2
4
S
Arthropoda
Malacostraca
Amphipoda
Corophiidae
148603
2
4
S
Arthropoda
Malacostraca
Amphipoda
Corophiidae
101574
101983
101498
102135
102137
2
2
2
2
2
4
3
3
3
3
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Corophiidae
Cyproideidae
Dexaminidae
Dexaminidae
Dexaminidae
102141
101736
103042
2
2
2
3
3
3
S
S
S
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Dexaminidae
Dulichiidae
Dulichiidae
102202
102317
103251
101554
102345
102347
102408
102431
102111
2
2
2
2
2
2
2
1
2
3
3
3
3
3
3
1
1
1
S
S
S
S
S
S
S
E
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Eusiridae
Haustoriidae
Hyperiidae
Iphimediidae
Iphimediidae
Iphimediidae
Ischyroceridae
Ischyroceridae
Ischyroceridae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3971
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Siphonoecetes
(Centraloecetes)
neapolitanus
Siphonoecetes
(Centraloecetes)
striatus
Leucothoe incisa
Leucothoe lilljeborgi
Leucothoe richiardii
Leucothoe spinicarpa
Liljeborgia
Liljeborgia kinahani
Liljeborgia pallida
Liljeborgia psaltrica
Listriella mollis
Hippomedon
bidentatus
Hippomedon
denticulatus
Hippomedon
massiliensis
Lepidepecreum
longicornis
Lysianassidae
Orchomene humilis
Orchomenella nana
Tryphosella sarsi
Tryphosella simillima
Tryphosites longipes
Acidostoma
nodiferum
Acidostoma obesum
Animoceradocus
semiserratus
Maera
Maera grossimana
Maera loveni
Maera schmidti
Othomaera othonis
Megaluropus agilis
Abludomelita
obtusata
Eriopisa elongata
Maerella tenuimana
Microprotopus
Microprotopus
maculatus
Odius carinatus
Deflexilodes acutipes
Deflexilodes gibbosus
Monoculodes
carinatus
Perioculodes
longimanus
Pontocrates
altamarinus
102112
2
1
S
Arthropoda
Malacostraca
Amphipoda
Ischyroceridae
102115
1
1
S
Arthropoda
Malacostraca
Amphipoda
Ischyroceridae
102460
102462
212784
102470
101582
102483
102485
102486
102488
102569
2
2
2
2
2
2
2
2
2
2
3
3
3
3
3
3
3
3
3
3
S
S
S
S
S
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Leucothoidae
Leucothoidae
Leucothoidae
Leucothoidae
Liljeborgiidae
Liljeborgiidae
Liljeborgiidae
Liljeborgiidae
Liljeborgiidae
Lysianassidae
102570
2
3
S
Arthropoda
Malacostraca
Amphipoda
Lysianassidae
102576
2
3
S
Arthropoda
Malacostraca
Amphipoda
Lysianassidae
102599
2
3
S
Arthropoda
Malacostraca
Amphipoda
Lysianassidae
101395
102665
102691
102771
102773
102779
102496
2
2
2
2
2
2
2
3
3
3
3
3
3
3
S
S
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Lysianassidae
Lysianassidae
Lysianassidae
Lysianassidae
Lysianassidae
Lysianassidae
Lysianassidae s.l.
102497
531364
2
2
3
3
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Lysianassidae s.l.
Maeridae
101675
102815
102820
102826
534781
102783
102788
2
2
2
2
2
2
2
3
3
3
3
3
3
2
S
S
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Maeridae
Maeridae
Maeridae
Maeridae
Maeridae
Megaluropidae
Melitidae
102807
102831
101561
102380
2
2
2
2
2
2
3
3
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Melitidae
Melitidae
Microprotopidae
Microprotopidae
102861
236538
236539
102882
1
2
2
2
3
3
3
3
E
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Odiidae
Oedicerotidae
Oedicerotidae
Oedicerotidae
102915
2
3
S
Arthropoda
Malacostraca
Amphipoda
Oedicerotidae
102916
2
3
S
Arthropoda
Malacostraca
Amphipoda
Oedicerotidae
3972
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Pontocrates arenarius
Synchelidium
haplocheles
Synchelidium
maculatum
Westwoodilla caecula
Westwoodilla
rectirostris
Gammaropsis cornuta
Gammaropsis
maculata
Gammaropsis nitida
Gammaropsis palmata
Gammaropsis sophiae
Megamphopus
cornutus
Photis longicaudata
Photis reinhardi
Harpinia
Harpinia agna
Harpinia ala
Harpinia antennaria
Harpinia crenulata
Harpinia dellavallei
Harpinia pectinata
Harpinia plumosa
Metaphoxus
Metaphoxus
pectinatus
Parametaphoxus
tulearensis
Phoxocephalus
holbolli
Pleustidae
Bathyporeia
Bathyporeia elegans
Bathyporeia
guilliamsoniana
Bathyporeia nana
Bathyporeia pelagica
Bathyporeia pilosa
Bathyporeia sarsi
Bathyporeia tenuipes
Scopelocheirus hopei
Stegocephaloides
christianiensis
Metopa
Metopa alderi
Metopa latimana
Stenothoe marina
Austrosyrrhoe
fimbriatus
Unciola crenatipalma
Unciola planipes
