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African Journal of Marine Science
ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/tams20
Zoogeography of marine Bryozoa around South
Africa
MK Boonzaaier-Davids , WK Florence & MJ Gibbons
To cite this article: MK Boonzaaier-Davids , WK Florence & MJ Gibbons (2020) Zoogeography of
marine Bryozoa around South Africa, African Journal of Marine Science, 42:2, 185-198
To link to this article: https://doi.org/10.2989/1814232X.2020.1765870
Published online: 11 Aug 2020.
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African Journal of Marine Science 2020, 42(2): 185–198
Printed in South Africa — All rights reserved
Copyright © NISC (Pty) Ltd
AFRICAN JOURNAL OF
MARINE SCIENCE
ISSN 1814-232X EISSN 1814-2338
https://doi.org/10.2989/1814232X.2020.1765870
African Journal of Marine Science is co-published by NISC (Pty) Ltd and Informa UK Limited (trading as Taylor & Francis Group)
Zoogeography of marine Bryozoa around South Africa
MK Boonzaaier-Davids1,2* , WK Florence2 and MJ Gibbons1
1 Department of Biodiversity and Conservation Biology, University of the Western Cape, Cape Town, South Africa
2 Research and Exhibitions Department, Iziko South African Museum, Cape Town, South Africa
* Corresponding author, e-mail: melissakaydavids@gmail.com
The zoogeography of marine Bryozoa around South Africa was investigated using published distribution
records, museum catalogues, and an examination of previously unworked bryozoan material in (mostly) museum
collections. Although a total of 276 valid species are recognised, it was not possible to unambiguously assess
geographic patterns of diversity. At all depth zones examined (shore and inner-shelf, 0–30 m; mid- and outer-shelf,
31–350 m; bathyal, >500 m), there was a clear geographic structure to communities that mirrored established
regional patterns of biogeography. Too few samples were collected from the shelf edge (351–500 m) and they
were consequently excluded from zoogeographic analysis. Communities on the shore and inner-shelf and on the
mid- and outer-shelf were more similar to each other than they were to bathyal communities, and the pronounced
structure in bathyal communities suggests heterogeneity in the deep sea around South Africa.
Keywords: Agulhas Current, Benguela Current, biogeography, cyphonautes, distribution, marine ecoregions, Namaqua marine bioregion,
species database
In the latest National Biodiversity Assessment (Skowno
et al. 2019), four broad biogeographic provinces have
been identified for nearshore taxa around South Africa,
and these are treated as the Delagoa (tropical), Natal
(subtropical), Agulhas (warm-temperate) and Southern
Benguela (cold-temperate) ecoregions. The Natal ecoregion
is split into two subregions, the ‘KwaZulu-Natal Bight’ in the
north and ‘southern KZN’ in the south, while the Southern
Benguela ecoregion is divided into a northern ‘Namaqua’
subregion and a southern ‘Cape’ subregion. Farther
offshore, two ecoregions are recognised: the Southwest
Indian and the Southeast Atlantic. The delineation of these
biogeographic provinces generally corresponds to the
distribution of major water masses and the interactions
between them (e.g. Maree et al. 2000; Spalding et al. 2007;
Gibbons et al. 2010a), although precise boundaries are the
subject of much discussion (Lombard et al. 2004; von der
Heyden 2009; Teske et al. 2011).
Owing to the strong association between water mass
and distribution (e.g. Promińska et al. 2018), variations in
patterns of marine biogeography among different taxa
are likely to be observed at different depths (Cleary et al.
2005; Hilário et al. 2015). Surface waters are subject to
wind stress, and regional water masses display regionally
discrete patterns of circulation linked to that wind stress
and its interactions with topography and bathymetry. In
contrast, the deeper ocean is less dynamic and more
homogeneous in nature, and water masses may be more
extensively distributed (see Zeppilli et al. 2016). That said,
our understanding of the role of depth in determining the
biogeography of marine organisms around South Africa
and southern Africa is poor. This is particularly true in the
deep sea where surveys are difficult and the expense of
collecting new data is relatively high (Tanner et al. 2018).
Bryozoans are aquatic, sessile, clonal animals that occur
as colonies attached to hard substrates, such as rocks, shells
and other bryozoans, and sometimes on soft substrates such
as algae (Bock 1982; Hayward and Ryland 1999). The phylum
is monophyletic and can be divided into three monophyletic
classes: Gymnolaemata, Stenolaemata and Phylactolaemata
(Fuchs et al. 2009; Taylor and Waeschenbach 2015).
Members of Phylactolaemata occur exclusively in fresh water,
while members of Stenolaemata (order Cyclostomatida) and
Gymnolaemata (orders Ctenostomatida and Cheilostomatida)
inhabit marine and/or estuarine environments (Hayward and
Ryland 1999; Bock and Gordon 2013). Most of our knowledge
about bryozoans around South Africa is taxonomic in nature,
though a few works have been published on biology and
ecology (e.g. Hayward and Cook 1979; Gray et al. 2005;
Florence et al. 2007).
In the last two decades, published taxonomic works on
South African bryozoans dealt with shallow-water (<30 m)
species collected along the west coast (Florence et al.
2007), some deep-water species (Florence 2016) and a
new species of Taylorius Gordon, 2014 (Oliver and Florence
2016). Investigations prior to that focused mostly on taxa in
intertidal and shallow subtidal waters, as well as deep-water
species from the east coast (Boonzaaier et al. 2014). The
first species-list of South African bryozoans was constructed
nearly a century ago by O’Donoghue (1924), but taxonomic
revisions of problematic genera or species, descriptions of
new species, and resolved synonymies have taken place
since then. An estimated 270 valid species are known from
South Africa (Florence et al. 2007), but our knowledge of the
Introduction
Published online 11 Aug 2020
Boonzaaier-Davids, Florence and Gibbons
186
bryozoan fauna from deep waters on the west and south
coasts remains poor.
Although most bryozoans can reproduce both sexually
and asexually, few species have planktotrophic larvae:
most release lecithrotrophic larvae that are resident in the
plankton for a short period of time (Ryland 1965; Taylor
1988). It might therefore be expected that the biogeography
of bryozoans around South Africa would display clear
provincial structure, as known for hydrozoans (Gibbons
et al. 2010a). Bryozoans are conspicuous components of
fouling communities, however, and are frequently recovered
from flotsam, which may serve to disperse populations in
the same way that planktotrophic larvae are distributed by
ocean currents (Cornelius 1992; Winston 2012).
The aim of this study was to examine the zoogeography
of bryozoans around South Africa, and to test congruity
in pattern with the depth zones described by Sink et al.
(2019). Published and unpublished data were used, as well
as validated museum specimens. The taxonomy employed
follows recent advances in the field.
Materials and methods
Study area
The South African coastline stretches from the Namibia
border (29°42ʹ S, 17°59ʹ E) to the Mozambique border
(26°51ʹ S, 32°49ʹ E) and is roughly divided into four regions,
namely the cool-temperate west coast, warm-temperate
south coast, subtropical southeast coast, and tropical east
coast. The ecoregion classification shown in Figure 1a
follows Skowno et al. (2019) and Sink et al. (2019).
Creating a species database
All published and unpublished information on the distribution
of extant marine bryozoans in South Africa was compiled
into a database. Three types of records were collated:
(i) observational records (no specimen, but species observed
and recorded); (ii) specimen records (catalogued specimens
in museums or institutions); and (iii) literature records (for
example, media, photographs and published research).
Specimen records were obtained from digitised catalogues
of deposited specimens within collections. Approximately
1 225 catalogued specimens housed at the Iziko South
African Museum (SAM) were extracted using Specify 6.5.02
software. Of these, 1 028 specimen records were used here,
together with 1 403 records from the Natural History Museum
(NHM) in London.
The works of Busk (1852, 1854, 1875, 1884, 1886),
O’Donoghue (1924, 1957), O’Donoghue and de Watteville (1935,
1937, 1944), Hayward and Cook (1979, 1983), and Florence
et al. (2007) were the main sources of literature records.
