Taxonomy and diversity of the sponge fauna from Walters Shoal,
a shallow seamount in the Western Indian Ocean region
By
Robyn Pauline Payne
A thesis submitted in partial fulfilment of the requirements for the degree of Magister
Scientiae in the Department of Biodiversity and Conservation Biology, University of the
Western Cape.
Supervisors: Dr Toufiek Samaai
Prof. Mark J. Gibbons
Dr Wayne K. Florence
The financial assistance of the National Research Foundation (NRF) towards this research is
hereby acknowledged. Opinions expressed and conclusions arrived at, are those of the author
and are not necessarily to be attributed to the NRF.
December 2015
Taxonomy and diversity of the sponge fauna from Walters Shoal, a shallow
seamount in the Western Indian Ocean region
Robyn Pauline Payne
Keywords
Indian Ocean
Seamount
Walters Shoal
Sponges
Taxonomy
Systematics
Diversity
Biogeography
ii
Abstract
Taxonomy and diversity of the sponge fauna from Walters Shoal, a shallow seamount in
the Western Indian Ocean region
R. P. Payne
MSc Thesis, Department of Biodiversity and Conservation Biology, University of the
Western Cape.
Seamounts are poorly understood ubiquitous undersea features, with less than 4% sampled
for scientific purposes globally. Consequently, the fauna associated with seamounts in the
Indian Ocean remains largely unknown, with less than 300 species recorded. One such
feature within this region is Walters Shoal, a shallow seamount located on the South
Madagascar Ridge, which is situated approximately 400 nautical miles south of Madagascar
and 600 nautical miles east of South Africa. Even though it penetrates the euphotic zone
(summit is 15 m below the sea surface) and is protected by the Southern Indian Ocean DeepSea Fishers Association, there is a paucity of biodiversity and oceanographic data. Thus, a
multidisciplinary cruise was initiated in May 2014 on the FRS Algoa as a component of the
African Coelacanth Ecosystem Programme. The research presented here focuses exclusively
on the diversity, bathymetric distribution patterns and biogeographic affiliations of the
sponge fauna of this seamount.
Sponges were sampled using SCUBA and a roughed epibenthic sled, from the peak and down
two opposing slopes of the seamount, to a depth of 500 m. Two hundred and fifty-five sponge
specimens were collected, comprising 78 operational taxonomic units (OTU’s), 23 of which
are known to science, 26 which are possibly new, 16 that could only be identified to higher
iii
taxonomic levels and 13 that could only be designated as morphospecies. Thirteen OTU’s are
formally described here, four which are known, and nine possibly new to science.
Sponge assemblages demonstrated no significant difference according to location on the
shoal, with several species shared by both the western and eastern flanks. In contrast, sponge
assemblages differed significantly according to depth, with the mesophotic zone (31 – 150 m)
acting as a transition between the shallow (15 – 30 m) and submesophotic (> 150 m) zones.
Species richness and the number of putative new species was highest in the submesophotic
zone. Biogeographical affiliations were found with both the Western Indo-Pacific and
Temperate Southern African realms based on the 23 known species recorded. No affiliations
were found with the West Wind Drift Island Province, as has been documented previously for
the fish fauna of this seamount, possibly due to the incomplete nature of the online database
(World Porifera Database) used to assess affinities. Thirty-nine percent of the known sponge
species found at Walters Shoal Seamount are widely distributed in the Indian Ocean, 35% are
found exclusively within the Western Indian Ocean region, with this study representing the
southernmost distribution record for several of these, and 26% have a restricted distribution
around South Africa.
December 2015
iv
Declaration
I declare that Taxonomy and diversity of the sponge fauna from Walters Shoal, a shallow
seamount in the Western Indian Ocean region is my own work, that is has not been submitted
for any degree or examination in any other university, and that all the sources I have used or
quoted have been indicated and acknowledged by complete references.
Robyn Pauline Payne
December 2015
Signed:
v
Acknowledgements
This thesis, as well as conference attendance to present findings (15th South African Marine
Science Symposium (SAMSS): 2014, 14th Deep-Sea Biology Symposium (DSBS): 2015 and
the 9th Western Indian Ocean Marine Science Association (WIOMSA) Scientific
Symposium: 2015), was made possible via funding provided by the African Coelacanth
Ecosystem Programme (ACEP (DST-NRF)), with bursary funding provided by the National
Research Foundation (NRF).
Resources for the cruise to collect samples included in this study were jointly provided by the
South African Institute for Aquatic Biodiversity (SAIAB), the South African Environmental
Observation Network (SAEON), the Department of Environmental Affairs (DEA) and the
Department of Agriculture, Forestry and Fisheries (DAFF). As such, I am grateful to all
involved in the cruise, including the captain of the RV Algoa (Geordie MacKenzie), officers
and crew, as well as the staff members from the institutions mentioned above whose
contributions made this project a reality.
More specifically, I would like to express my sincere gratitude to my supervisors: Dr Toufiek
Samaai (DEA), Prof Mark J. Gibbons (University of the Western Cape) and Dr Wayne K.
Florence (Iziko Museum) for their ongoing assistance, guidance and advice. Special thanks
are extended to Dr Toufiek Samaai for introducing me to the fascinating world of sponge
taxonomy. A word of thanks is also extended to Liesl Janson (DEA) and Seshnee Maduaray
(DEA), for their continued help with sponge processing techniques, thesis advice and
friendship; Dr Tanya Haupt-Schuter (DEA), for advice and reading an earlier version of this
manuscript; Miranda Waldron (Electron Microscope Unit, UCT) for help with the scanning
electron microscope and Marcel van den Berg (DEA) for assistance with Surfer 9 and
compiling the bathymetric figures of Walters Shoal Seamount.
vi
My sincerest thanks go to the editors of the World Porifera Database (especially Rob van
Soest) for their quick response, access to data and key taxonomic papers, interest in future
collaborations, overall friendliness and willingness to assist.
I would be remiss if I didn’t extend thanks to all the people that have brightened my days at
the Department of Environmental Affairs, including: Asma Damon, Darrell Anders, Imtiyaaz
Malick, Jabulile Nhleko, Keshnee Pillay, Leon Jacobs, Marco Worship, Dr Maya Pfaff, Dr
Stephen Kirkman and Taryn Joshua.
Special thanks are extended to my closest friend, Zoleka Filander (DEA), for her belief in me
and mentorship. Finally, I wish to thank my family: my parents, grandmother and two sisters
for never-ending support and encouragement.
vii
Contents
Title Page
Keywords
Abstract
Declaration
Acknowledgements
Contents Page
List of Figures
List of Tables
Chapter 1 – Introduction
1.1 A deep-sea habitat: the seamount
1.2 Seamounts of the Indian Ocean
1.3 Why sponges?
1.4 Hypotheses
i
ii
iii
v
vi
viii
ix
xi
1 – 21
2
10
15
20
Chapter 2 – Methodology
2.1 Collection
2.2 Taxonomic procedure
2.3 Sample storage
2.4 Location and depth analyses
2.5 Biogeography analyses
22 – 26
22
22
25
25
26
Chapter 3 – Results
3.1 Systematics
3.2 Descriptions
3.3 Location and depth affiliations
3.4 Biogeographical affiliations
27 – 66
27
28
62
64
Chapter 4 – Discussion
4.1 Diversity
4.2 Location and depth affiliations
4.3 Biogeographical affiliations
4.4 Study limitations and future work
4.5 Conclusion
67 – 77
67
69
72
75
77
Figures
Tables
References
Appendix
78 – 100
101 – 131
132 – 160
161 – 175
viii
List of Figures
Fig. 1:
Map showing the location of Walters Shoal Seamount within the
78
bathymetric context of the Western Indian Ocean region.
Fig. 2:
Bathymetric map of Walters Shoal Seamount.
79
Fig. 3:
Sponge specimen sampling strategies: SCUBA dives and the use of a
80
roughed epibenthic sled.
Fig. 4:
Bathymetric map of Walters Shoal Seamount with sampling locations.