Tmetonyx cicada
Urothoe
102918
102924
2
2
3
2
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Oedicerotidae
Oedicerotidae
102928
2
2
S
Arthropoda
Malacostraca
Amphipoda
Oedicerotidae
102932
102937
2
2
3
3
B
B
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Oedicerotidae
Oedicerotidae
148545
102364
2
2
2
2
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Photidae
Photidae
102367
102369
102371
102377
1
2
2
2
2
2
2
3
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Photidae
Photidae
Photidae
Photidae
102383
102387
101716
102957
102958
102960
102963
102966
102972
102973
101720
102983
2
2
2
2
2
2
2
2
2
2
2
2
1
1
3
3
3
3
3
3
3
3
3
3
S
S
S
S
S
S
S
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Photidae
Photidae
Phoxocephalidae
Phoxocephalidae
Phoxocephalidae
Phoxocephalidae
Phoxocephalidae
Phoxocephalidae
Phoxocephalidae
Phoxocephalidae
Phoxocephalidae
Phoxocephalidae
549079
2
3
S
Arthropoda
Malacostraca
Amphipoda
Phoxocephalidae
102989
2
3
S
Arthropoda
Malacostraca
Amphipoda
Phoxocephalidae
101404
101742
103058
103060
2
2
2
2
3
3
3
3
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Pleustidae
Pontoporeiidae
Pontoporeiidae
Pontoporeiidae
103064
103066
103068
103073
103076
102720
103102
2
2
2
2
2
2
2
3
3
3
3
3
3
3
S
S
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Pontoporeiidae
Pontoporeiidae
Pontoporeiidae
Pontoporeiidae
Pontoporeiidae
Scopelocheiridae
Stegocephalidae
101764
103116
103125
103166
103179
2
2
2
2
2
3
3
3
3
3
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Stenothoidae
Stenothoidae
Stenothoidae
Stenothoidae
Synopiidae
102057
102061
102736
101789
2
2
2
2
3
3
1
3
B
B
B
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Unciolidae
Unciolidae
Uristidae
Urothoidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3973
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Urothoe brevicornis
Urothoe elegans
Urothoe marina
Urothoe poseidonis
Bodotria scorpioides
Iphinoe
Iphinoe serrata
Iphinoe tenella
Iphinoe trispinosa
Diastylidae
Diastylis
Diastylis bradyi
Diastylis cornuta
Diastylis laevis
Diastylis lucifera
Diastylis neapolitana
Diastylis rathkei
Diastylis rugosa
Diastyloides biplicatus
Hemilamprops roseus
Lamprops fasciatus
Eudorella emarginata
Eudorella truncatula
Eudorellopsis deformis
Leucon (Leucon)
nasica
Cumella (Cumella)
pygmaea
Nannastacus
unguiculatus
Monopseudocuma
gilsoni
Pseudocuma
Pseudocuma
(Pseudocuma)
longicorne
Pseudocuma
(Pseudocuma) simile
Alpheus glaber
Athanas nitescens
Aristaeomorpha
foliacea
Atelecyclus
rotundatus
Calocaris macandreae
Callianassa
Callianassa
subterranea
Callianassidae
Pestarella tyrrhena
Corystes cassivelaunus
Crangon
Crangon allmanni
Crangon crangon
Crangonidae
Philocheras bispinosus bispinosus
103226
103228
103233
103235
110445
110391
110460
110461
110462
110380
110398
110472
110474
110481
110483
110484
110487
110488
110494
110514
110516
110524
110535
110536
110618
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
2
3
3
3
3
3
3
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Amphipoda
Amphipoda
Amphipoda
Amphipoda
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Cumacea
Urothoidae
Urothoidae
Urothoidae
Urothoidae
Bodotriidae
Bodotriidae
Bodotriidae
Bodotriidae
Bodotriidae
Diastylidae
Diastylidae
Diastylidae
Diastylidae
Diastylidae
Diastylidae
Diastylidae
Diastylidae
Diastylidae
Diastylidae
Lampropidae
Lampropidae
Leuconidae
Leuconidae
Leuconidae
Leuconidae
110567
2
3
S
Arthropoda
Malacostraca
Cumacea
Nannastacidae
110574
2
3
S
Arthropoda
Malacostraca
Cumacea
Nannastacidae
422916
2
3
S
Arthropoda
Malacostraca
Cumacea
Pseudocumatidae
110427
110627
2
2
3
3
S
S
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Cumacea
Cumacea
Pseudocumatidae
Pseudocumatidae
110628
2
3
S
Arthropoda
Malacostraca
Cumacea
Pseudocumatidae
107477
107486
158326
4
2
1
4
2
4
B
S
E
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Decapoda
Decapoda
Decapoda
Alpheidae
Alpheidae
Aristeidae
107273
5
4
R
Arthropoda
Malacostraca
Decapoda
Atelecyclidae
107726
107072
107729
4
4
4
4
4
4
B
B
B
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Decapoda
Decapoda