Lesser sources included The Bryozoa Home Page (http://
www.bryozoa.net) and the World Register of Marine Species
(WoRMS, http://www.marinespecies.org). Additionally, over 1 000
uncatalogued specimens from the collections of the SAM were
identified to species-level, where possible, and associated locality
information (geographic coordinates and depth) was compiled.
Data-cleaning
A ‘usable’ record contained a valid species name, locality
name and/or locality coordinates (latitude/longitude),
and depth. Some evident morpho-species (species
morphologically different from related species) were
identified at the genus level, where possible, and also
included in the dataset. Data were cleaned in a three-step
process: (i) updating of resolved synonymies; (ii) inferring
omitted locality data; and (iii) removing dubious or
duplicated records. Where possible, synonymies were
resolved using the WoRMS Taxon Match Tool (http://
marinespecies.org/aphia.php?p=match) and the literature.
Most records from the 19th and early 20th centuries did
not have detailed information on locality, and the use of
nearest place names was the norm. In these cases, the
coordinates for localities were obtained using GEOLocate
(https://geo-locate.org). Where specimens were collected
by hand, either from the intertidal zone or scraped from, for
example, a plate in harbours, depth was inferred as <10 m.
Bryozoans collected from the hulls of ships, in a random
collection event, or from beach casts were excluded from the
dataset. Dubious or doubtful observational records, including
locality records on land, and records not verified by an
expert, were also omitted from the dataset.
After data-cleaning, records from over 600 samples falling
within South Africa’s exclusive economic zone (EEZ) were used
to analyse distribution patterns. A sample refers to a collection
from a single locality or station made at a single point in time,
and may contain more than one species. A sample reflects
dedicated sampling effort along the coast, as opposed to a
record, which is regarded here as an ad hoc collection. Of the
approximately 1 700 ‘usable’ records, more than 800 records
were from both the literature and current collections (housed
at the SAM), while the database from the SAM yielded an
additional 74 unpublished records and that from the NHM
an additional three records, as the majority of records from
this institution were already published.
Geographic assignment of data
Instead of a full-degree square or quarter-degree-grid cell
(QDGC) (1° latitude × 1° longitude) that would misrepresent
the sparsely distributed data points, a level-2 QDGC (0.25°
latitude × 0.25° longitude or 16 squares per degree square)
was created using the vector grid tool in QGIS 2.2.0 (see
Larsen et al. 2009). This resulted in the coastline being divided
into 0.25 (i.e. level-2) QDGCs, each with its output number
(Figure 1b). This layer was projected into ArcGIS 10.2 and
was combined with the data-point layer created from the
species database.
The available literature suggests that bryozoan diversity
changes with depth (e.g. Rouse et al. 2014; Denisenko et al.
2016). In this study, existing bryozoan samples in South Africa
had been taken at varying depth intervals, and, in many cases,
inadequate sampling effort across depth intervals meant
that the available depth data were not adequate to deduce
systematic depth-related gradients in species distribution.
Consequently, the data were categorised into depth
zones, following Sink et al. (2019), as shore and inner-shelf
(0–30 m), mid- and outer-shelf (31–350 m), and bathyal
(>500 m), in addition to being assigned a marine ecoregion
(see Figure 1a). The few samples that were collected from
the shelf edge (351–500 m) were all collected along the west
coast; they make no contribution to our understanding of
zoogeography and hence were excluded from further analysis.
African Journal of Marine Science 2020, 42(2): 185–198
187
Figure 1: Map of the Southern African subregion showing (a) marine ecoregions (from Sink et al. [2019]), with some of the localities
mentioned in the main text, and (b) level-2 quarter-degree-grid cells (QDGCs) and their in-text numbering. Two main transition zones (i.e.
zones lying between two regions) are False Bay in the Western Cape Province and East London in the Eastern Cape Province (Skowno
et al. 2019)
AFRICA
South
Africa
SOUTH
AFRICA
Kosi Bay
St Lucia
Southern Benguela
Southwest Indian Deep Ocean
Southeast Atlantic Deep Ocean
Natal
Agulhas
Delagoa
Durban
Scotburgh
Hluleka
East London
Port Elizabeth
False Bay
Agulhas
Cape Peninsula
Cape Point
Table Bay
Langebaan
Saldanha Bay
Lambert’s Bay
SOUTH AFRICA
ATLANTIC
OCEAN
INDIAN OCEAN
(a)
(b)
30° S
30° E20° E
20° E
Boonzaaier-Davids, Florence and Gibbons
188
Data from the joint layer (with unique QDGCs and
associated data points) were extracted and, following
Samaai (2006), used to create a data matrix of species
presence/absence. Latitudinal gaps between records, per
depth zone, were filled on the assumption that the absence
of records reflected a lack of sampling (interpolated data).
Species that were present in only a single QDGC, following
interpolation, were excluded from further analysis, unless
they occupied an isolated position at the extremes of the
grid, latitudinally or bathymetrically.
Statistical analyses
To test for differences in the species composition of QDGCs
by ecoregion and depth zone, a similarity matrix was first
generated between each using the Bray–Curtis index.
This was visualised using cluster analysis (group average)
and nonmetric multidimensional scaling (nmMDS). The
cluster analysis was run with SIMPROF, which determines
the subclusters in the hierarchical group-average cluster
analysis that can be interpreted as distinguishable. To test
whether there were differences between the composition
of the a priori geographic and bathymetric regions, data
were tested using a two-factor PERMANOVA (Anderson
et al. 2008). The factors Depth (zone) and Ecoregion were
fixed; the permutation of residuals was conducted using
a reduced model, and Type III sums of squares were
calculated. All analyses were computed using PRIMER 7.
Results
The dataset included 276 valid species of Bryozoa, belonging
to 148 genera, 74 families, three orders (Cyclostomatida,
Ctenostomatida and Cheilostomatida) and two classes
(Stenolaemata and Gymnolaemata). The three most-species-
rich families were Adeonidae (31 species), Phidoloporidae
(22 species) and Bugulidae (14 species). The most diverse
genera were Adeonella (26 species), Rhynchozoon
(9 species) and Reteporella (8 species). The full species
list used for this study is provided in the Appendix, though
a detailed, annotated species list is being prepared for
separate publication.
Table 1 shows the distribution of effort and observed
richness across the different a priori ecoregions and depth
zones. There is clearly an incomplete and inconsistent
coverage of the greater region, with certain areas (e.g.
Agulhas mid-shelf) receiving much attention whereas
some other areas have received little to no attention (e.g.
Agulhas bathyal, Natal shallow, Natal bathyal and Delagoa)
(Table 1). This suggests that efforts to definitively map
patterns of diversity across the greater region would, at
present, be premature, though there are clear relationships
between effort in each depth zone and the number of
species collected, across regions (Table 1).
Results of the PERMANOVA reveal highly significant
differences in the structure of bryozoan communities by
a priori ecoregion and depth zone, and by the interaction
between them (Table 2). This is partly visualised by the
nmMDS plot, which shows a pronounced difference in
communities between bathyal QDGCs and those from
shallower depths (Figure 2); and, in general, communities
from the mid- and outer-shelf are distinct from those on the
shore and over the inner-shelf.