81
Fig. 5:
Sheet completed per sponge specimen to denote macroscopical features.
82
Fig. 6:
Map showing the ecoregions, as defined by Spalding et al. (2007),
83
surrounding Walters Shoal Seamount that were included in the
biogeographical analyses.
Fig. 7:
Plate of Agelas ceylonica Dendy, 1905.
84
Fig. 8:
Plate of Ptilocaulis sp. •
85
Fig. 9:
Plate of Callyspongia (Callyspongia) sp. •
86
Fig. 10:
Plate of Lissodendoryx (Lissodendoryx) pygmaea (Burton, 1931).
87
Fig. 11:
Plate of Fibulia ectofibrosa (Lévi, 1963).
88
Fig. 12:
Plate of Latrunculia (Biannulata) sp. •
89
Fig. 13:
Plate of Clathria (Clathria) sp. •
90
ix
Fig. 14:
Plate of Halichondria (Halichondria) sp. •
91
Fig. 15:
Plate of Aaptos sp. •
92
Fig. 16:
Plate of Tethya sp. •
93
Fig. 17:
Plate of Ancorina sp. •
94
Fig. 18:
Plate of Chelotropella sp. •
95
Fig. 19:
Plate of Penares intermedia (Dendy, 1905).
96
Fig. 20:
Non-metric MDS ordination of sampling locations according to location.
97
Fig. 21:
Non-metric MDS ordination of sampling locations according to depth.
98
Fig. 22:
Biogeographical affinities of the 23 known sponge species recorded
99
from Walters Shoal Seamount.
Fig. 23:
The low spatial complexity and growth profile of Walters Shoal
100
Seamount.
x
List of Tables
Table 1:
Invertebrate (including sponge) collection sampling strategy.
101
Table 2:
Microwave 5mm/2 layer method for sponge specimen histology
102
processing.
Table 3:
Ecoregions included in the biogeographical analyses.
103
Table 4:
Sponge species documented from Walters Shoal Seamount per sampling
104
location.
Table 5:
Sponge species documented from Walters Shoal Seamount.
106
Table 6:
Spicule dimensions of Agelas ceylonica Dendy, 1905.
112
Table 7:
Spicule dimensions of Ptilocaulis sp. •
112
Table 8:
Spicule dimensions of Callyspongia (Callyspongia) sp. •
112
Table 9:
Spicule dimensions of Lissodendoryx (Lissodendoryx) pygmaea
113
(Burton, 1931).
Table 10:
Spicule dimensions of Fibulia ectofibrosa (Lévi, 1963).
113
Table 11:
Spicule dimensions of Clathria (Clathria) sp. •
114
Table 12:
Spicule dimensions of Halichondria (Halichondria) sp. •
114
Table 13:
Spicule dimensions of Aaptos sp. •
115
Table 14:
Spicule dimensions of Tethya sp. •
115
xi
Table 15:
Spicule dimensions of Ancorina sp. •
116
Table 16:
Spicule dimensions of Penares intermedia (Dendy, 1905).
117
Table 17:
Walters Shoal Seamount sponge species list per location.
118
Table 18:
SIMPER results – percentage contribution of each species that overall
120
contribute to at least 60% of the difference between the western and
eastern flank of Walters Shoal Seamount.
Table 19:
Walters Shoal Seamount sponge species list per depth zone.
121
Table 20:
SIMPER results – species that contribute to 90% (100% in
123
submesophotic zone) of sampling location similarity in each depth zone.
Table 21:
Walters Shoal Seamount sponges – percent contribution of higher
124
taxonomic levels (families and genera) per depth zone.
Table 22:
Walters Shoal Seamount – sponge families per depth zone.
125
Table 23:
Walters Shoal Seamount – sponge genera per depth zone.
126
Table 24:
SIMPER results – percentage contribution of each species that overall
127
contribute to at least 60% of the difference between depth zones.
Table 25:
Biogeographical affinities of the Walters Shoal Seamount sponge fauna
129
based on the 23 known species from this study.
Table 26:
The most represented sponge families and genera per ecoregion that was
130
found to have biogeographical affiliations with Walters Shoal Seamount.
Table A:
Taxonomic sponge species list per ecoregion included in the
161
biogeographical analyses, compiled from the World Porifera Database.
xii
‘On the undersea mountains live myriads of animals, particularly attached forms, which of
all deep-sea organisms are least accessible to the biologist.’
– Marshall (1979)
‘Red, orange, violet, or yellow, they [sponges] stand out against the
whiteness of the sand or are projected on the greenish background of rocks and look like
fantastically beautiful shrubbery planted by an unknown hand in a submarine garden.’
– Galtsoff (1960)
xiii
Chapter 1 – Introduction
The deep sea is the largest ecosystem on Earth, constituting approximately 90% of the ocean
(Gage & Tyler 1991, Ramirez-Llodra et al. 2010). Yet, due to the remote nature of this
environment and the expense of research, including the fickle nature of funding (Gage &
Tyler 1991, Rex & Etter 2010), only 5% of the deep sea has been investigated via remote
equipment, and less than 0.01% of the deep-sea floor has been sampled in any detail
(Ramirez-Llodra et al. 2010). Thus, this region is the least explored and understood
ecosystem on Earth (Ramirez-Llodra et al. 2010). The lack of scientific knowledge about the
deep sea not only permits creative speculation from researchers in this field (Batson 2003),
but also enabled paradigms to be perpetuated far longer than justified (Gage & Tyler 1991).
This is illustrated in the previously prolonged opinion that the unlit, cold and energy-deprived
nature of the deep sea shaped an environment that was not conducive for life (Rex 1981, Rex
& Etter 2010, Ramirez-Llodra et al. 2011).
To date, from samples collected, aided increasingly by new technology and a relatively recent
international research effort, it has been revealed that the deep sea supports one of the highest
levels of biodiversity on earth (Smith et al. 2008, Blaustein 2010, Costello et al. 2010,
Ramirez-Llodra et al. 2010, Rex & Etter 2010). This ecosystem is also comprised of a variety
of distinct habitats, with twenty eight new habitats having been discovered in the deep sea
since the 1840s (Ramirez-Llodra et al. 2010). Subsequently, this ecosystem is now viewed as
comprising vast expanses of continental slope and abyssal plains, interspersed with other
geological features that host unique faunal communities (Ramirez-Llodra et al. 2010). The
seamount habitat is one such feature.
1
1.1 A deep-sea habitat: the seamount
Seamounts are submerged inactive volcanoes, otherwise known as undersea mountains,
which are predominantly found on the oceanic crust (Gage & Tyler 1991, Batson 2003,
Wessel 2007, Ramirez-Llodra et al. 2010). The definition of this geological feature is subject
to a significant amount of inconsistency and ambiguity (Pitcher et al. 2007, Staudigel et al.
2010), largely due to differences in the way that scientists from different disciplines define
them (Staudigel et al. 2010). These features were originally defined as isolated peaks with an
elevation greater than 1000 m above the seafloor (Menard 1964, Rogers 1994, Pitcher et al.
2007) due to difficulties in distinguishing smaller seamounts from the seafloor topography
(Schmidt & Schmincke 2000, Staudigel et al. 2010). However, with no geological or
ecological reason to separate smaller volcanic features from larger ones (Schmidt &
Schmincke 2000, Wessel 2007, Consalvey et al. 2010, Clark et al. 2011, Yesson et al. 2011),
this 1000 m constraint has been relaxed (Pitcher et al. 2007). Currently, the definition
includes most features which rise more than 100 m from the seafloor (Smith & Cann 1992,
Schmidt & Schmincke 2000, Staudigel et al. 2010, Kvile et al. 2014), with this cut-off chosen
as features of this size can largely be recognised as individual volcanoes (Staudigel et al.
2010).