Decapoda
Axiidae
Callianassidae
Callianassidae
106800
238027
107277
107007
107551
107552
106782
108207
4
4
5
2
2
2
2
2
4
4
4
4
4
4
4
4
B
B
R
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Callianassidae
Callianassidae
Corystidae
Crangonidae
Crangonidae
Crangonidae
Crangonidae
Crangonidae
3974
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Philocheras trispinosus
Diogenes pugilator
Galathea
Galathea dispersa
Galathea intermedia
Eualus cranchii
Inachus
Inachus dorsettensis
Macropodia
Macropodia deflexa
Macropodia rostrata
Jaxea nocturna
Ebalia
Ebalia cranchii
Ebalia granulosa
Ebalia tuberosa
Nephrops norvegicus
Anapagurus
Anapagurus
bicorniger
Anapagurus laevis
Paguridae
Pagurus
Pagurus bernhardus
Pagurus cuanensis
Palaemon elegans
Pandalus montagui
Pilumnus hirtellus
Liocarcinus
Liocarcinus depurator
Liocarcinus holsatus
Liocarcinus marmoreus
Liocarcinus navigator
Liocarcinus pusillus
Pisidia longicornis
Carcinus maenas
Processa
Processa acutirostris
Processa canaliculata
Processa elegantula
Processa modica modica
Processa nouveli holthuisi
Solenocera
membranacea
Thia scutellata
Upogebia deltaura
Upogebia pusilla
Upogebia stellata
Upogebiidae
Anthura gracilis
Astacilla longicornis
Cirolana cranchi
Conilera cylindracea
Eurydice pulchra
Eurydice spinigera
Natatolana borealis
107562
107199
106834
107148
107150
156083
106905
107327
205077
107338
107345
107737
106889
107294
107298
107301
107254
106849
107213
2
1
4
4
4
2
1
1
1
1
1
4
1
1
1
1
4
1
1
4
4
3
3
3
4
3
3
3
3
3
4
3
3
3
3
4
4
4
S
E
B
B
B
S
E
E
E
E
E
B
E
E
E
E
B
E
E
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Crangonidae
Diogenidae
Galatheidae
Galatheidae
Galatheidae
Hippolytidae
Inachidae
Inachidae
Inachidae
Inachidae
Inachidae
Laomediidae
Leucosiidae
Leucosiidae
Leucosiidae
Leucosiidae
Nephropidae
Paguridae
Paguridae
107218
106738
106854
107232
107235
107614
107651
107418
106925
107387
107388
107390
107392
107393
107188
107381
107054
107681
107682
107684
108343
108344
107120
1
1
1
1
1
1
1
2
5
5
5
5
5
5
1
5
1
1
1
1
1
1
1
4
4
4
4
4
4
4
3
4
4
4
4
4
4
3
4
4
4
4
4
4
4
4
E
E
E
E
E
E
E
S
R
R
R
R
R
R
E
R
E
E
E
E
E
E
E
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Paguridae
Paguridae
Paguridae
Paguridae
Paguridae
Palaemonidae
Pandalidae
Pilumnidae
Polybiidae
Polybiidae
Polybiidae
Polybiidae
Polybiidae
Polybiidae
Porcellanidae
Portunidae
Processidae
Processidae
Processidae
Processidae
Processidae
Processidae
Solenoceridae
107281
107739
107741
107742
106803
118467
119024
118839
118842
118852
148637
118859
2
3
3
3
3
2
4
2
1
2
2
2
4
4
4
4
4
2
3
3
2
3
3
3
S
DC
DC
DC
DC
S
B
S
E
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Decapoda
Decapoda
Decapoda
Decapoda
Decapoda
Isopoda
Isopoda
Isopoda
Isopoda
Isopoda
Isopoda
Isopoda
Thiidae
Upogebiidae
Upogebiidae
Upogebiidae
Upogebiidae
Anthuridae
Arcturidae
Cirolanidae
Cirolanidae
Cirolanidae
Cirolanidae
Cirolanidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3975
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Gnathia maxillaris
Gnathia oxyuraea
Gnathiidae
Paragnathia formica
Idotea
Janira maculosa
Sphaeroma serratum
Gastrosaccus spinifer
Haplostylus normani
Nebalia bipes
Apseudes spinosus
Apseudes talpa
Apseudopsis latreillii
Akanthophoreus gracilis
Pseudoparatanais batei
118994
118995
118278
119001
118454
118732
118973
120020
148698
147032
136284
136285
247077
136340
136457
2
2
2
2
2
2
2
2
1
1
2
2
2
2
2
3
3
3
3
3
3
3
3
1
3
2
2
2
2
2
S
S
S
S
S
S
S
S
E
E
S
S
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Malacostraca
Isopoda
Isopoda
Isopoda
Isopoda
Isopoda
Isopoda
Isopoda
Mysida
Mysida
Nebaliacea
Tanaidacea
Tanaidacea
Tanaidacea
Tanaidacea
Tanaidacea
Tanaopsis graciloides
136458
2
2
S
Arthropoda
Malacostraca
Tanaidacea
Pseudosphyrapus
anomalus
Tanaissus lilljeborgi
Stomatopoda
Longipedia coronata
Cylindroleberis mariae
Philomedes
(Philomedes)
lilljeborgi
Philomedes brenda
Myodocopida
Achelia echinata
Callipallene brevirostris
Nymphon brevirostre
Anoplodactylus
Anoplodactylus
petiolatus
Anoplodactylus
virescens
Pycnogonida
Priapulus caudatus
Ascidiella aspersa
Eugyra arenosa
Styelidae
Branchiostoma
lanceolatum