Analysing the data within each depth zone separately,
a clear biogeographic pattern was observed across all
Shore and
inner-shelf (<30 m)
Mid- and outer-shelf
(31–350 m)
Shelf edge
(351–500 m)
Bathyal
(>500 m) Total
Ecoregion Subregion Spp. Samples Spp. Samples Spp. Samples Spp. Samples Spp. Samples
Southern
Benguela
Namaqua 8 5 18 5 8 4 0 0 28 14
Cape 91 83 60 31 14 8 3 3 135 125
Agulhas 87 61 192 123 0 0 0 0 222 184
Natal sKZN 0 0 2 2 0 0 0 0 2 2
KZNB 16 3 46 8 3 1 0 0 62 12
Delagoa 0 0 0 0 0 0 0 0 0 0
Southeast Atlantic 0 0 0 0 0 0 6 6 9 6
Southwest Indian 0 0 0 0 33 2 27 27 81 29
Table 1: Changes in the numbers of bryozoan species and samples collected across each a priori marine ecoregion and depth zone around
South Africa. See Figure 1a for delimitation of the marine ecoregions. Note: Samples collected from the shelf edge or from the Delagoa or
the southern part of the Natal marine ecoregions were not analysed owing to poor coverage. KZNB = KwaZulu-Natal Bight; sKZN = southern
KwaZulu-Natal
Source df SS MS Pseudo-F p(perm) p(MC) Estimate
Ecoregion 4 89274 22319 23.998 0.001 0.001 1 510.6
Depth 1 32712 32712 35.173 0.001 0.001 1 291.1
Ecoregion × Depth 3 35919 11973 12.874 0.001 0.001 1 468.6
Residuals 104 672 930.04 930.04
Total 113 36075
Table 2: PERMANOVA results showing the effect of a priori ecoregion, depth zone, and the interaction between ecoregion
and depth zone, on the structure of bryozoan ‘communities’ around South Africa. Data in the right-hand column show
estimates of the components of variation. df = degrees of freedom; SS = sum of squares; MS = mean sum of squares
African Journal of Marine Science 2020, 42(2): 185–198
189
depth zones (Figures 3–5), with an orderly arrangement of
QDGCs that matches geographic position. In the case of
the shore and inner-shelf fauna (Figure 3), distinct clusters
are evident for the Namaqua, Cape, Agulhas and Natal
ecoregions, with boundaries in the vicinity of Lambert’s
Bay, Table Bay, and between East London and Durban,
respectively. The results of the pairwise tests computed in
PERMANOVA reveal significant differences between all
ecoregions in this depth zone (data not shown).
Over the mid- and outer-shelf depth zone (Figure 4)
there was a clear subdivision between the Natal ecoregion
and the balance of QDGCs (~8% similarity), with those of
the Agulhas ecoregion clustering more closely with the
Southern Benguela ecoregion (~27%). As noted in the
analyses of the shore and inner-shelf fauna, there was some
mismatch in the clustering of the a priori ecoregions and
the QDGCs, with overlap in adjacent QDGCs. The breaks
between the Namaqua and Cape subregions of the Southern
Benguela ecoregion occurred in the vicinity of Saldanha
Bay, and between the Cape subregion and the Agulhas
ecoregion in False Bay. The break between the Agulhas
and Natal ecoregions occurred in the vicinity of Scottburgh
in southern KZN. The results of the pairwise tests computed
in PERMANOVA reveal significant differences between all
ecoregions in this depth zone (data not shown).
Figure 2: Nonmetric multidimensional scaling plot showing the
similarity in the bryozoan composition (presence, absence)
between level-2 quarter-degree-grid cells (QDGCs), by a priori
marine ecoregion and depth zone around South Africa. Ecoregions:
Agulhas = Agulhas ecoregion; KZNB = KwaZulu-Natal Bight
subregion of the Natal ecoregion; Namaqua and Cape = subregions
of the Southern Benguela ecoregion; SEAtl = Southeast Atlantic
ecoregion; SWInd = Southwest Indian ecoregion. Depth zones:
Inner = shore and inner-shelf; Mid/Outer = mid- and outer-shelf;
Bathy = bathyal
Figure 3: Ordination plot showing the similarity in the bryozoan composition (presence, absence) between QDGCs by a priori marine
ecoregion across the shore and inner-shelf depth zone (<30 m) around South Africa. Clustering by group-average linkage. Dotted lines
indicate those samples considered as indistinguishable using SIMPROF. Agulhas = Agulhas ecoregion; KZNB = KwaZulu-Natal Bight
subregion of the Natal ecoregion; Namaqua and Cape = subregions of the Southern Benguela ecoregion
Boonzaaier-Davids, Florence and Gibbons
190
Interestingly, excluding the latitudinal outliers in the
bathyal region (Figure 5), the split between the fauna of
the Southeast Atlantic and Southwest Indian ecoregions
occurred in the vicinity of East London (QDGCs #10 and
#11), at approximately 11% similarity. Once again, the
results of the pairwise tests computed in PERMANOVA
reveal significant differences between all ecoregions in this
depth zone (data not shown).
Discussion
The gaps noted here in terms of geographic and bathymetric
coverage of the Bryozoa samples are similar, in part, to
those of other studies (Sink et al. 2005; Samaai 2006;
Samaai et al. 2010). There is a paucity of samples from deep
water off the west, south and southeast coasts of South
Africa, but not from off the east coast, for which area major
taxonomic works were published in the reports of Hayward
and Cook (1979, 1983), documenting deep-water collections
from the SAM’s Meiring Naudé cruises. The lack of samples
from shallow waters along the east coast is unexplained and
likely attributed to dubious records from the SAM database
that were excluded from the dataset, highlighting the need
for consistency and accuracy in digitising museum records
for biodiversity research. However, given the accessibility of
these habitats, the paucity of samples could be rectified.
Despite the gaps between some QDGCs in the present
analysis, the use of interpolation reduces their impact.
That said, it does artificially increase the similarity between
adjacent QDGCs and it contributes to their smooth
geographical ordering in nmMDS plots. The results of the
present analysis conform in general to the biogeographic
patterns observed by numerous others investigating
shallow-water benthic communities (e.g. Emanuel et al.
1992; Awad et al. 2002; Scott et al. 2012) and taxa such as
algae (Bolton and Stegenga 2002; Anderson et al. 2009),
fish (Turpie et al. 2000), hydrozoans (Millard 1978; Gibbons
et al. 2010a) and echinoderms (Thandar 1989). The results
are also consistent with the results of work on pelagic taxa:
euphausiids (Gibbons 1995) and siphonophores (Gibbons
and Thibault-Botha 2005). Discrepancies between our
data and previously published work relate to the position
of breaks between bioregions (see Table 3), which are
generally trivial and probably a reflection of differences in
methodological resolution rather than being real. This is
particularly true for deep-water environments, where large
spatial and knowledge gaps exist (Sink et al. 2019), and for
some taxa that remain understudied and undersurveyed
Figure 4: Ordination plot showing the similarity in the bryozoan composition (presence, absence) between QDGCs by a priori marine
ecoregion across the mid- and outer-shelf depth zone (31–350 m) around South Africa. Clustering by group-average linkage. Dotted lines
indicate those samples considered as indistinguishable using SIMPROF. Agulhas = Agulhas ecoregion; KZNB = KwaZulu-Natal Bight
subregion of the Natal ecoregion; Namaqua and Cape = subregions of the Southern Benguela ecoregion
African Journal of Marine Science 2020, 42(2): 185–198
191
(e.g. Awad et al. 2002; Gibbons et al. 2010a; Griffiths et al.
2010; Mead et al. 2011; Sink et al. 2019).
Hydrozoans and bryozoans, although phylogenetically
very distinct (Telford et al. 2015), both have (for the most
part) benthic, polymorphic colonies based on clonal
individuals, and as such hydrozoans provide a useful proxy
for bryozoans. Most thecate and anthoathecate hydrozoans
have a restricted larval dispersal (Gibbons et al. 2010b),
and these taxa show a strong biogeographic pattern of
distribution around South Africa (Gibbons et al. 2010a),
a pattern that is linked tightly to the prevailing mesoscale
oceanography. In contrast, hydrozoans that are either
holopelagic (no benthic stage) or meroplanktonic (with an
alternation between a sessile benthic stage and a pelagic
medusa stage) tend to show a weaker biogeographic
pattern than ‘strictly benthic’ taxa around South Africa.
This was argued by Gibbons et al. (2010b) to reflect the
important role of the dispersive medusa stage and its
interaction with macroscale oceanography.