Seamounts may occur in isolation, clusters or chains (Schmidt & Schmincke 2000, Batson
2003, Mladenov 2013), but overall they constitute approximately 2.6 – 4.7% of the seafloor
(Ramirez-Llodra et al. 2010,Yesson et al. 2011). These features are ubiquitous and are
distributed unevenly among the ocean basins (Kitchingman & Lai 2004, Wessel 2007,
Consalvey et al. 2010, Wessel et al. 2010), with most found in the Pacific (Wessel 2007).
That said, the global distribution and abundance of seamounts is difficult to estimate
(Schmidt & Schmincke 2000, Kitchingman & Lai 2004, Consalvey et al. 2010, Yesson et al.
2
2011) as these factors are dependent on the resolution of bathymetric maps used, as well as
how these features are defined in a given study (Kitchingman & Lai 2004).
Yesson et al. (2011) notes several studies that have attempted to determine global seamount
abundance. Although estimates vary, most suggest that, based on extrapolation, more than
100 000 large seamounts exist (Wessel 2001, Wessel 2007, Wessel et al. 2010), with this
number rising into the millions when the smallest seamounts are included (Hillier & Watts
2007, Wessel 2007). More recently, Yesson et al. (2011) used high resolution satellite
bathymetry data to compile the largest global set of seamounts and knolls. These authors
identified 33 452 seamounts (using the 1000 m cut-off) worldwide.
Regardless of the widespread nature of this habitat, less than 300 (0.4 – 4%) seamounts have
been directly sampled for scientific purposes globally (Kvile et al. 2014). This lack of
sampling could possibly be attributed to logistical difficulties associated with their steep,
rocky topography (Rex & Etter 2010, Williams et al. 2015). In spite of this, an increasing
amount of work is being done on seamounts, with some progress being made with regards to
documenting and understanding the biodiversity and connectivity of the biological
communities that inhabit them (Clark et al. 2010).
The biodiversity of seamount communities
Seamounts have unique hydrographic conditions, brought about by their raised topography
and complex rocky substratum that differs considerably from the soft sediments of the
surrounding deep-sea floor (Glover & Smith 2003, Ramirez-Llodra et al. 2010).
Consequently, these features are regions of increased productivity, which support abundant
3
benthic and pelagic communities (Batson 2003, Glover & Smith 2003, Ramirez-Llodra et al.
2011).
Sessile epifaunal suspension feeders generally colonise the slopes of these underwater
features (Marshall 1979, Wilson & Kaufman 1987, Samadi et al. 2007, Consalvey et al.
2010, Ramirez-Llodra et al. 2010), taking advantage of current amplifications that increase
food supply (Genin 2004, Ramirez-Llodra et al. 2010), remove sediment (Genin 2004) and
play a role in larval transport (Consalvey et al. 2010). This group of organisms is dominated
by cnidarians (Samadi et al. 2007, Consalvey et al. 2010), although sponges, crinoids,
molluscs, ascidians and cirripeds are also prominent (Rogers 1994, Samadi et al. 2007,
Consalvey et al. 2010). These benthic assemblages act as biogenic environments that host
numerous mobile species and which form an element of the surprisingly complex seamount
food web (Samadi et al. 2007).
The ichthyofauna represent another component of the seamount ecosystem, and a total of
almost 800 associated fish species have been recorded by Morato et al. (2004). Seamounts
generally support an elevated plankton and fish biomass when compared to surrounding
waters, especially in oligotrophic oceans (Clark et al. 2010). This is attributed to the
enhanced productivity over these features, which is a difficult phenomenon to understand due
to sparse data availability (Batson 2003, Ramirez-Llodra et al. 2010).
It was first thought that hydrographic events, such as upwelling and eddies, around
seamounts enhanced local surface primary productivity, fuelling higher trophic levels
(Consalvey et al. 2010). To date, there is little evidence to support this theory, with upwelling
rarely penetrating the photic layer nor persisting long enough to enable zooplankton growth
(Genin & Dower 2007, White et al. 2007). Current theories suggest that the food supply is
imported from elsewhere (Consalvey et al. 2010), including topographically trapped
4
vertically migrating zooplankton and/or horizontally advected micronekton (Genin 2004,
Genin & Dower 2007, Kvile et al. 2014).
Nonetheless, the enhanced productivity of seamounts attracts numerous top-level predators
(Worm et al. 2003). These include tuna, billfish, sharks, cetaceans, pinnipeds, turtles and
seabirds (Batson 2003, Holland & Grubbs 2007, Kaschner 2007, Litvinov 2007, Santos et al.
2007, Thompson 2007). This attraction could also be attributed to the role these features
might play in navigation (Holland & Grubbs 2007, Kaschner 2007), or as a breeding ground
(Litvinov 2007).
With such diverse assemblages of benthic organisms, ichthyofauna and other visiting mobile
species, seamounts are often referred to as biodiversity hotspots (McClain 2007, Samadi et al.
2007). As a result, there is substantial research interest in this habitat, often motivated by the
notion that seamounts host unique communities that are dissimilar to those that inhabit the
surrounding deep sea (Stocks & Hart 2007).
The distinctness and connectivity of seamount communities
A key question in seamount research is the extent to which seamounts represent isolated
habitats with unique communities (Stocks & Hart 2007). Initially, it was thought that
seamounts act as stepping stones across the ocean basins, facilitating species dispersal
(Hubbs 1959, Stocks & Hart 2007, Shank 2010). Conversely, numerous records of endemic
seamount species (Wilson & Kaufmann 1987, de Forges et al. 2000), suggest that these
features act more like biological islands as a result of geographic isolation and unique
physical conditions (Clark et al. 2010). To date, evidence supporting the ability of these
5
factors to create a distinct community is mixed (Stocks & Hart 2007, Consalvey et al. 2010,
Shank 2010).
Plankton and pelagic fish species that inhabit seamounts seem to be similar to, or the same as,
those from nearby oceanic pelagic communities, and endemics are not reported often (Stocks
& Hart 2007, Shank 2010). On the other hand, the benthic fish and invertebrates seem to
differ slightly more from the surrounding seafloor or continental margins and have higher
rates of endemism (Stocks & Hart 2007). This could be attributed to the distinctness of the
seamount habitat, and thus environmental factors, when compared to the surrounding area
(Stocks & Hart 2007). The benthic community is also determined by depth, according to
environmental gradients (such as temperature and oxygen concentration) that are associated
with this factor (Stocks & Hart 2007, Clark et al. 2010, Consalvey et al. 2010).
Seamounts generally span a spectrum of endemicity (Stocks & Hart 2007), and too little work
has been done on these features to enable or support any generalizations on this topic (Shank
2010). This is also the case for many other theories that have been ascribed to seamounts
(Kvile et al. 2014). Describing these deep-sea habitats as biodiversity hotspots,
biogeographical islands and oases which host lush sponge or coral gardens are tenets, many
of which have become prevalent in the literature and the minds of those working on these
features (Rowden et al. 2010). Yet, their accuracy has been called into question (McClain
2007, Rowden et al. 2010), with Samadi et al. (2007) suggesting that many seamount ‘traits’,
such as archaism and endemism, may be artefacts of the increased sampling and work done
on seamounts when compared to other deep-sea environments. This increased knowledge of
seamounts is a by-product of fisheries studies (Brewin et al. 2007, Samadi et al. 2007,
Consalvey et al. 2010), with commercial fishing having the largest negative anthropogenic
impact on this habitat (Clark et al. 2010).
6
Anthropogenic threats to seamounts
In the late 1960s and 1970s, the former Soviet Union began an intensive global search for
seamount fishery resources (Clark et al. 2007). These searches were conducted systematically
by offshore trawler fleets in the Pacific, Atlantic and Indian Oceans (Clark et al. 2007), which
may have been partially motivated by the declarations of the 200 nautical mile exclusive
economic zones (EEZs) around most nations’ productive coastal waters (Watson et al. 2007).
After finding large aggregations of fish and invertebrates, commercial fisheries developed in
a number of regions, with many countries pursuing fisheries on seamounts (Clark et al. 2007,
Morato & Clark 2007, Consalvey et al. 2010). These were aided by significant technological
advancements in the 1980s and 1990s, especially with regards to navigation (Brewin et al.