Actinia equina
Urticina felina
Edwardsia
Edwardsia claparedii
Edwardsiidae
Scolanthus
callimorphus
Halcampa
chrysanthellum
Calliactis parasitica
Sagartia troglodytes
Actiniaria
136319
2
2
S
Arthropoda
Malacostraca
Tanaidacea
Gnathiidae
Gnathiidae
Gnathiidae
Gnathiidae
Idoteidae
Janiridae
Sphaeromatidae
Mysidae
Mysidae
Nebaliidae
Apseudidae
Apseudidae
Apseudidae
Leptognathiidae
Paratanaoidea
incertae sedis
Paratanaoidea
incertae sedis
Sphyrapidae
136486
14355
116368
238708
128460
2
4
2
2
2
2
4
3
3
4
S
B
S
S
S
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Malacostraca
Malacostraca
Maxillopoda
Ostracoda
Ostracoda
Tanaidacea
Tanaissuidae
Harpacticoida
Myodocopida
Myodocopida
Longipediidae
Cylindroleberididae
Philomedidae
127718
2104
134599
134643
150520
134592
134723
2
2
2
1
1
1
1
3
3
2
1
1
4
4
S
S
S
E
S
E
E
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Arthropoda
Ostracoda
Ostracoda
Pycnogonida
Pycnogonida
Pycnogonida
Pycnogonida
Pycnogonida
Myodocopida
Myodocopida
Pantopoda
Pantopoda
Pantopoda
Pantopoda
Pantopoda
Philomedidae
134730
1
4
E
Arthropoda
Pycnogonida
Pantopoda
Phoxichilidiidae
1302
101160
103718
103764
103450
104906
1
4
1
2
1
4
4
2
1
3
1
3
E
B
E
S
E
B
Arthropoda
Cephalorhyncha
Chordata
Chordata
Chordata
Chordata
Pycnogonida
Priapulida
Ascidiacea
Ascidiacea
Ascidiacea
Leptocardii
Phlebobranchia
Stolidobranchia
Stolidobranchia
Priapulidae
Ascidiidae
Molgulidae
Styelidae
Branchiostomidae
100803
100834
100730
100880
100665
100910
1
1
2
2
2
2
1
1
2
2
2
2
E
E
S
S
S
S
Cnidaria
Cnidaria
Cnidaria
Cnidaria
Cnidaria
Cnidaria
Anthozoa
Anthozoa
Anthozoa
Anthozoa
Anthozoa
Anthozoa
Actiniaria
Actiniaria
Actiniaria
Actiniaria
Actiniaria
Actiniaria
Actiniidae
Actiniidae
Edwardsiidae
Edwardsiidae
Edwardsiidae
Edwardsiidae
100916
2
2
S
Cnidaria
Anthozoa
Actiniaria
Halcampidae
100946
100994
1360
2
1
2
2
1
2
S
E
S
Cnidaria
Cnidaria
Cnidaria
Anthozoa
Anthozoa
Anthozoa
Actiniaria
Actiniaria
Actiniaria
Hormathiidae
Sagartiidae
3976
Ammotheidae
Callipallenidae
Nymphonidae
Phoxichilidiidae
Phoxichilidiidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Cerianthus
Cerianthus lloydii
Pteroeides griseum
Virgularia mirabilis
Asterias rubens
Astropecten irregularis
Psammechinus miliaris
Echinocyamus pusillus
Echinidea
Brissopsis lyrifera
Echinocardium
Echinocardium cordatum
Echinocardium
flavescens
Spatangus purpureus
Spatangoida
Labidoplax buskii
Leptosynapta
Leptosynapta
bergensis
Leptosynapta
inhaerens
Oestergrenia digitata
Synaptidae
Leptopentacta
Leptopentacta elongata
Leptopentacta elongata
Leptopentacta
tergestina
Thyone fusus
Amphilepis norvegica
Acrocnida brachiata
Amphipholis squamata
Amphiura
Amphiura
(Ophiopeltis)
securigera
Amphiura chiajei
Amphiura filiformis
Ophiocomina nigra
Ophiothrix fragilis
Ophiocten affinis
Ophiura
Ophiura albida
Ophiura ophiura
Ophiuridae
Maxmuelleria gigas
Echiurus echiurus
Thalassema
thalassemum
Echiurida
Echiura
Glandiceps talaboti
Enteroptneusta
Ensis magnus
100782
283798
181504
128539
123776
123867
124319
124273
510534
124373
123426
124392
124394
2
2
2
2
1
2
1
2
4
4
4
4
4
1
1
2
2
3
3
3
3
3
3
3
3
3
S
S
S
S
E
S
E
S
B
B
B
B
B
Cnidaria
Cnidaria
Cnidaria
Cnidaria
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Anthozoa
Anthozoa
Anthozoa
Anthozoa
Asteroidea
Asteroidea
Echinoidea
Echinoidea
Echinoidea
Echinoidea
Echinoidea
Echinoidea
Echinoidea
Ceriantharia
Ceriantharia
Pennatulacea
Pennatulacea
Forcipulatida
Paxillosida
Camarodonta
Clypeasteroida
Echinoida
Spatangoida
Spatangoida
Spatangoida
Spatangoida
Cerianthidae
Cerianthidae
Pennatulidae
Virgulariidae
Asteriidae
Astropectinidae
Parechinidae
Echinocyamidae
124418
123106
124455
123449
124462
4
4
2
2
2
3
3
3
3
3
B
B
S
S
S
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinoidea
Echinoidea
Holothuroidea
Holothuroidea
Holothuroidea
Spatangoida
Spatangoida
Apodida
Apodida
Apodida
Synaptidae
Synaptidae
Synaptidae
124465
2
3
S
Echinodermata
Holothuroidea
Apodida
Synaptidae
152547
123182
123481
124635
124635
124636
2
2
2
2
2
2
3
3
3
3
3
3
S
S
S
S
S
S
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Holothuroidea
Holothuroidea
Holothuroidea
Holothuroidea