The majority of bryozoan species are brooders and
produce non-feeding, lecithotrophic larvae, which generally
settle within a few hours of release and typically display
limited dispersal (Hayward and Ryland 1999; Clarke
and Lidgard 2000). Most bryozoans therefore lack a
long-distance dispersive larval stage, a cyphonautes larva,
and it might then be expected that they would show patterns
of distribution akin to hydrozoans with a similar life history, as
suggested by our results. However, it might also be expected
that such patterns would be shown only in shallower waters
and that they would fall away in deeper waters, to be
replaced by patterns more similar to those of meroplanktonic
or holopelagic hydrozoans. This difference reflects the
fact that life in shallow water is strongly influenced by local
atmospheric conditions, where the oceanography is dynamic
and subject to local winds interacting with local topography
to create local mesoscale conditions to which species need
to adapt. For example, changes in water temperatures can
affect the settlement of larvae in many marine fishes and
invertebrates (O’Connor et al. 2007).
By contrast, deeper waters tend to move more sluggishly,
over greater distances, and are more uniform in structure
(e.g. Lutjeharms 2006). Some epibenthic invertebrate taxa
show considerable variation in community assemblages,
even within the same depth range, as was found in
deep water along the west coast of South Africa (Lange
and Griffiths 2014). However, a number of deep-water
organisms have wide distributions even though their larvae
lack extensive dispersal powers (Hilário et al. 2015), and
Figure 5: Ordination plot showing the similarity in the bryozoan composition (presence, absence) between QDGCs by a priori marine
ecoregion across the bathyal (>500 m) depth zone around South Africa. Clustering by group-average linkage. Dotted lines indicate those
samples considered as indistinguishable using SIMPROF. SEAtl = Southeast Atlantic ecoregion; SWInd = Southwest Indian ecoregion
Boonzaaier-Davids, Florence and Gibbons
192
hydrodynamic processes presumably play an important role
(e.g. Mariani et al. 2006).
Generally, bryozoan richness declines with depth and is
largely a function of hard substrate availability (Eggleston
1972). This is reflected in the fact that the deep-sea
bed consists of mostly sand and mud, and is therefore
dominated by the conescharelliform and setoselliniform
species generally exclusive to deep-water environments
(Hayward and Cook 1979), such as Characodoma protrusum
(Thornely, 1905), Galeopsis circella (Hayward & Cook,
1979) and Galeopsis bispiramina (Hayward & Cook, 1979).
That deep-water bryozoans do not show such (anticipated)
widespread distributions around South Africa suggests
that, at the depths investigated, there is still significant
heterogeneity in, and mesoscale structure to, the deep-water
environment around the coast.
As mentioned, most bryozoans lack a dispersive
cyphonautes larva, although its presence is thought to
represent the ancestral condition (Taylor 1988; Watts and
Thorpe 2006). Indeed, the evolution of brooded larvae
in cheilostomes is widely credited for the high diversity
of the group, and species with planktotrophic larvae
generally represent less than 5% of regional faunas (Taylor
1988). Off South Africa, the suborder Malacostegina,
which contains all bryozoans with cyphonautes larvae, is
represented by only two species of Electridae (Conopeum,
Harpecia) and three species of Membraniporidae (Biflustra,
Jellyella, Membranipora), and this prevents us from
re-analysing the data by life-history strategy (Gibbons
et al. 2010b). Why species with relatively long-lived and
dispersive larvae should be so uncommon around South
Africa, and so much more uncommon than hydrozoans
with a similar life history (Gibbons et al. 2010b), is
intriguing and worthy of further attention.
Acknowledgements — We are grateful to Iziko Museums of South
Africa for hosting the first author (MKBD) during her doctoral study.
The Department of Biodiversity and Conservation Biology at the
University of the Western Cape (South Africa) is acknowledged for
providing academic and logistical support. MKBD would like to thank
Francuois Müller (Agricultural Research Council) for assistance
with ArcGIS. Funding was provided by the South African National
Research Foundation through the Professional Development
Programme. MJG would like to thank Anne-Gro Salvanes of
the University of Bergen (Norway) and the study programme
PRIMALearn for supporting him while he consolidated the manuscript.
ORCID
Melissa K Boonzaaier-Davids: https://orcid.org/0000-0001-7049-7356
Wayne K Florence: https://orcid.org/0000-0003-0224-2874
Mark J Gibbons: https://orcid.org/0000-0002-8320-8151
References
Anderson MJ, Gorley RN, Clarke KR. 2008. PERMANOVA+ for
PRIMER: guide to software and statistical methods. Plymouth,
UK: PRIMER-E.
Anderson RJ, Bolton JJ, Stegenga H. 2009. Using the
biogeographical distribution and diversity of seaweed species to
Taxa Location of faunal breaks Sources
Bryozoans West coast: Saldanha Bay
South coast: False Bay
East coast: Scottburgh, East London
This study
Marine invertebrates
(several taxa)
Southwest coast: Cape Point
East coast: East London, Durban
Emanuel et al. (1992)
Algae West coast: Langebaan/Saldanha Bay
Southeast coast: Tsitsikamma Coastal National
Park (west of Port Elizabeth)
East coast: Hluleka
Bolton and Stegenga
(2002)
Cnidarians –
Octocorals
Southeast coast: Port Elizabeth
East coast: Durban, St Lucia
Awad et al. (2002)
Awad et al. (2002)
Awad et al. (2002)
Molluscs –
Chitons
Southwest coast: Cape Peninsula
East coast: East London area
Bivalves East coast: Durban area
Prosobranch gastropods East coast: East London area
Opisthobranch gastropods South coast: False Bay
East coast: St Lucia
Polychaetes South coast: False Bay
Echinoderms South coast: False Bay
East coast: Durban
Thandar (1989)
Awad et al. (2002)
Crustaceans –
Amphipods, isopods
South coast: False Bay Awad et al. (2002)
Crabs South coast: False Bay
East coast: Kosi Bay
Vertebrates –
Fishes
Southwest coast: Cape Point
South coast: De Hoop Nature Reserve
(east of Agulhas)
Turpie et al. (2000)
Table 3: Localities of faunal breaks (see Figure 1a) of marine taxa in South Africa, based on this study
and previous studies
African Journal of Marine Science 2020, 42(2): 185–198
193
test the efficacy of marine protected areas in the warm-temperate
Agulhas Marine Province, South Africa. Diversity and Distributions
15: 1017–1027.
Awad A, Griffiths CL, Turpie JK. 2002. Distribution and endemicity
patterns of benthic marine invertebrates in South Africa applied
to the selection of priority conservation areas. Diversity and
Distributions 8: 129–145.
Bock PE. 1982. Bryozoans (Phylum bryozoa or Ectoprocta).
In: Shepherd SA, Thomas IM (eds), Marine invertebrates
of southern Australia: Part I. Adelaide, South Australia:
D.J. Woolman, Government Printer. pp 319–394.
Bock PE, Gordon DP. 2013. Phylum Bryozoa Ehrenberg, 1831.
Zootaxa 3703: 67–74.
Bolton JJ, Stegenga H. 2002. Seaweed species diversity in South
Africa. South African Journal of Marine Science 24: 9–18.
Boonzaaier MK, Florence WK, Spencer Jones ME. 2014. Historical
review of South African bryozoology: a legacy of European
endeavour. In: Wyse Jackson PN, Spencer Jones ME (eds),
Annals of bryozoology 4: aspects of the history of research
on bryozoans. Dublin, Ireland: International Bryozoology
Association. pp 1–34.
Busk G. 1852. Catalogue of marine Polyzoa in the collection of the
British Museum. Part I. Cheilostomata. London: Trustees of the
British Museum.
Busk G. 1854. Catalogue of marine Polyzoa in the collection of the
British Museum. Part II. Cheilostomata. London: Trustees of the
British Museum.