2007, Clark & Koslow 2007, Clark et al. 2010), which was important when fishing the
rugged terrain of the seamount habitat (Batson 2003).
Many of these fisheries have not been sustainable, with a number of them exhibiting a boomand-bust pattern (Clark et al. 2007). Fulton et al. (2007) notes that fish populations of certain
seamounts are often exhausted within five to ten years of exploitation, and probably take
decades to recover. The vulnerability of these fish populations is often due to their life history
and ecological characteristics (Morato & Clark 2007). The species concerned are often longlived, have a late age at maturity, low fecundity and sporadic reproduction (Clark 2001,
Brewin et al. 2007, Morato et al. 2008). They are also concentrated in a relatively small area
and need large spawning aggregations for successful recruitment (Brewin et al. 2007),
enabling big catches and a quick depletion of stock size (Clark & Koslow 2007, Clark et al.
2010). Other negative effects include a reduction in genetic diversity, the removal of apex
predators via bycatch (Batson 2003) and the discharge of processing waste (Clark & Koslow
2007).
7
The benthic seamount habitat and its associated fauna are also very vulnerable to the effects
of fishing (Clark & Koslow 2007, Consalvey et al. 2010, Clark et al. 2015), especially with
regards to bottom trawling (Clark & Koslow 2007, Clark et al. 2007). Demersal fauna is
often dominated by large, slow-growing sessile animals (Batson 2003, Clark & Koslow
2007), which have a limited spatial extent, low larval output, possibly limited recruitment
between seamounts, and a very localised distribution (Samadi et al. 2007, Consalvey et al.
2010). When removed as bycatch (Batson 2003), the seamount may take decades to recover
(Consalvey et al. 2010, Clark et al. 2015). The benthic species composition, abundance, age
composition, size structure and overall structural complexity may also be impacted (Clark &
Koslow 2007). Indirect effects include sediment re-suspension and mixing (Batson 2003,
Clark & Koslow 2007). Finally, endemic species may be at an increased risk of extinction
(Samadi et al. 2007).
Other possible threats to the seamount habitat include the mining of cobalt-rich
ferromanganese crusts (Ramirez-Llodra et al. 2010), invasive organisms, pollution, rising
carbon dioxide levels (Guinotte et al. 2006 ) and climate change (Batson 2003).
Historically, seamounts have not been well protected (Fulton et al. 2007, Consalvey et al.
2010). Unregulated, extensive commercial fishing often occurs on the high seas (both in the
past and presently), with these areas falling outside of any nations jurisdiction (Consalvey et
al. 2010). When regulations are in place, enforcement on the high seas is also a challenge
(Consalvey et al. 2010). In addition, there has also been little obligation to collect
information that is important with regards to effective management (Fulton et al. 2007).
Other issues include incorrect reports on fishing activities (Clark et al. 2007), the inability to
relate catch statistics to a specific seamount (Watson et al. 2007) and a lack of scientific and
fisheries data (Clark et al. 2007), all of which are important for fisheries models (Brewin et
al. 2007).
8
A major factor which hinders the successful management of this deep-sea habitat is the sparse
nature of data at both national and international levels (Davies et al. 2007, Clark et al. 2011).
For example, the impact of fishery-based disruption on seamount communities is difficult to
measure, with little known of their recovery process (Clark & Koslow 2007, Consalvey et al.
2010, Clark et al. 2015). This places pressure on scientists to obtain information in order to
suggest appropriate management plans (Clark et al. 2010).
Future research
Since the time that the exploitation of seamounts began, the field of seamount biology has
grown, especially in recent decades (Brewin et al. 2007). Despite this, we still know little
about these deep-sea habitats and the communities they contain. The fauna of these habitats
are poorly documented (Samadi et al. 2007, Consalvey et al. 2010) and the structure of whole
assemblages is only known from relatively few seamounts worldwide (Samadi et al. 2007).
The lack of taxonomic expertise, slow description rates of new species and varied sampling
methods also limit what can be done with the sparse data that are available (Samadi et al.
2007, Ramirez-Llodra et al. 2010). Further bias has also arisen due the greater sampling of
larger fauna (Clark et al. 2010). Moreover, certain seamount types and locations are
understudied, such as deep seamounts, and those in the equatorial regions or at high latitudes
(Clark et al. 2010).
Seamounts may be one of the last major frontiers of exploration on Earth, especially from a
geographic, ecological and geological point of view (Wessel et al. 2010). The purpose of
seamount conservation and management has ignited a new era of multidisciplinary research
and international collaboration (Brewin et al. 2007, Kvile et al. 2014). Examples of this
include the Global Census of Marine Life on Seamounts (CenSeam) (Stocks et al. 2012) and
9
the Seamount Ecosystem Evaluation Framework (SEEF) (Kvile et al. 2014), which have
elevated the seamount habitat in the public eye (Consalvey et al. 2010). The current
knowledge of seamounts would be enhanced by the standardisation of sample collection and
data sharing (Consalvey et al. 2010), as would future research in understudied regions, such
as the Indian Ocean (Clark et al. 2010, Consalvey et al. 2010).
1.2 Seamounts of the Indian Ocean
An intermediate number of seamounts occur in the Indian Ocean (Ingole & Koslow 2005),
with these deep-sea habitats being the most poorly explored of this region (Wafar et al.
2011). Overall, Sautya et al. (2011) suggests that 15 seamounts have been investigated
biologically in this Ocean, but only four of these (Equator Seamount, Fred Seamount, Mount
Error Guyot and Walters Shoal Seamount) have well documented benthos, and only single
records are known from the others. Thus, the fauna of seamounts remain effectively unknown
in the Indian Ocean (Rogers et al. 2009, Sautya et al. 2011, Kvile et al. 2014), with the
number of species recorded from these features currently less than 300 (Wafar et al. 2011).
This limited understanding of biodiversity, both generally in this ocean, and of the seamounts
it contains, can be attributed to the lack of funding and capabilities (human, technical and
institutional) in its surrounding countries (Wafar et al. 2011).
Marine research in the Indian Ocean is intertwined with its colonial past, with most work to
date having been done by European scientists (Wafar et al. 2011). Extensive sampling was
carried out during the International Indian Ocean Expedition (Rogers et al. 2009), but the
main source of information regarding seamount biology has been scientific and/or fisheries
reports of past Soviet and French expeditions, which focused predominantly on ichthyofauna
and plankton according to their interest in seamount fisheries (Romanov 2003, Ingole &
10
Koslow 2005, Rogers et al. 2009, Letessier et al. 2015). In addition, work on the seamounts
of this region often remains in an unpublished state, in grey literature and/or is often
unavailable in English, making it difficult to find and access (Kvile et al. 2014).
Walters Shoal: a shallow seamount in the Western Indian Ocean region
Compared to other seamounts in the Indian Ocean, quite a few studies have been carried out
on Walters Shoal. This shallow seamount is located on the South Madagascar Ridge at
33°13'S, 43°51'E and lies approximately 400 nautical miles south of Madagascar and 600
nautical miles east of South Africa (Fig. 1). During the Pleistocene (and possibly the Tertiary)
period, Walters Shoal was exposed to subaerial erosion (Schlich et al. 1974). Today, this
seamount forms part of a benthic protected area voluntarily closed to trawl fishing by the
Southern Indian Ocean Deep-sea Fishers Association (SIODFA) (Shotton 2006, Rogers et al.
2009, Letessier et al. 2015).
Rogers et al. (2009) attributes the past and present interest in this seamount to its close
proximity to land and to the commercial fishery focus in this region. Its accessibility could
also play a role, with the shallow seamount lying approximately 15 m below the sea surface
(Rogers 2012, Pollard & Read 2015, Fig. 2). Accordingly, this atypically domed structure
penetrates the euphotic zone, enabling its shallowest depths to be covered in rhodolithforming coralline encrusting algae (Kensley 1969, Collette & Parin 1991) and coral
(Romanov 2003).