Holothuroidea
Holothuroidea
Apodida
Apodida
Dendrochirotida
Dendrochirotida
Dendrochirotida
Dendrochirotida
Synaptidae
Synaptidae
Cucumariidae
Cucumariidae
Cucumariidae
Cucumariidae
124670
125057
236130
125064
123613
125195
2
4
4
4
4
4
3
3
3
3
3
3
S
B
B
B
B
B
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Holothuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Dendrochirotida
Ophiurida
Ophiurida
Ophiurida
Ophiurida
Ophiurida
Cucumariidae
Amphilepididae
Amphiuridae
Amphiuridae
Amphiuridae
Amphiuridae
125073
125080
125027
125131
124850
123574
124913
124929
123200
110367
110377
110375
4
4
2
2
2
2
2
2
2
5
3
3
3
3
2
2
2
2
2
2
2
4
2
2
B
B
S
S
S
S
S
S
S
R
DC
DC
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echinodermata
Echiura
Echiura
Echiura
Ophiuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Ophiuroidea
Echiuroidea
Echiuroidea
Echiuroidea
Ophiurida
Ophiurida
Ophiurida
Ophiurida
Ophiurida
Ophiurida
Ophiurida
Ophiurida
Ophiurida
Bonelliida
Echiurida
Echiurida
Amphiuridae
Amphiuridae
Ophiocomidae
Ophiotrichidae
Ophiuridae
Ophiuridae
Ophiuridae
Ophiuridae
Ophiuridae
Bonelliidae
Echiuridae
Echiuridae
110345
1269
137612
178738
160539
3
3
5
5
2
2
2
4
4
2
DC
DC
R
R
S
Echiura
Echiura
HemiChordata
HemiChordata
Mollusca
Echiuroidea
Echiurida
Enteropneusta
Enteropneusta
Bivalvia
Enteropneusta
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Brissidae
Loveniidae
Loveniidae
Loveniidae
Spatangidae
Spengelidae
Pharidae
3977
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
140103
1
2
E
Mollusca
Bivalvia
140108
1
2
E
Mollusca
Bivalvia
138333
2
2
S
Mollusca
Bivalvia
Ensis directus
140732
2
2
S
Mollusca
Bivalvia
Ensis ensis
140733
2
2
S
Mollusca
Bivalvia
Pharus legumen
140736
2
2
S
Mollusca
Bivalvia
Phaxas adriaticus
156343
2
2
S
Mollusca
Bivalvia
Phaxas pellucidus
140737
2
2
S
Mollusca
Bivalvia
139450
140291
181373
3
2
3
2
2
2
UC/DC
S
DC
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
[unassigned]
Euheterodonta
[unassigned]
Euheterodonta
[unassigned]
Euheterodonta
[unassigned]
Euheterodonta
[unassigned]
Euheterodonta
[unassigned]
Euheterodonta
[unassigned]
Euheterodonta
[unassigned]
Euheterodonta
[unassigned]
Euheterodonta
Anomalodesmata
Anomalodesmata
Anomalodesmata
138549
141644
152378
141651
140025
140253
138818
138823
138831
492197
140237
140242
140282
140283
140287
234161
234162
152905
152424
138552
141655
141659
141662
141668
141672
141676
139410
138211
140430
140431
140432
140440
140461
140467
3
3
3
3
2
1
2
2
2
2
1
1
2
2
2
3
3
2
2
3
3
3
3
3
3
3
2
2
2
2
2
2
2
2
2
2
2
2
2
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
3
2
2
DC
DC
DC
DC
S
E
S
S
S
S
E
E
S
S
S
DC
UC/DC
S
S
DC
DC
DC
DC
DC
DC
DC
S
S
S
S
S
S
S
S
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Anomalodesmata
Anomalodesmata
Anomalodesmata
Anomalodesmata
Arcoida
Arcoida
Carditoida
Carditoida
Carditoida
Carditoida
Limoida
Limoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Lucinoida
Myoida
Myoida
Myoida
Myoida
Myoida
Mytiloida
Mytiloida
Mytiloida
Hiatella arctica
Saxicavella
jeffreysi
Ensis
Cuspidaria obesa
Lyonsia norwegica
Cochlodesma
praetenue
Thracia
Thracia convexa
Thracia phaseolina
Thracia villosiuscula
Glycymeris glycymeris
Limopsis cristata
Astarte borealis
Astarte montagui
Goodallia triangularis
Arcturellina
Limatula gwyni
Limatula subauriculata
Lucinella divaricata
Lucinoma borealis
Myrtea spinifera
Axinulus croulinensis
Axinulus eumyarius
Mendicula ferruginosa
Mendicula pygmaea
Thyasira
Thyasira biplicata
Thyasira equalis
Thyasira flexuosa
Thyasira obsoleta
Thyasira sarsi
Thyasira succisa
Corbula gibba
Mya
Mya arenaria
Mya truncata
Sphenia binghami
Crenella decussata
Modiolula phaseolina
Modiolus modiolus
3978
Family
Hiatellidae
Hiatellidae
Pharidae
Pharidae
Pharidae
Pharidae
Pharidae
Pharidae
Cuspidariidae
Lyonsiidae
Periplomatidae
Thraciidae
Thraciidae
Thraciidae
Thraciidae
Glycymerididae
Limopsidae
Astartidae
Astartidae
Astartidae
Carditidae
Limidae
Limidae
Lucinidae
Lucinidae
Lucinidae
Thyasiridae
Thyasiridae
Thyasiridae
Thyasiridae
Thyasiridae
Thyasiridae
Thyasiridae
Thyasiridae
Thyasiridae
Thyasiridae
Thyasiridae
Corbulidae