Busk G. 1875. Catalogue of the marine Polyzoa in the collection of
the British Museum. Part III. Cyclostomata. London: Trustees of
the British Museum.
Busk G. 1884. Report on the Polyzoa collected by H.M.S. Challenger
during the years 1873–1876. Part 1. The Cheilostomata. Report
on the Scientific Results of the Voyage of the H.M.S. ‘Challenger’.
Zoology 10: 1–216.
Busk G. 1886. Report on the Polyzoa collected by H.M.S. Challenger
during the years 1873–1876. Part II. The Cyclostomata,
Ctenostomata and Pedicellinea. Report on the Scientific Results of
the Voyage of the H.M.S. ‘Challenger’. Zoology 17: 1–47.
Clarke A, Lidgard S. 2000. Spatial patterns of diversity in the sea:
bryozoan species richness in the North Atlantic. Journal of
Animal Ecology 69: 799–814.
Cleary DFR, Becking LE, de Voogd NJ, Renema W, de Beer M,
van Soest RWM, Hoeksema BW. 2005. Variation in the diversity
and composition of benthic taxa as a function of distance
offshore, depth and exposure in the Spermonde Archipelago,
Indonesia. Estuarine, Coastal and Shelf Science 65: 557–570.
Cornelius PFS. 1992. The Azores hydroid fauna and its origin, with
discussion of rafting and medusa suppression. Arquipélago –
Life and Earth Science 10: 75–99.
Denisenko NV, Hayward PJ, Tendal OS, Sørensen J. 2016.
Diversity and biogeographic patterns of the bryozoan fauna of
the Faroe Islands. Marine Biology Research 12: 360–378.
Eggleston D. 1972. Factors influencing the distribution of
sub-littoral ectoprocts off the south of the Isle of Man (Irish Sea).
Journal of Natural History 6: 247–260.
Emanuel BP, Bustamante RH, Branch GM, Eekhout S, Odendaal FJ.
1992. A zoogeographic and functional approach to the selection of
marine reserves on the west coast of South Africa. In: Payne AIL,
Brink KH, Mann KH, Hilborn R (eds), Benguela trophic functioning.
South African Journal of Marine Science 12: 341–354.
Florence WK. 2016. Some deep-water cheilostome Bryozoa from
the south coast of South Africa. African Natural History 12: 5–11.
Florence WK, Hayward PJ, Gibbons MJ. 2007. Taxonomy of
shallow-water Bryozoa from the west coast of South Africa.
African Natural History 3: 1–58.
Fuchs J, Obst M, Sundberg P. 2009. The first comprehensive
molecular phylogeny of Bryozoa (Ectoprocta) based on
combined analyses of nuclear and mitochondrial genes.
Molecular Phylogenetics and Evolution 52: 225–233.
Gibbons MJ. 1995. Observations on euphausiid assemblages of
the south coast of South Africa. South African Journal of Marine
Science 16: 141–148.
Gibbons MJ, Thibault-Botha D. 2005. Epipelagic siphonophores off
the east coast of South Africa. African Journal of Marine Science
27: 129–139.
Gibbons MJ, Buecher E, Thibault-Botha D, Helm RR. 2010a.
Patterns in marine hydrozoan richness and biogeography around
southern Africa: implications of life cycle strategy. Journal of
Biogeography 37: 606–616.
Gibbons MJ, Janson LA, Ismail A, Samaai T. 2010b. Life cycle
strategy, species richness and distribution in marine Hydrozoa
(Cnidaria: Medusozoa). Journal of Biogeography 37: 441–448.
Gray CA, McQuaid CD, Davies-Coleman MT. 2005. A symbiotic
shell-encrusting bryozoan provides subtidal whelks with chemical
defence against rock lobsters. African Journal of Marine Science
27: 549–556.
Griffiths CL, Robinson TB, Lange L, Mead A. 2010. Marine
biodiversity in South Africa: an evaluation of current states of
knowledge. PLoS ONE 5: e12008.
Hayward PJ, Cook PL. 1979. The South African Museum’s Meiring
Naudé cruises. Part 9, Bryozoa. Annals of the South African
Museum 79: 43–130.
Hayward PJ, Cook PL. 1983. The South African Museum’s Meiring
Naudé cruises. Part 13, Bryozoa II. Annals of the South African
Museum 91: 1–161.
Hayward PJ, Ryland JS. 1999. Cheilostomatous Bryozoa, Part 2,
Hippothooidea – Celleporoidea. Issue 14. Shrewsbury, UK: Field
Studies Council for The Linnean Society of London and The
Estuarine and Coastal Science Association.
Hilário A, Metaxas A, Gaudron SM, Howell KL, Mercier A,
Mestre NC et al. 2015. Estimating dispersal distance in the deep
sea: challenges and applications to marine reserves. Frontiers in
Marine Science 2: 1–14.
Lange L, Griffiths CL. 2014. Large-scale spatial patterns within
soft-bottom epibenthic invertebrate assemblages along the west
coast of South Africa, based on the Nansen trawl survey. African
Journal of Marine Science 36: 111–124.
Larsen R, Holmern T, Prager SD, Maliti H, Røskaft E. 2009. Using
the extended quarter-degree grid-cell system to unify mapping and
sharing of biodiversity data. African Journal of Ecology 47: 382–392.
Lombard AT, Strauss T, Harris J, Sink K, Attwood C, Hutchings L.
2004. South African National Spatial Biodiversity Assessment
2004: Technical report, volume 4: marine component. Pretoria,
South Africa: South African National Biodiversity Institute.
Lutjeharms JRE. 2006. The ocean environment off southeastern
Africa: a review. South African Journal of Science 102: 419–427.
Maree RC, Whitfield AK, Booth AJ. 2000. Effect of water
temperature on the biogeography of South African estuarine
fishes associated with the subtropical/warm-temperate
subtraction zone. South African Journal of Science 96: 184–188.
Mariani S, Uriz MJ, Turon X, Alcoverro T. 2006. Dispersal
strategies in sponge larvae: integrating the life history of larvae
and the hydrologic component. Oecologia 149: 174–184.
Mead A, Carlton JT, Griffiths CL, Rius M. 2011. Revealing the scale
of marine bioinvasions in developing regions: a South African
re-assessment. Biological Invasions 13: 1991–2008.
Millard NAH. 1978. The geographical distribution of southern
African hydroids. Annals of the South African Museum 74:
159–200.
O’Connor MI, Bruno JF, Gaines SD, Halpern BS, Lester SE,
Kinlan BP, Weiss JM. 2007. Temperature control of larval
dispersal and the implications for marine ecology, evolution, and
conservation. Proceedings of the National Academy of Sciences
of the United States of America 104: 1266–1271.
Boonzaaier-Davids, Florence and Gibbons
194
O’Donoghue CH. 1924. The Bryozoa (Polyzoa) collected by the
S.S. “Pickle.” Fisheries and Marine Biological Survey of South
Africa. Report No. 3 for the year 1992. Special Report 10:
1–63.
O’Donoghue CH. 1957. Some South African Bryozoa. Transactions
of the Royal Society of South Africa 35: 71–95.
O’Donoghue CH, de Watteville D. 1935. A collection of Bryozoa
from South Africa. Journal of the Linnean Society, Zoology 39:
203–218.
O’Donoghue CH, de Watteville D. 1937. Notes on South African
Bryozoa. Zoologischer Anzeiger 117: 12–22.
O’Donoghue CH, de Watteville D. 1944. Additional notes on
Bryozoa from South Africa. Annals of the Natal Museum 10:
407–432.
Oliver JC, Florence WK. 2016. A new species of Taylorius
(Bryozoa: Escharinidae) from the east coast of South Africa.
African Natural History 12: 1–4.
Promińska A, Falck E, Walczowski W. 2018. Interannual variability
in hydrography and water-mass distribution in Hornsund, an
Arctic fjord in Svalbard. Polar Research 37: 1–19.