Walters Shoal was sampled in 1964 during the International Ocean Expedition by the RV
Anton Bruun, giving rise to the discovery of several invertebrates. Clark (1972) described a
new endemic subspecies of crinoid, Comanthus wahlbergi tenuibrachia (currently
11
Comanthus wahlbergi), while Kensley (1975) noted a new endemic isopod, Jaeropsis
waltervadi. An endemic species of alpheid shrimp, Alpheus waltervadi, was also discovered
on the shoal, and the presence of four other decapods was recorded (Kensley 1969, Kensley
1981). The coral Enallopsammia amphelioides was collected (in addition to a few fish) in
1976 using the French vessel, Marion Dufresne (Zibrowius 1982), while the search for
fishery resources by both French and Soviet vessels led to the finding of many fish (and some
crustacean) species (Collette & Parin 1991, Romanov 2003, Rogers et al. 2009). The 17th
cruise of the Soviet oceanographic vessel, Vityaz in 1988 – 1989 provided more details on the
ichthyofauna inhabiting Walters Shoal. Collette & Parin (1991) recorded 20 fish species
obtained down to approximately 400 m, while 52 cephalopod species were collected on, over
and around the seamount (Nesis 1994). A few new endemic fish species were also discovered
(Poss & Collette 1990, Collette et al. 1991, Iwamoto et al. 2004), while work regarding the
brachiopods of Walters Shoal has also arisen based on a few collections during this cruise
(Zezina 2010). In addition, macroplankton collected was included in the work by
Vereshchaka (1995), which was a comprehensive summary of several investigations
regarding macroplankton found in the near-bottom layer of seamounts and slopes in the
Indian Ocean. Studies on the distribution patterns of Walters Shoal benthic and water-column
fauna were carried out by the P.P. Shirshov Institute of Oceanology in the 1980s (T. N.
Molodtsova, personal communication, September 2, 2015). However, these works including
Parin et al. (1993) and Detinova & Sagaidachny (1994) are largely inaccessible. These data
may be available on OBIS (Ocean Biogeographic Information System, available:
www.iobis.org/mapper/) according to T.N. Molodtsova (personal communication, September
2, 2015), which may account for the 288 taxa recorded from Walters Shoal Seamount by
Sautya et al. (2011).
12
More recently, a commercial fishing trip aboard the Spanish fishing vessel Iannis, led to the
discovery of a new species of spiny lobster, Palinurus barbarae (Groeneveld et al. 2006). In
2009, the RV Dr Fridtjof Nansen undertook a cruise aimed at understanding the pelagic
biology and physical oceanographic setting of the seamounts on the Southwest Indian Ocean
Ridge, including a sampling location on or near Walters Shoal Seamount (see Rogers et al.
2009). Studies from the data and samples collected have led to recent publications on
physical oceanography (Read & Pollard 2015), circulation (Pollard & Read 2015), the
distribution of micronektonic crustaceans (Letessier et al. 2015) and cephalopod diversity
(Laptikhovsky et al. 2015). Rogers et al. (2009) also noted the presence of marine mammals,
including sperm whales, humpback whales and short-finned whales. Blue whales and fin
whales were also possibly observed. These accounts support sightings of humpback whales
by Collette & Parin (1991) and Shotton (2006), suggesting that Walters Shoal may be an
important migratory area between feeding and breeding grounds (Shotton 2006). In addition,
tracking data have revealed that Walters Shoal is an important foraging ground for both the
red-tailed tropicbird and Barau’s petrel (Le Corre et al. 2012), probably due to upwelling and
local enrichments.
The previous work done on the fish (Collette & Parin 1991, Iwamoto et al. 2004) and
cephalopod (Nesis 1994) fauna led to the current understanding of the biogeographical
affiliations of Walters Shoal Seamount. Collette & Parin (1991) found the shallow-water fish
fauna to be composed of three elements, including endemics (to the West Wind Drift or
Indian Ocean islands and seamounts within the region, or just to Walters Shoal; six to seven
species), widespread temperate or subtropical species (six to seven) and tropical Indo-West
Pacific reef species (six). No Antarctic and Subantarctic species were found and there was
little similarity to the fishes of South Africa (Collette & Parin 1991). These authors suggest
that the fish fauna of Walters Shoal link the Tristan-Gough Province (Southern South
13
America Cold Temperate Region) with the Amsterdam-St. Paul Province (Southern African
Warm Temperate Region) as defined by Briggs (1974) into a single biogeographic province,
which they named the West Wind Drift Islands Province (WWDIP). This province includes
Tristan da Cunha, Gough Island, Vema Seamount, Walters Shoal, UN-2 (unnamed seamount
south of Madagascar) and the St Paul and Amsterdam islands (Nesis 2003). Most of this
chain lies along the edge of the West Wind Drift (WWD), which is an eastward-flowing
Subantarctic surface current, with a northern boundary defined by the Subtropical
Convergence Zone (Iwamoto et al. 2004). Similar findings were documented with the
cephalopod fauna (Nesis 2003), with Iwamoto et al. (2004) suggesting that the fish fauna
found at Walters Shoal can be explained by its location within the northern oscillatory region
of the WWD, thus comprising both subtropical and Subantarctic elements, as seen with the
grenadier fauna.
The work by Parin et al. (1993) (as cited by Iwamoto et al. 2004) included both shallowwater fish and invertebrates. These authors suggest that the source faunas for Walters Shoal
were the tropical Western Indian Ocean, southernmost South Africa and islands of the WWD.
On the other hand, they found that subtropical, antitropical and southern peripheral species
dominated on the continental slope and midwaters.
Although Walters Shoal has been relatively well sampled, there is still a paucity of available
biodiversity and oceanographic data. Thus, a multidisciplinary cruise was launched in May to
June 2014 as a component of the third phase of the African Coelacanth Ecosystem
Programme (ACEP III). Sponsored by the National Research Foundation (NRF) and
supported by the Department of Environmental Affairs (DEA) Oceans and Coasts Branch,
participants included members of the Department of Agriculture Forestry and Fisheries
(DAFF), the South African Environmental Observation Network (SAEON), DEA and
students from both Rhodes University (RU) and the University of the Western Cape (UWC).
14
As one of the few South African expeditions to explore this unique feature, the aim of the
cruise was to gain detailed information on the benthic fauna (invertebrates and fish)
associated with the photic and subphotic zone, while also collecting information on the
physical and chemical environment. Combined, this data would provide a better
understanding of the Walters Shoal ecosystem.
This thesis falls under the above-mentioned larger project, with the aim to investigate the
diversity and distribution of the sponge fauna from Walters Shoal, while also assessing the
possible connectivity between this shallow seamount and adjacent regions. The four main
objectives of this study are as follows:
I) To sample and identify the sponges collected from Walters Shoal Seamount.
II) To describe a subset of the sponges collected from this seamount in order to illustrate
some of the potentially new species found in this study.
III) To determine whether sponge assemblages differ according to location (western vs.
eastern flank) and depth (shallow, mesophotic, submesophotic) on the seamount.
IV) To further investigate the biogeographical affiliations of Walters Shoal Seamount,
especially within the larger Western Indian Ocean and West Wind Drift context.
1.3 Why sponges?
Sponges (phylum Porifera), are considered to be amongst the first and simplest metazoans
(Batson 2003, Pechenik 2009) and although they lack the complexity observed in other
animal taxa, they comprise a highly successful and variable group (Marshall 1979, Gage &
Tyler 1991). Found in a range of environmental conditions, 98% of sponge species are
15
marine (Pechenik 2009) and inhabit all depths (Galstoff 1960, Bell & Carballo 2008, van
Soest et al. 2012).
In addition to their ubiquitous nature, sponges act as prominent, ecologically significant and
competitive components of marine benthic communities (Branch & Branch 1981, Barnes &
Bell 2002, Samaai 2006, van Soest 2007, Pechenik 2009). These organisms may serve as a
food source for demersal grazers and other predators (Kelly-Borges 1997, van Soest 2007), as
well as acting as a biological habitat and/or hosts for associated (sometimes symbiotic)
species (Jones & Gates 2010) including fish, macrofauna and microbes (Galstoff 1960,
Batson 2003, Schuchert & Reiswig 2006, van Soest 2007, Pechenik 2009, van Soest et al.