Myidae
Myidae
Myidae
Myidae
Mytilidae
Mytilidae
Mytilidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Mytilus
Mytilus edulis
Nuculana minuta
Yoldiella lucida
Ennucula tenuis
Nucula
Nucula hanleyi
Nucula nitidosa
Nucula nucleus
Nucula sulcata
Nucula turgida
Nuculidae
Anomiidae
Monia patelliformis
Aequipecten
opercularis
Palliolum striatum
Palliolum tigerinum
Pectinidae
Arctica islandica
Acanthocardia
Acanthocardia
aculeata
Acanthocardia
echinata
Acanthocardia
paucicostata
Cardiidae
Cerastoderma edule
Laevicardium crassum
Parvicardium exiguum
Parvicardium
minimum
Parvicardium scabrum
Donax vittatus
Kelliella miliaris
Vesicomya abyssicola
Hemilepton nitidum
Lepton squamosum
Lutraria lutraria
Mactra
Mactra stultorum
Spisula
Spisula elliptica
Spisula solida
Spisula subtruncata
Devonia perrieri
Epilepton clarkiae
Kurtiella bidentata
Montacuta
phascolionis
Montacuta substriata
Tellimya ferruginosa
Tellimya tenella
Gari
Gari costulata
138228
140480
140577
142002
140584
138262
140588
140589
140590
140592
152274
204
214
153027
140687
1
1
2
2
2
2
2
2
2
2
2
2
1
2
2
1
1
3
3
3
3
3
3
3
3
3
3
1
2
3
E
E
S
S
S
S
S
S
S
S
S
S
E
S
S
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Mytiloida
Mytiloida
Nuculanoida
Nuculanoida
Nuculida
Nuculida
Nuculida
Nuculida
Nuculida
Nuculida
Nuculida
Nuculida
Pectinoida
Pectinoida
Pectinoida
Mytilidae
Mytilidae
Nuculanidae
Yoldiidae
Nuculidae
Nuculidae
Nuculidae
Nuculidae
Nuculidae
Nuculidae
Nuculidae
Nuculidae
Anomiidae
Anomiidae
Pectinidae
140709
140710
213
138802
137732
138990
2
2
2
2
2
2
3
3
3
2
2
2
S
S
S
S
S
S
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Pectinoida
Pectinoida
Pectinoida
Veneroida
Veneroida
Veneroida
Pectinidae
Pectinidae
Pectinidae
Arcticidae
Cardiidae
Cardiidae
138992
2
2
S
Mollusca
Bivalvia
Veneroida
Cardiidae
138993
2
2
S
Mollusca
Bivalvia
Veneroida
Cardiidae
229
138998
139004
139008
139010
2
2
2
2
2
2
2
2
2
2
S
S
S
S
S
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Cardiidae
Cardiidae
Cardiidae
Cardiidae
Cardiidae
139012
139604
152396
464333
246148
140218
140295
138158
140299
138159
140300
140301
140302
140365
140366
345281
140374
2
2
4
4
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
S
S
B
B
S
S
S
S
S
S
S
S
S
S
S
S
S
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Cardiidae
Donacidae
Kelliellidae
Kelliellidae
Lasaeidae
Lasaeidae
Mactridae
Mactridae
Mactridae
Mactridae
Mactridae
Mactridae
Mactridae
Montacutidae
Montacutidae
Montacutidae
Montacutidae
140377
146952
152397
138388
140868
2
2
2
2
2
2
2
2
2
2
S
S
S
S
S
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Montacutidae
Montacutidae
Montacutidae
Psammobiidae
Psammobiidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
3979
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Gari depressa
Gari fervensis
Gari tellinella
Abra
Abra alba
Abra nitida
Abra prismatica
Azorinus chamasolen
Solecurtus scopula
Angulus
Angulus fabula
Angulus incarnatus
Angulus pygmaeus
Angulus tenuis
Arcopagia crassa
Macoma balthica
Moerella donacina
Peronaea planata
Tellina distorta
Tellina nitida
Turtonia minuta
Chamelea gallina
Chamelea striatula
Clausinella fasciata
Dosinia
Dosinia exoleta
Dosinia lupinus
Gouldia minima
Mysia undata
Petricolaria
pholadiformis
Polititapes virgineus
Timoclea ovata
Venerupis corrugata
Venerupis corrugata
Venus casina
Chaetoderma
nitidulum
Caudofoveata
Odostomia scalaris
Gibbula cineraria
Bittium reticulatum
140869
140870
140873
138474
141433
141435
141436
141541
141543
146491
152829
594114
152828
146492
141577
141579
147021
605934
141585
141589
141875
141907
141908
141909
138636
141911
141912
141916
140728
156961
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Psammobiidae
Psammobiidae
Psammobiidae
Semelidae
Semelidae
Semelidae
Semelidae
Solecurtidae
Solecurtidae
Tellinidae
Tellinidae
Tellinidae
Tellinidae
Tellinidae
Tellinidae
Tellinidae
Tellinidae
Tellinidae
Tellinidae
Tellinidae
Turtoniidae
Veneridae
Veneridae
Veneridae
Veneridae
Veneridae
Veneridae
Veneridae
Veneridae
Veneridae
507877
141929
181364
181364
141934
139106
2
2
2
2
2
2
2
2
2
2
2
2
S
S
S
S
S
S
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Bivalvia