Rouse S, Spencer Jones ME, Porter JS. 2014. Spatial and
temporal patterns of bryozoan distribution and diversity in the
Scottish sea regions. Marine Ecology 34: 85–102.
Ryland JS. 1965. Catalogue of main fouling organisms (found on
ships coming into European waters). Vol. 2: Polyzoa. Paris:
Organisation for Economic Co-operation and Development.
Samaai T. 2006. Biodiversity “hotspots”, patterns of richness and
endemism, and distribution of marine sponges in South Africa
based on actual and interpolation data: a comparative approach.
Zootaxa 1358: 1–37.
Samaai T, Gibbons MJ, Kerwath S, Yemane D, Sink K. 2010.
Sponge richness along a bathymetric gradient within the
iSimangaliso Wetland Park, South Africa. Marine Biodiversity 40:
205–217.
Scott RJ, Griffiths CL, Robinson TB. 2012. Patterns of endemicity
and range restriction among southern African coastal marine
invertebrates. African Journal of Marine Science 34: 341–347.
Sink KJ, Branch GM, Harris JM. 2005. Biogeographic patterns in
rocky intertidal communities in KwaZulu-Natal, South Africa.
African Journal of Marine Science 27: 81–96.
Sink KJ, van der Bank MG, Majiedt PA, Harris LR, Atkinson LJ,
Kirkman SP, Karenyi N (eds). 2019. South African National
Biodiversity Assessment 2018: Technical report, volume 4:
marine realm. Pretoria, South Africa: South African National
Biodiversity Institute.
Skowno AL, Poole CJ, Raimondo DC, Sink KJ, van Deventer H,
van Niekerk L et al. 2019. National Biodiversity Assessment
2018: the status of South Africa’s ecosystems and biodiversity.
Synthesis report. Pretoria, South Africa: South African National
Biodiversity Institute.
Spalding MD, Fox HE, Allen GR, Davidson N, Ferdaña ZA,
Finlayson M et al. 2007. Marine ecoregions of the world: a
bioregionalization of coastal and shelf areas. BioScience 57:
573–583.
Tanner JE, Althaus F, Sorokin SJ, Williams A. 2018. Benthic
biogeographic patterns in the southern Australian deep sea:
do historical museum records accord with recent systematic,
but spatially limited, survey data? Ecology and Evolution 8:
11423–11433.
Taylor PD. 1988. Major radiation of cheilostome bryozoans:
triggered by the evolution of a new larval type? Historical Biology
1: 45–64.
Taylor PD, Waeschenbach A. 2015. Phylogeny and diversification
of bryozoans. Palaeontology 58: 585–599.
Telford MJ, Budd GE, Philippe H. 2015. Phylogenomic insights into
animal evolution. Current Biology 25: R876–R887.
Teske PR, von der Heyden S, McQuaid CD, Barker NP. 2011.
A review of marine phylogeography in southern Africa. South
African Journal of Science 107: 43–53.
Thandar AS. 1989. Zoogeography of the southern African
echinoderm fauna. South African Journal of Zoology 24:
311–318.
Turpie JK, Beckley LE, Katua SM. 2000. Biogeography and the
selection of priority areas for conservation of South African
coastal fishes. Biological Conservation 92: 59–72.
von der Heyden S. 2009. Why do we need to integrate population
genetics into South African marine protected area planning?
African Journal of Marine Science 31: 263–269.
Watts PC, Thorpe JP. 2006. Influence of contrasting larval
developmental types upon the population-genetic structure of
cheilostome bryozoans. Marine Biology 149: 1093–1101.
Winston JE. 2012. Dispersal in marine organisms without a pelagic
larval phase. Integrative and Comparative Biology 52: 447–457.
Zeppilli D, Pusceddu A, Trincardi F, Danovaro R. 2016. Seafloor
heterogeneity influences the biodiversity-ecosystem functioning
relationships in the deep sea. Scientific Reports 6: 1–12.
Manuscript received April 2020 / revised April 2020 / accepted April 2020
African Journal of Marine Science 2020, 42(2): 185–198
195
Family and species Range
ADEONIDAE
Adeona costata O’Donoghue, 1924 E
Adeonella abdita Hayward & Cook, 1983 E
Adeonella sp. nov. (Boonzaaier-Davids, Florence &
Gibbons in press)
E
Adeonella alia Hayward & Cook, 1983 E
Adeonella aspera Hayward, 1988 E
Adeonella circumspecta Hayward, 1988 E
Adeonella concinna Hayward, 1988 E
Adeonella confusanea Hayward & Cook, 1983 E
Adeonella conspicua Hayward & Cook, 1983 E
Adeonella coralliformis O’Donoghue, 1924 E
Adeonella cracens Hayward & Cook, 1979 E
Adeonella decipiens Hayward & Cook, 1983 E
Adeonella distincta Hayward & Cook, 1983 E
Adeonella expansa O’Donoghue, 1924 E
Adeonella falcicula Hayward, 1981 E
Adeonella fuegensis (Busk, 1852) S
Adeonella gibba Hayward & Cook, 1983 E
Adeonella guttata Hayward, 1988 E
Adeonella infirmata Hayward & Cook, 1983 E
Adeonella ligulata O’Donoghue, 1924 E
Adeonella lobata Hayward, 1988 E
Adeonella majuscula Hayward & Cook, 1979 E
Adeonella pluscula Hayward, 1988 E
Adeonella purpurea Hayward, 1988 E
Adeonella regularis Busk, 1884 E
Adeonella similis Hayward, 1988 E
Adeonella tuberosa Hayward, 1988 E
Adeonellopsis meandrina (O’Donoghue & de
Watteville, 1944)
E
Dimorphocella moderna Hayward & Cook, 1983 E
Laminopora bimunita (Hincks, 1891) I
Laminopora jellyae (Levinsen, 1909) E
AETEIDAE
Aetea anguina (Linnaeus, 1758) C
ALCYONIDIIDAE
Alcyonidium chondroides O’Donoghue & de
Watteville, 1937
E
Alcyonidium ustroides Busk, 1886 E
Alcyonidium nodosum O’Donoghue & de Watteville,
1944
S
Alcyonidium rhomboidale O’Donoghue, 1924 E
ALYSIDIIDAE
Alysidium parasiticum Busk, 1852 I
Catenicula corbulifera O’Donoghue, 1924 –
Catenicula compacta O’Donoghue & de Watteville,
1944
E
ARACHNOPUSIIDAE
Arachnopusia corniculata Hayward & Cook, 1983 E
ASPIDOSTOMATIDAE
Aspidostoma livida Hayward & Cook, 1983 E
Aspidostoma magna Hayward & Cook, 1979 E
Aspidostoma sp. nov. (Boonzaaier-Davids, Florence
& Gibbons in press)
E
BATOPORIDAE
Batopora lagaaiji Hayward & Cook, 1979 E
Batopora murrayi Cook, 1966 S
Batopora nola Hayward & Cook, 1979 E
Lacrimula burrowsi Cook, 1966 I
Lacrimula pyriformis Cook, 1966 S
Appendix: An updated species list of 276 marine Bryozoa, indicating new species (in bold) and newly recorded genera (with an asterisk) in
South Africa from this study (excluding morpho-species). Global distribution range or affinities, where applicable, are indicated as: endemic
(E), Atlantic (A), Indo-Pacific (I), widespread (W), cosmopolitan (C), or scattered (S)
Family and species Range
BEANIIDAE
Beania inermis (Busk, 1852) S
Beania magellanica (Busk, 1852) W
Beania minuspina Florence, Hayward & Gibbons,
2007