2012). Some symbiotic microbes may play a part in the nitrogen cycle and possibly
contribute organic production in nutrient impoverished environments (van Soest et al. 2012),
while their hosts (as active suspension feeders) enable benthic-pelagic coupling (van Soest
2007, van Soest et al. 2012). Furthermore, sponges may act as bio-eroders (Kelly-Borges
1997, Holmes 2000, van Soest 2007, van Soest et al. 2012) and environmental quality
indicators (Diaz & Rützler 2009).
From an anthropogenic point of view, sponges played an important role in ancient society,
and continue to do so today. In the past, sponges were used as household items, for personal
hygiene, for the relief of pain, for treating disease and in art (van Soest 2007, Voultsiadou
2007, van Soest et al. 2012). More recently, interest in sponges has largely arisen due to their,
and/or their symbionts, production of novel chemical compounds, which may have potential
biomedical and anti-fouling applications (Batson 2003, van Soest 2007, Pechenik 2009). In
addition, the silica structures made by sponges (spicules) have instigated further interest due
to their unique optical and mechanical properties, which may enable the manufacture of
advanced materials (Sundar et al. 2003, Weaver et al. 2003). Finally, further study into
16
sponges may lead to a greater understanding of life on Earth in an evolutionary context (van
Soest et al. 2012).
Global sponge diversity
According to van Soest et al. (2012), approximately 8 500 valid sponge species are known,
with most of these (around 80%) belonging to the class Demospongiae. However, our
knowledge of sponge diversity is incomplete and at least double this number of species is
thought likely to exist (van Soest 2007, van Soest et al. 2012). Although global patterns in
sponge species diversity remain rudimentary (van Soest 1994, Wörheide et al. 2005), recent
work by van Soest et al. (2012) suggests that these diversity patterns are similar to those
recognised in other marine animal groups, i.e. more species in tropical regions, and fewer in
colder areas of the global ocean. Yet, this pattern only emerges when looking at the most
elevated spatial marine realms (as defined by Spalding et al. (2007)) or highest taxonomic
ranks (Gage & Tyler 1991, van Soest 1994, Barnes & Bell 2002). At all spatial and
taxonomic levels, sponge diversity data demonstrate a strong bias according to collection and
taxonomy efforts (van Soest 1994, Barnes & Bell 2002, van Soest et al. 2012). The majority
of sponges occur in regional or local areas of endemism, mainly because of the limited
swimming capabilities of their larvae, asexual reproduction (van Soest et al. 2012), and
environmental variables including light and turbidity (Wörheide et al. 2005). Thus, van Soest
et al. (2012) suggest that a regional approach may currently provide more insight into the
biogeographic history of sponges.
Regional expeditions and work on sponge biodiversity has increased over the past two
decades (van Soest et al. 2012). As a result, many outputs including regional sponge guides,
databases, inventories, websites and CD’s have been realised (van Soest 2007, van Soest et
17
al. 2012). Other online databases focus on the natural compounds and symbionts of sponges,
as well as barcoding and DNA-based identification (van Soest 2007, van Soest et al. 2012).
The most internationally significant advancements include a comprehensive, multi-author,
guide to the identification of sponges (the Systema Porifera), edited by Hooper & van Soest
(2002), and the subsequent, regularly updated, searchable online database (the World Porifera
Database, van Soest et al. 2015).
To date, much work still needs to be done, with more scientific focus placed on economically
important species including molluscs, fish and crustaceans (Samaai, 2006, Costello et al.
2010). This also may be partly due to difficulties in sponge identification, associated with
morphological plasticity, and a shortage in the relevant taxonomic capacity (Branch &
Branch 1981, Kelly-Borges 1997, Barnes & Bell 2002, Batson 2003, Samaai 2006, Costello
et al. 2010, Jones & Gates 2010). Other factors that hamper our knowledge of global sponge
diversity include the often dated, scattered (and sometimes inaccessible) nature of the
taxonomic literature, the lag between documenting, describing and distributing information
on collected specimens, the numerous specimens awaiting description in museums
worldwide, as well as the neglect of certain taxa (e.g. Calcarea) (Wörheide et al. 2005, van
Soest et al. 2012). The lack of unsubstantiated and unpublished presence records, as well as
collection effort, also plays a role, with many regions and habitats remaining largely
undersampled (Wörheide et al. 2005, van Soest 2007, Costello et al. 2010, van Soest et al.
2012).
Seamount-inhabiting sponges
Globally, very little is known about seamount sponges (Vieira et al. 2010), with studies
predominantly documenting sponge fauna diversity and/or describing new species (e.g.
18
Vieira et al. 2010, Cristobo et al. 2015, Kelly et al. 2015). Even less is known of seamountinhabiting sponges in the Indian Ocean, with Sautya et al. (2011) suggesting that, prior to
their study, there were only reports on ‘Porifera’ and ‘Hexactinellida’ from two Indian Ocean
seamounts each in the literature.
Relatively comprehensive studies were carried out by Lévi (1969) on Vema Seamount
(South-East Atlantic), who recorded 28 species (15 new, 53% endemic), SchlacherHoenlinger et al. (2005) who documented 16 (seven new) ‘lithistid’ sponges from South
Pacific seamounts, with the fauna dominated by ‘spot endemics’ (species restricted to a single
site) and the work done by Xavier & van Soest (2007). The latter authors assessed the
diversity and biogeographical affiliations of the demosponge fauna of Gettysburg and
Ormonde Seamounts on the Gorringe Bank (North-East Atlantic), finding 23 species, with 36
species recorded overall. This study also documented range extensions, a moderate faunal
similarity (around 50% shared species) with adjacent locations and demosponge distribution
patterns consistent with those observed for the mollusc and fish fauna of these seamounts. In
contrast to these faunas, the sponge assemblage demonstrated a relatively high level of
endemism (28%).
As documented in other sessile benthic assemblages on seamounts (e.g. Bo et al. 2011,
Sautya et al. 2011, Thresher et al. 2014, McClain & Lundsten 2015), the sponge fauna
inhabiting these features often demonstrates significant differences (e.g. diversity,
abundance) with position on the seamount and depth, often according to local
geomorphology and hydrodynamic conditions (Bo et al. 2011). Examples include studies by
Henrich et al. (1992, Vesterisbanken Seamount), Pereira et al. (2015, Condor Seamount) and
Xavier et al. (2015, Schultz Seamount).
19
The current state of knowledge of seamount-inhabiting sponges indicates a diverse fauna that
is highly endemic, with existing estimates conservative, as many sponge collections have yet
to be sorted and identified (Schlacher-Hoenlinger et al. 2005, Vieira et al. 2010).
To date, this thesis constitutes the only study dedicated exclusively to the diversity,
distribution and biogeographical affiliations of the sponge fauna, not only of Walters Shoal
Seamount, but also possibly from the seamount habitat in the Indian Ocean.
1.4 Hypotheses
Based on the previous research carried out on Walters Shoal Seamount, as well as ‘accepted’
or general principles (i.e. individual seamounts may show great variability) from other
seamount studies worldwide, several hypotheses can be proposed regarding the sponge fauna
of Walters Shoal:
I) The sponge fauna will be diverse as found in previous studies on seamount-inhabiting
sponges (Lévi 1969, Schlacher-Hoenlinger et al. 2005, Xavier & van Soest 2007). These
studies report assemblages with less than 40 species as diverse.
II) Range extensions and sponge species new to science will be discovered due to the
undersampled and underworked state of the sponge fauna, not only of Walters Shoal, or of
seamounts in the Indian Ocean (Sautya et al. 2011), but also of the Western Indian Ocean
region in general (Kelly-Borges 1997, Richmond 2001).
III) Sponge assemblages will not demonstrate a significant difference according to
location (western vs. eastern flank) on the seamount, due to its small size and the retentive
nature of the waters above it (Nesis 1994, Gopal 2007).