Caudofoveata
Veneroida
Veneroida
Veneroida
Veneroida
Veneroida
Chaetodermatida
Veneridae
Veneridae
Veneridae
Veneridae
Veneridae
Chaetodermatidae
151365
141014
141782
139054
2
4
1
2
2
3
3
4
S
B
E
S
Mollusca
Mollusca
Mollusca
Mollusca
Caudofoveata
Gastropoda
Gastropoda
Gastropoda
Epitonium turtonis
139738
2
2
S
Mollusca
Gastropoda
Eulima bilineata
566940
4
3
B
Mollusca
Gastropoda
Eulima glabra
139805
4
3
B
Mollusca
Gastropoda
135
4
3
B
Mollusca
Gastropoda
Vitreolina philippi
139903
4
3
B
Mollusca
Gastropoda
Turritella communis
141872
4
3
B
Mollusca
Gastropoda
Hedylopsis spiculifera
140085
4
3
B
Mollusca
Gastropoda
Eulimidae
3980
[unassigned]
Caenogastropoda
[unassigned]
Caenogastropoda
[unassigned]
Caenogastropoda
[unassigned]
Caenogastropoda
[unassigned]
Caenogastropoda
[unassigned]
Caenogastropoda
[unassigned]
Caenogastropoda
Acochlidiacea
Pyramidellidae
Trochidae
Cerithiidae
Epitoniidae
Eulimidae
Eulimidae
Eulimidae
Eulimidae
Turritellidae
Hedylopsidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Akera bullata
Cylichna alba
Cylichna cylindracea
Philine
Philine aperta
Philine scabra
Retusa
Retusa truncatula
Roxania utriculus
Scaphander lignarius
Aporrhais pespelecani
Caecum glabrum
Crepidula fornicata
Galeodea
echinophora
Hydrobiidae
Ceratia
Ceratia proxima
Hyala vitrea
Euspira pulchella
Lunatia catena
Lunatia montagui
Naticidae
Alvania
Alvania cancellata
Alvania cimicoides
Alvania subsoluta
Alvania testae
Rissoa
Rissoa parva
Lamellaria latens
Buccinum undatum
Neptunea antiqua
Mangelia attenuata
Raphitoma linearis
Nassarius incrassatus
Nassarius reticulatus
Onchidoris
Cuthona
Acteon tornatilis
Turbonilla
Turbonilla acuta
Turbonilla crenata
Rissoella
Rissoella diaphana
Rissoella opalina
Lepidochitona
(Lepidochitona)
cinerea
Leptochiton asellus
Antalis entalis
Antalis vulgaris
Entalina tetragona
Nematoda
Cerebratulus
Lineidae
138734
139474
139476
138339
140744
140761
138432
141138
139486
139488
138760
138952
138963
139023
2
2
2
2
2
2
2
2
2
2
2
1
1
1
3
3
3
3
3
3
3
3
3
3
3
2
1
4
S
S
S
S
S
S
S
S
S
S
S
E
E
E
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Anaspidea
Cephalaspidea
Cephalaspidea
Cephalaspidea
Cephalaspidea
Cephalaspidea
Cephalaspidea
Cephalaspidea
Cephalaspidea
Cephalaspidea
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Akeridae
Cylichnidae
Cylichnidae
Philinidae
Philinidae
Philinidae
Retusidae
Retusidae
Scaphandridae
Scaphandridae
Aporrhaidae
Caecidae
Calyptraeidae
Cassidae
120
138082
140128
140129
140539
150590
150639
145
138439
141165
141168
141247
141251
138456
141365
140172
138878
138920
139265
139371
140503
140513
138288
138543
138691
138421
141052
141057
144173
141147
141149
140143
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1
1
2
2
2
2
2
2
2
1
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
4
4
4
4
3
3
1
2
3
2
2
2
3
3
3
2
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
S
E
S
S
S
S
S
S
S
E
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Mollusca
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Gastropoda
Polyplacophora
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Littorinimorpha
Neogastropoda
Neogastropoda
Neogastropoda
Neogastropoda
Neogastropoda
Neogastropoda
Nudibranchia
Nudibranchia
Chitonida
Hydrobiidae
Iravadiidae
Iravadiidae
Iravadiidae
Naticidae
Naticidae
Naticidae
Naticidae
Rissoidae
Rissoidae
Rissoidae
Rissoidae
Rissoidae
Rissoidae
Rissoidae
Velutinidae
Buccinidae
Buccinidae
Conidae
Conidae
Nassariidae
Nassariidae
Onchidorididae
Tergipedidae
Acteonidae
Pyramidellidae
Pyramidellidae
Pyramidellidae
Rissoellidae
Rissoellidae
Rissoellidae
Tonicellidae
140199
150534
196380
139691
799
122348
122314
1
3
3
2
2
4
4
2
2
2
2
2
3
3
E
UC
UC/DC
S
S
B
B
Mollusca
Mollusca
Mollusca
Mollusca
Nematoda
Nemertea
Nemertea
Polyplacophora
Scaphopoda
Scaphopoda
Scaphopoda
Lepidopleurida
Dentaliida
Dentaliida
Gadilida
Leptochitonidae
Dentaliidae
Dentaliidae
Entalinidae
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Anopla
Anopla
Cerebratulidae
Cerebratulidae
3981
s et al.
A.M. Queiro
Marine Invertebrate Bioturbation Classification
Table 1. Continued.