E
Beania rediviva Hayward & Cook, 1983 E
Beania uniarmata O’Donoghue & de Watteville, 1944 E
Beania vanhoeeni Kluge, 1914 E
BIFAXARIIDAE
Bifaxaria submucronata Busk, 1884 S
Diplonotos inornatus (Hayward, 1981) S
Domosclerus corrugatus (Busk, 1884) S
Raxifabia longicaulis (Harmer, 1957) S
BITECTIPORIDAE
Bitectipora umboavicula Florence, Hayward &
Gibbons, 2007
E
Hippomonavella formosa (MacGillivray, 1887) S
Hippomonavella sp. nov. (Boonzaaier-Davids,
Florence & Gibbons in press)
E
Schizosmittina lizzya Florence, Hayward & Gibbons,
2007
E
BRYOCRYPTELLIDAE
Porella capensis O’Donoghue, 1924 E
BUGULIDAE
Bicellariella bonsai Florence, Hayward & Gibbons,
2007
E
Bicellariella chuakensis (Waters, 1913) I
Bugula neritina (Linnaeus, 1758) C
Bugula robusta MacGillivray, 1869 I
Bugulella gracilis (Nichols, 1911) W
Bugulella problematica Hayward & Cook, 1983 E
Bugulina avicularia (Linnaeus, 1758) S
Bugulina abellata (Thompson in Gray, 1848) W
Cornucopina novissima Hayward, 1981 I
Himantozoum leontodon (Busk, 1884) W
Kinetoskias cyathus (Wyville Thomson, 1873) A
Kinetoskias elegans Menzies, 1963 A
Kinetoskias pocillum Busk, 1881 W
Virididentula dentata (Lamouroux, 1816) W
BUSKIIDAE
Cryptopolyzoon concretum (Dendy, 1889) I
CALLOPORIDAE
Amphiblestrum pontifex Hayward & Cook, 1983 E
Amphiblestrum triangularis (O’Donoghue, 1924) S
Callopora jamesi O’Donoghue & de Watteville, 1944 E
Crassimarginatella marginalis (Kirkpatrick, 1888) I
CALWELLIDAE
Onchoporella buskii (Harmer, 1923) E
CANDIDAE
Caberea darwinii Busk, 1884 S
Caberea darwinii occlusa Hastings, 1943 E
Eupaxia quadrata (Busk, 1884) I
Hoplitella armata (Busk, 1852) E
Menipea crispa (Pallas, 1766) E
Menipea marionensis Busk, 1884 E
Menipea ornata (Busk, 1852) E
Menipea triseriata Busk, 1852 E
Notoplites candoides Hayward & Cook, 1979 E
Notoplites cassidula Hayward & Cook, 1979 E
Tricellaria varia Hayward & Cook, 1979 E
Boonzaaier-Davids, Florence and Gibbons
196
Family and species Range
CATENICELLIDAE
Catenicella elegans Busk, 1852 W
Cornuticella taurina (Busk, 1852) I
Costaticella carotica Hayward & Cook, 1979 E
CELLARIIDAE
Cellaria mandibulata Hincks, 1882 I
Cellaria paradoxa Hayward & Cook, 1979 E
Cellaria punctata (Busk, 1852) I
Smitticellaria tectiformis (Hayward & Cook, 1979) E
CELLEPORIDAE
Cellepora pustulata Busk, 1881 –
Celleporina solida Florence, Hayward & Gibbons, 2007 E
Galeopsis bispiramina (Hayward & Cook, 1979) E
Galeopsis circella (Hayward & Cook, 1979) E
Galeopsis pentagonus (d’Orbigny, 1842) W
Lagenipora echinacea Marcus, 1922 S
Lagenipora spinifera O’Donoghue, 1924 E
Nigrapercula mutabilis (Canu & Bassler, 1929) I
Turbicellepora avicularis (Hincks, 1860) S
Turbicellepora canaliculata (Busk, 1881) A
Turbicellepora conica (Busk, 1881) E
Turbicellepora coralliformis Hayward, 1980 A
Turbicellepora protensa Hayward & Cook, 1979 E
Turbicellepora redoutei (Audouin, 1826) I
Turbicellepora valligera Hayward & Cook, 1983 E
CHAPERIIDAE
Chaperia acanthina (Lamouroux, 1825) –
Chaperia capensis (Busk, 1884) E
Chaperia septispina Florence, Hayward & Gibbons,
2007
E
Chaperiopsis sp. nov. (Boonzaaier-Davids, Florence
& Gibbons in press)
–
Chaperiopsis chelata Fernandez Pulpeiro & Reverter
Gil, 1998
E
Chaperiopsis cylindracea (Busk, 1884) S
Chaperiopsis familiaris Hayward & Cook, 1983
comb. nov. (Boonzaaier-Davids, Florence & Gibbons
in press)
E
Chaperiopsis multida (Busk, 1884) E
Chaperiopsis stephensoni (O’Donoghue & de
Watteville, 1935)
E
Notocoryne cervicornis Hayward & Cook, 1979 E
CHORIZOPORIDAE
Chorizopora brongniartii (Audouin, 1826) C
CLEIDOSCHASMATIDAE
Characodoma cribritheca (Busk, 1884) E
Characodoma protrusum (Thornely, 1905) S
Cleidochasma anis (Milne Edwards, 1836) –
CONESHARELLINIDAE
Conescharellina africana Cook, 1966 I
CRIBRILINIDAE
Cribrilaria innominata (Couch, 1844) S
Cribrilaria venusta (Canu & Bassler, 1925) S
Cribrilina dispersa O’Donoghue & de Watteville, 1937 E
Cribrilina simplex O’Donoghue & de Watteville, 1935 E
Figularia ssa (Hincks, 1880) I
Figularia philomela (Busk, 1884) I
Glabrilaria africana (Hayward & Cook, 1983) E
Inversiscaphos setifer Hayward & Cook, 1979 E
Khulisa gen. nov., sp. nov. (Boonzaaier-Davids,
Florence & Gibbons in press)*
E
CRISIIDAE
Bicrisia edwardsiana (d’Orbigny, 1841) W
Appendix: (cont.)
Family and species Range
Crisia elongata Milne Edwards, 1838 W
Crisidia cornuta (Linnaeus, 1758) A
Mesonea radians (Lamarck, 1816) I
CRYPTOSULIDAE
Cryptosula pallasiana (Moll, 1803) W
CUPULADRIIDAE
Discoporella umbellata (Defrance, 1823) A
Discoporella umbellata ‘peyroti’ Hayward & Cook 1983 E
Reussirella multispinata (Canu & Bassler, 1923) A
DENSIPORIDAE
Favosipora sp. nov. (Boonzaaier-Davids, Florence &
Gibbons in press)*
E
DIAPEROECIIDAE
Nevianipora pulcherrima (Kirkpatrick, 1890) W
ELECTRIDAE
Conopeum reticulum (Linnaeus, 1767) C
Electra pilosa (Linnaeus, 1767) C
ENTALOPHORIDAE
?Mecynoecia australis (Busk, 1852) I
Mecynoecia clavaeformis (Busk, 1875) E
Mecynoecia delicatula (Busk, 1875) I
ESCHARINIDAE
Hippomenella avicularis (Livingstone, 1926) I
Taylorius waiparaensis (Brown, 1952) –
EUTHYRISELLIDAE
Tropidozoum burrowsi Cook & Chimonides, 1981 S
EXOCHELLIDAE
Escharoides contorta (Busk, 1854) E
Escharoides custodis Florence, Hayward & Gibbons,
2007
E
Escharoides distincta Hayward & Cook, 1979 E
FARCIMINARIIDAE
Columnella accincta Hayward, 1981 E
Columnella magna (Busk, 1884) W
Farciminellum hexagonum (Busk, 1884) I
FLUSTRELLIDRIDAE
Elzerina blainvillii Lamouroux, 1816 I
FLUSTRIDAE
Carbasea elegans Busk, 1852 I
Carbasea mediocris Hayward & Cook, 1979 E
Gregarinidra spinuligera (Hincks, 1891) E
FOVEOLARIIDAE
Dactylostega prima Hayward & Cook, 1983 E
Dactylostega tubigera (Busk, 1884) E
Foveolaria imbricata (Busk, 1884) E
GIGANTOPORIDAE
Gephyrophora polymorpha Busk, 1884 E
HELIODOMIDAE
Heliodoma implicata Calvet, 1906 A
Setosellina roulei Calvet, 1906 A
HETEROPORIDAE
Heteropora pelliculata Waters, 1879 –
HIPPOPORIDRIDAE
Fodinella spinigera (Philipps, 1900) S
Hippoporella labiata Hayward & Cook, 1983 E
Hippoporidra senegambiensis (Carter, 1882) A
HIPPOTHOIDAE
Celleporella annularis (Pallas, 1766) W
Celleporella hyalina (Linnaeus, 1767) S
Hippothoa musivaria Hayward & Fordy, 1982 E
HORNERIDAE
Hornera americana d’Orbigny, 1842 A
Hornera erugata Hayward & Cook, 1983 E
Appendix: (cont.)