20
IV) Sponge assemblages will demonstrate a significant difference according to depth
(shallow, mesophotic, submesophotic) on the seamount. Seamount benthic communities
are often determined by this factor, according to associated environmental gradients,
including temperature and oxygen concentration (Stocks & Hart 2007, Clark et al. 2010,
Consalvey et al. 2010). Previous works on seamount sessile benthic assemblages have noted
such a difference (e.g. Bo et al. 2011, Sautya et al. 2011, Thresher et al. 2014, McClain &
Lundsten 2015), including those on sponge fauna (Henrich et al. 1992, Pereira et al. 2015,
Xavier et al. 2015).
V) Sponge faunal affinities will be with surrounding regions including the tropical
Western Indian Ocean, southernmost South Africa and the West Wind Drift Islands
Province, as was found for the fish fauna by Collette & Parin (1991), the cephalopod fauna
by Nesis (2003), as well as both the fish and invertebrate fauna by Parin et al. (1993).
21
Chapter 2 – Methodology
2.1 Collection
Sponges were collected from Walters Shoal Seamount during a single cruise aboard the RV
Algoa from 15 May to 13 June 2014 (cruise number 208). Collections were carried out in a
random-stratified regime, following Clark et al. (2004), using SCUBA and a roughed
epibenthic sled, from the peak and down two opposing slopes (west and east) of the seamount
(Table 1, Fig. 3, Fig. 4). Nine sled transects were undertaken, three in each depth strata
(following Lesser et al. (2009)), including shallow water (15 – 30 m), the mesophotic zone
(31 – 150 m) and the submesophotic zone (>150 m). Two SCUBA dives were carried out in
shallow water (29 m), while eight sponge specimens found in a lobster trap (39 m) deployed
on the trip, were also included.
Once collected, specimens were labelled and frozen, to retain colour following Hooper
(2003), for processing onshore.
2.2 Taxonomic procedure
In the laboratory, the macroscopical features of each specimen were described (Fig. 5), with
the aid of Boury-Esnault & Rützler (1997). Subsequently, a TS number (personalised number
for collection of Toufiek Samaai) was assigned and digital colour photographs were taken of
each specimen before being preserved in 96% ethanol.
22
Spicules
For the study of spicules by light and scanning electron microscopy (SEM), a small section
(~3 mm3) of tissue (including both ectosome and choanosomal regions) was placed in a test
tube with a few drops of nitric acid. Once the tissue had digested, the spicules were rinsed in
distilled water (centrifuged for 1 min at 3000 rpm) three times consecutively, twice with
distilled water and once with 96% ethanol. Spicule samples were then stored at room
temperature in 96% ethanol.
For light microscopy, spicule extracts were re-suspended and pipetted onto microscope
slides, and air-dried at 40⁰C. Subsequently, the mounting medium Entellan was added to the
slides, followed by cover slips. These slides were then allowed to air dry at room temperature
until the mountant had hardened. A Carl Zeiss AxioCam ERc 5s camera (mounted on a
compound microscope) and ZEN 2012 software were used to measure ten spicules from each
spicule category (per specimen). These dimensions are given as mean length (range) x mean
width (range) followed by the number of spicule measurements taken (n). Spicule dimensions
from other specimens obtained in this study, and from the literature where possible, are
included to determine the level of intraspecific variability (Samaai & Gibbons 2005).
For SEM, spicule extracts were placed on film negative fixed to aluminium studs with
superglue. Once the ethanol had evaporated, the studs were sputter-coated with goldpalladium and images taken using a FEI Nova NanoSEM 230 equipped with a field emission
gun and digital imaging software programme. Such microscopy was necessary to perceive
small but important spicule variations that confer specific identity (Hooper 1996).
23
Skeletal arrangement
A perpendicular section of tissue (~5 mm3), including both ectosome and choanosomal
regions, was collected from each specimen (where possible) and stored in 96% ethanol in
order to document the skeletal structure and spicule arrangement. After the sample had been
processed through a series of dehydrating and cleaning agents (Table 2), it was embedded in
paraffin wax. Using a microtome, a section of ~30 – 90 μm was cut from the embedded
sample and the wax removed via washing in Xylene (in a fume cupboard). After being
mounted on a slide with Entellan, the skeletal arrangement was photographed using the
equipment and software previously mentioned. Alternatively, where the above was not
possible or did not reveal certain structures, a perpendicular section of tissue was coated with
nitric acid and heated at 40⁰C to remove tissue.
Digital images were combined on a black background, aligned and cleaned (when
appropriate) using PowerPoint and Photoshop CS5.
Identification
Identifications to the lowest operational taxonomic unit (OTU) were possible through the
consideration of the macroscopic features, spicule array and skeletal arrangement in
conjunction with the consultation of the taxonomic literature. The work by Hooper & van
Soest (2002) and the World Porifera Database (van Soest et al. 2015) was especially useful
with regards to identifying specimens to genus level and documenting updated classifications
respectively. The majority of specimens were identified to the genus level and compared to
documented species within the region of interest (Table 3, Table A in appendix). When
specimens were found to differ from these species, or represent the first record of a genus in
24
the region of interest, they were denoted as sp. • and likely constitute species new to science,
which will be subsequently described for publication. Several specimens could only be
identified to higher taxonomic levels (i.e. order, family or tentative genus). These were
denoted as sp. and require further investigation. Finally, specimens that lacked enough
diagnostic material for identification, but were morphologically distinct, were designated as
morphospecies (M).
2.3 Sample storage
Samples of all material will eventually be housed in the Natural History collection of the
South African Iziko Museum in Cape Town, and accession numbers will be provided by this
institution once deposited. Voucher samples will be kept in the private collection of Dr
Toufiek Samaai presently of the Department of Environmental Affairs, Oceans and Coasts
Branch.
2.4 Location and depth analyses
To determine whether the sponge fauna of Walters Shoal Seamount is associated with
location (western vs. eastern flank) and depth (shallow, mesophotic, submesophotic; as
defined above), Bray-Curtis coefficients based on a presence/absence (non-detection) matrix
of the OTU’s found at each sampling location (Table 4) were calculated using PRIMER
v.6.1.11 (Plymouth Routines in Multivariate Ecological Research; Clarke & Gorley 2006).
The two-way crossed analysis of similarity (ANOSIM) and non-metric multidimensional
scaling (nMDS) ordination routines (see O’Hara (2007)) were performed to assess and
visualise the sponge faunal similarities between sampling locations for both factors
25
respectively. ANOSIM is an approximate equivalent of the standard ANOVA (analysis of
variance), enabling a non-parametric test for statistically significant differences in the sponge
assemblage composition between sample groups specified by the location and depth factor
levels (Clarke & Gorley 2006), with the significance of this statistical test assigned here at the
5% level. SIMPER (similarity percentage analysis) is an exploratory analysis which indicates
the species principally responsible for differences between sets of samples (Clarke & Gorley
2006) and was thus used to assess the extent of similarity both within and between the
location and depth factors, while also identifying the species contributing to the observed
(dis)similarity.
2.5 Biogeography analyses
To comment on the biogeographical affiliations of the Walters Shoal Seamount sponge fauna,
it was compared to that of the surrounding regions pertinent to the hypotheses proposed
(Table 3, Fig. 6, Table A in appendix). Species lists were extracted for these regions, from the
World Porifera Database (van Soest et al. 2015), according to the MEOW (Marine
Ecoregions of the World) biogeographical classification scheme as defined by Spalding et al.
(2007). The similarity between these regions and Walters Shoal was assessed by calculating
the ratio of shared species (known sponge species documented from this study) between each
region and Walters Shoal Seamount and the total number of species recorded from Walters
Shoal, following Xavier & van Soest (2007). Extracted lists were also used to determine the
contribution of each family and genus to the sponge fauna of regions found to have
biogeographical affiliations with Walters Shoal, for comparison with all OTU’s recorded in
this study.