Scientific Name
Aphia ID
Ri
Mi
Fti
Phylum
Class
Order
Family
Nemertea
Tubularia
Phoronida
Phoronis
Phoronis muelleri
Phoronis pallida
Turbellaria
Platyhelminthes
Pachymatisma
johnstonia
Golfingia
Golfingia
(Golfingia)
elongata
Golfingia (Golfingia)
iniqua
Golfingia (Golfingia)
vulgaris vulgaris
Nephasoma
(Nephasoma)
minutum
Thysanocardia
procera
Onchnesoma
Onchnesoma
magnibathum
Onchnesoma
steenstrupii
steenstrupii
Phascolion
(Phascolion)
strombus
strombus
Sipunculidea
Sipuncula
Aspidosiphon
(Aspidosiphon)
muelleri muelleri
Ericthonius
Goneplax
rhomboides
Isaeidae
Melitidae
152391
600947
1789
128545
128549
128551
794
793
134057
4
1
2
2
2
2
2
2
1
3
1
1
1
1
1
2
2
1
B
E
S
S
S
S
S
S
E
Nemertea
Ochrophyta
Phoronida
Phoronida
Phoronida
Phoronida
Platyhelminthes
Platyhelminthes
Porifera
Demospongiae
Astrophorida
Geodiidae
136021
175026
4
4
3
3
B
B
Sipuncula
Sipuncula
Sipunculidea
Sipunculidea
Golfingiida
Golfingiida
Golfingiidae
Golfingiidae
136042
4
3
B
Sipuncula
Sipunculidea
Golfingiida
Golfingiidae
410724
4
3
B
Sipuncula
Sipunculidea
Golfingiida
Golfingiidae
136060
4
3
B
Sipuncula
Sipunculidea
Golfingiida
Golfingiidae
136063
4
3
B
Sipuncula
Sipunculidea
Golfingiida
Golfingiidae
136024
136064
4
4
3
3
B
B
Sipuncula
Sipuncula
Sipunculidea
Sipunculidea
Golfingiida
Golfingiida
Phascolionidae
Phascolionidae
410742
2
2
S
Sipuncula
Sipunculidea
Golfingiida
Phascolionidae
410749
2
2
S
Sipuncula
Sipunculidea
Golfingiida
Phascolionidae
1296
1268
410717
4
4
4
3
3
3
B
B
B
Sipuncula
Sipunculida
Sipunculidea
101567
107292
2
4
1
4
S
B
101388
101397
2
2
3
2
S
S
Bacillariophyceae
Phoronidae
Turbellaria
Mi scores: 1 for organisms that live in fixed tubes; 2 indicates limited movement; 3 indicates slow, free movement through the sediment matrix;
4 indicates free movement, that is, via burrow system. Ri scores: 1 for epifauna; 2 for surficial modifiers; 3 for upward and downward conveyors;
4 for biodiffusors; and 5 for regenerators. Reworking types (Fti): “S” for surficial modifiers; “B” for biodiffusors; “UC” and “DC” for upward and
downward conveyors; “R” for regenerators; and “E” for epifauna.
Thus far, variation in burrowing behavior across life
stages has been poorly documented in the literature.
However, in some species, juveniles and adults are
known to exhibit different burial behavior, which can
also be modified during reproductive stages (Aguzzi and
Sarda 2007; Schwalb and Pusch 2009). If such changes in
3982
behavior are known to occur, different trait scores
should be attributed as appropriate to different genders
or life stages.
3 Where no species level information exists, taxa are
assumed to have a similar bioturbation mode to others,
which are closely related taxonomically. The paucity of
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
s et al.
A.M. Queiro
information on many bioturbators (Teal et al. 2008)
necessitates matching some species with the closest
possible species or taxonomic group (e.g., genus or
family). Regardless of whether taxonomic relatedness is
a good indicator of the ecological characteristics of a
species (Bevilacqua et al. 2012), the table presented
here does not account for changes in taxonomic classification over time. Such changes in taxonomy could
alter an organism’s taxonomic relatedness to other taxa
and therefore its assumed bioturbation classification.
For example, Alitta virens (Sars 1835) and Hediste
diversicolor (M€
uller 1776) may have been classified as
being functionally similar, as both these species were
formerly classified under the genus Nereis (Linnaeus
1758). Recent changes to their taxonomic classification
now better reflect fundamental differences in bioturbation modes (i.e., gallery diffusor and conveyor, respectively, Francßois et al. 2002). Nevertheless, in the
absence of specific information, we consider it likely
that genetically and physically similar taxa are likely to
be functionally similar. As BPc is a biomass-weighted
sum of traits from many species, we consider it unlikely that small changes to trait assignments of individual species would greatly influence large-scale
assessments. The current structure of the table reflects
the taxonomic classification of the species at the time
of analysis and will need updating as taxonomic inventories are refined.
Accepting the limitations imposed by the assumptions
underpinning BPc, the classification list we have assembled will facilitate the calculation of BPc across most
temperate coast and shelf benthic environments in the
North Atlantic. This aspects makes a strong case for a
wider implementation of BPc as a standardized indicator: the ability to build on existing data (abundance
and biomass) to fill gaps about bioturbation patterns
where direct assessments are not, or cannot, been routinely carried out. Further efforts will need to take
place for other regions of the world or particular circumstances. There is also tremendous potential for
application to historical datasets, to estimate bioturbation rates in the past, providing insight into how this
process has helped to shape sedimentary ecosystems
over time (Solan et al. 2008). Finally, the Marine Strategy Framework Directive (2008/56/EC) now requires an
integrated understanding and management of regional
scale patterns of marine ecosystem functioning, based
on the use of inexpensive, rapid indicators. Few functional indicators currently exist for European waters,
and initial tests of the suitability and applicability of
BPc suggest that it holds promise as a tool for informing management and policy (Van Hoey et al. 2010; Birchenough et al. 2012).
ª 2013 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Marine Invertebrate Bioturbation Classification
Acknowledgments
This publication was initiated and facilitated by the Study
Group Climate Related Benthic Processes in the North
Sea (SGCBNS), which is an expert group of the International Council for the Exploration of the Sea (ICES).
Funding was supported by the Western Channel Observatory, part of the UK Natural Environmental Research
Council’s National Capability (Queir
os, Somerfield and
Widdicombe), the Department of the Environment, Food
and Rural Affairs under Science-Level Agreement SLA31
(Birchenough), the Conseil Regional d’Aquitaine and the
University of Bordeaux 1 (Romero-Ramirez), and the
Flemish Fund for Scientific Research (Van Colen). Caroline Louise McNeill and Sarah Dashfield are thanked for
their contributions towards one of the datasets analyzed.
Celine Labrune and Jean-Michel Amouroux are thanked
for contributions towards another.
Conflict of Interest
None declared.
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