African Journal of Marine Science 2020, 42(2): 185–198
197
Family and species Range
INCERTAE SEDIS
Klugeustra jonesii Florence, Hayward & Gibbons,
2007
E
Lepralia clithridiata O’Donoghue, 1924 E
LACERNIDAE
Arthropoma cecilii (Audouin, 1826) W
Arthropoma lioneli Florence, 2016 E
Phonicosia circinata (MacGillivray, 1869) I
LANCEOPORIDAE
Calyptotheca capensis (O’Donoghue & de Watteville,
1937)
E
Calyptotheca nivea (Busk, 1884) W
Calyptotheca porelliformis (Waters, 1918) E
Emballotheca ambigua Hayward & Cook, 1983 E
LEIOSALPINGIDAE
Leiosalpinx inornata (Goldstein, 1882) E
LEKYTHOPORIDAE
Turritigera stellata Busk, 1884 S
LEPRALIELLIDAE
Celleporaria capensis (O’Donoghue & de Watteville,
1935)
E
Celleporaria tridenticulata (Busk, 1881) S
LICHENOPORIDAE
Disporella buski (Harmer, 1915) W
Disporella novaehollandiae (d’Orbigny, 1853) S
Patinella radiata (Audouin, 1826) S
Patinella verrucaria (Linnaeus, 1758) S
MACROPORIDAE
Macropora africana Hayward & Cook, 1983 E
MALILLOPORIDAE
Anoteropora inarmata Cook, 1966 E
Anoteropora latirostris Silén, 1947 W
MARGARETTIDAE
Margaretta levinseni (Canu & Bassler, 1930) I
Margaretta opuntioides (Pallas, 1766) I
MEMBRANIPORIDAE
Biustra sp. nov. (Boonzaaier-Davids, Florence &
Gibbons in press)*
E
Jellyella tuberculata (Bosc, 1802) W
Membranipora rustica Florence, Hayward & Gibbons,
2007
E
MICROPORELLIDAE
Fenestrulina elevora Florence, Hayward & Gibbons,
2007
E
Fenestrulina indigena Hayward & Cook, 1983 E
Flustramorpha angusta Hayward & Cook, 1979 E
Flustramorpha abellaris (Busk, 1854) E
Flustramorpha marginata (Krauss, 1837) A
Microporella sp. nov. (Boonzaaier-Davids, Florence
& Gibbons in preparation)
E
Microporella madiba Florence, Hayward & Gibbons,
2007
E
MICROPORIDAE
Micropora sp. nov. (Boonzaaier-Davids, Florence
& Gibbons in press)
E
Micropora latiavicula Florence, Hayward & Gibbons, 2007 E
Micropora similis Hayward & Cook, 1983 I
ONCOUSOECIIDAE
Eurystrotos planus Florence, Hayward & Gibbons, 2007 E
PETALOSTEGIDAE
Petalostegus bicornis (Busk, 1884) I
Appendix: (cont.)
Family and species Range
PETRALIELLIDAE
Mucropetraliella asymmetrica Hayward & Cook, 1983 E
PHIDOLOPORIDAE
Iodictyum osculum Hayward & Cook, 1983 E
Phidolopora sp. nov. (Boonzaaier-Davids, Florence &
Gibbons in press)*
E
Plesiocleidochasma perspicuum (Hayward & Cook,
1983)
E
Reteporella bullata (Hayward & Cook, 1979) E
Reteporella clancularia Hayward & Cook, 1979 E
Reteporella dinotorhynchus Hayward & Cook, 1979 E
Reteporella gilchristi (O’Donoghue, 1924) S
Reteporella sp. nov. (Boonzaaier-Davids, Florence &
Gibbons in press)
E
Reteporella lata (Busk, 1884) E
Reteporella magellensis (Busk, 1884) A
Reteporella verecunda (Hayward & Cook, 1983) E
Rhynchozoon abscondum Florence, Hayward &
Gibbons, 2007
E
Rhynchozoon beatulum Hayward & Cook, 1983 E
Rhynchozoon documentum Hayward & Cook, 1983 E
Rhynchozoon fulgidum O’Donoghue & de Watteville,
1935
E
Rhynchozoon incallidum Hayward & Cook, 1983 E
Rhynchozoon longirostris (Hincks, 1881) I
Rhynchozoon oscitans Hayward & Cook, 1983 E
Rhynchozoon ptarmicum Hayward & Cook, 1983 E
Rhynchozoon stomachosum Hayward & Cook, 1983 E
Schizoretepora tessellata (Hincks, 1878) S
Stephanollona ignota (Hayward & Cook, 1983) I
PLAGIOECIIDAE
Desmeplagioecia lineata (MacGillivray, 1885) I
Plagioecia patina (Lamarck, 1816) S
RHABDOZOIDAE
Rhabdozoum stephensoni O’Donoghue & de Watteville,
1944
E
ROMANCHEINIDAE
Escharella anatirostris (O’Donoghue, 1924) E
Escharella discors Hayward & Cook, 1983 E
Escharella serratilabris (O’Donoghue, 1924) E
SCHIZOPORELLIDAE
Schizoporella inconspicua Hincks, 1891 E
Schizoporella unicornis (Johnston in Wood, 1844) W
SCRUPARIIDAE
Scruparia ambigua (d’Orbigny, 1841) C
Scruparia chelata (Linnaeus, 1758) S
Scruparia spiralis Hasenbank, 1932 E
SMITTINIDAE
Parasmittina novella Hayward & Cook, 1983 E
Parasmittina tropica (Waters, 1909) W
Smittina ferruginea Hayward & Cook, 1983 E
Smittina landsborovii (Johnston, 1847) S
Smittina sitella Hayward & Cook, 1983 E
Smittoidea calcarata Hayward & Cook, 1983 E
Smittoidea circumspecta Hayward & Cook, 1983 E
Smittoidea errata Hayward & Cook, 1983 E
Smittoidea hexagonalis (O’Donoghue, 1924) E
STEGINOPORELLIDAE
Steginoporella buskii Harmer, 1900 W
STOMACHETOSELLIDAE
Stomachetosella balani (O’Donoghue & de Watteville,
1944)
E
Appendix: (cont.)
Boonzaaier-Davids, Florence and Gibbons
198
Family and species Range
THALAMOPORELLIDAE
Thalamoporella spiravicula Florence, Hayward &
Gibbons, 2007
E
TRYPOSTEGIDAE
Trypostega sp. nov. (Boonzaaier-Davids, Florence &
Gibbons in press)
E
Trypostega venusta (Norman, 1864) W
TUBULIPORIDAE
Exidmonea atlantica (Forbes in Johnston, 1847) A
Exidmonea crassimargo (Canu & Bassler, 1929) E
Idmidronea contorta (Busk, 1875) E
Idmidronea parvula (Canu & Bassler, 1929) –
Tennysonia stellata Busk, 1867 E
VESICULARIIDAE
Amathia gracilis (Leidy, 1855) C
Amathia lendigera (Linnaeus, 1758) W
WATERSIPORIDAE
Watersipora subtorquata (d’Orbigny, 1852) W
Appendix: (cont.)