26
Chapter 3 – Results
3.1 Systematics
A total of 255 sponge specimens were collected from Walters Shoal Seamount, comprising
78 operational taxonomic units (OTU’s) (Table 4, Table 5). There were representatives of six
subgenera, 40 genera, three subfamilies, 28 families, one suborder, 14 orders, four subclasses
and two classes. Twenty-three species (29.5%) are known and could be included in the
biogeographical analyses. Twenty-six species (33.3%) were compared to species of the same
genera within the region of interest (Table 3, Table A in appendix) and were found to differ,
or represent the first record of a genus in the region of interest, and thus likely represent
species new to science. Ten (12.8%), four (5.1%) and two (2.6%) species could only be
identified to order, family and tentative genus level respectively and therefore require further
investigation. Finally, 13 species (16.7%) could only be designated as morphospecies due to a
lack of diagnostic material, but could still be included in location and depth analyses.
The dominant group was the class1 Demospongiae, which comprised 63 species (80.8%)
overall. Within this group, the subclass Heteroscleromorpha was well represented,
comprising 59 species (75.6%), while the subclasses Keratosa and Verongimorpha comprised
two species (2.6%) each. The class Calcarea was represented by two species (2.6%), both
within the subclass Calcinea.
The orders Tetractinellida (13 species), Poecilosclerida (11 species) and Suberitida (11
species) were most speciose and together accounted for 44.9% of all species. The orders
1
Higher taxa names follow the revised classification proposed by Morrow & Cárdenas (2015) and
recognised by the World Porifera Database (van Soest et al. 2015).
27
Axinellida and Haplosclerida were also relatively well represented, with eight and seven
species documented respectively. Three species obtained were from the order Bubarida,
while the orders Agelasida, Biemnida, Clathrinida and Tethyida comprised two species each.
Finally, one species was obtained for each of the orders Chondrosiida, Dendroceratida,
Dictyoceratida and Verongiida.
The majority of species represent a single genus each. However, four species were designated
to Stelletta, while three species each were designated to the genera Phakellia and
Protosuberites. Two species each were designated to the genera Amorphinopsis,
Callyspongia, Eurypon and Tedania.
3.2 Descriptions
For the purposes of this thesis, the taxonomic descriptions of only 13 Demospongiae species
from Walters Shoal Seamount are given below. Of these, four are re-described from fresh
material and nine are described as new (denoted as sp. •).
Phylum Porifera Grant, 1836
Class Demospongiae Sollas, 1885
Subclass Heteroscleromorpha Cárdenas, Perez & Boury-Esnault, 2012
Order Agelasida Hartman, 1980
Family Agelasidae Verrill, 1907
Genus Agelas Duchassaing & Michelotti, 1864
28
Agelas ceylonica Dendy, 1905 (Fig. 7 A – F, Table 6)
Synonymy
None.
Material examined. TS 2309 (WSL-INV47), TS 2313 (WSL-INV48), TS 2317
(WSL-INV46(2)): Walters Shoal Seamount, Grid WSL022, Station ALG10954, collected via
sled (no 2) by the RV Algoa, (33°10.9' S; 43°48.6' E) - (33°11.2' S; 43°50.2' E), duration 41
min, depth 170 – 72 m, 29 May 2014. TS 2441 (WSL-INV74(7)), TS 2443 (WSL-INV74(9)),
TS 2452 (WSL-INV74(18)), TS 2455 (WSL-INV74(21)), TS 2456 (WSL-INV74(22)), TS
2549 (WSL-INV74(31)): Walters Shoal Seamount, Grid WSL024, Station ALG10956,
collected via sled (no 3) by the RV Algoa, (33°08.8' S; 43°49.1' E) - (33°09.0' S; 43°50.5' E),
duration 33 min, depth 348 – 103 m, 29 May 2014.
Description. Repent-ramose form, which binds together with biogenic debris,
creating a conglomerate (not shown). Length 6.5 cm, diameter 1.3 cm and thickness 0.7 cm.
Surface rough and fuzzy, with small, randomly scattered oscules (round apertures), ranging
from <1 mm – 1 mm in diameter. Consistency and texture is soft and spongy, compressible
and not easily torn. Colour in situ brownish orange, pale orange in preservative.
Skeleton. Choanosomal skeleton comprises an isotropic reticulation consisting of a
uniform network of spongin fibres, echinated by verticillate acanthostyles, with blunt ends
embedded in the fibre. These fibres are also rarely cored with verticillate acanthostyles
(embedded lengthwise in fibre). Ascending fibres usually echinated while transverse fibres
not, but this is not always the case. Interconnecting transverse fibres are often uncored, and
29
form irregular meshes of 30 – 100 µm in diameter. Spongin is sparsely scattered through the
mesohyl. No ectosomal specialisation.
Spiculation. Megascleres. Verticillate acanthostyles in two size classes: I) 191.5
(163.9 – 216.5) x 9.0 (6.3 – 11.0) µm, n = 10, with 15 – 22 whorls of spines; II) 115.6 (89.6 –
148.0) x 4.3 (3.1 – 5.5) µm, n = 10, with 12 – 17 whorls of spines. Microscleres. Absent.
Substratum, Depth range and Ecology. Nine specimens found in two sleds, one
which was dominated by hard live rock with many bivalves and sponges, the other host to
predominantly dead shells and hydrozoans. Depth range: 72 – 348 m.
Geographic Distribution. Found extensively throughout the Indian Ocean, including
Walters Shoal Seamount.
Remarks. The present material conforms well to Agelas ceylonica, which was
described by Dendy (1905) from the Gulf of Mannar as consisting of ‘a few slender,
anastomosing, sub-cylindrical branches, arising from an irregular, proliferous basal crust
attached to a calcareous nodule’. Dendy (1905) describe this species as having verticillate
spined styles of approximately 240 x 20 µm, while Lévi (1961) describe specimens from the
Seychelles as having two classes of ‘acanthostyles’ (I) 80 – 275 x 5 – 15 µm, with 16 – 21
whorls; II) 100 – 300 x 6 – 15 µm, with 13 – 23 whorls). The skeletal structure of Agelas
ceylonica is also consistent with the material here, having a fibre network echinated by
verticillate acanthostyles, with these also found occasionally embedded lengthwise in the
fibre.
30
The descriptions of the other species of this genus found in the region of interest
(Table 3): Agelas bispiculata Vacelet, Vasseur & Lévi,1976 (Verticillate acanthostyles: I)
320 – 400 x 14 – 17 µm, with 20 whorls; II) 55 – 120 x 6 – 10 µm, with 11 – 15 whorls),
Agelas marmarica Lévi, 1958 (Verticillate acanthostyles: 230 x 10 µm, with 19 – 21 whorls)
and Agelas mauritiana (Carter, 1883) (Verticillate acanthostyles according to Lévi (1961):
150 – 160 x 8 – 12 µm, with 16 – 20 whorls) also seem to be quite similar, especially with
regards to morphology. However, the present material differs from the above due to the
presence of very distinct, elongated spines on the smaller verticillate acanthostyles of these
species, which also cover the head of the spicule, unlike those described here which have
reduced spines and smooth heads.
Order Axinellida Lévi, 1953
Family Axinellidae Carter, 1875
Genus Ptilocaulis Carter, 1883
Ptilocaulis sp. • (Fig. 8 A – E, Table 7)
Material examined. TS 2440 (WSL-INV74(6)), TS 2448 (WSL-INV74(14)), TS
2546 (WSL-INV74(28)), TS 2570 (WSL-INV74(52)): Walters Shoal Seamount, Grid
WSL024, Station ALG10956, collected via sled (no 3) by the RV Algoa, (33°08.8' S;
43°49.1' E) - (33°09.0' S; 43°50.5' E), duration 33 min, depth 348 – 103 m, 29 May 2014. TS
2458 (WSL-INV114(1)): Walters Shoal Seamount, Grid WSL047, Station ALG10979,
collected via sled (no 9) by the RV Algoa, (33°09.7' S; 43°58.4' E) - (33°09.8' S; 43°57.0' E),
duration 50 min, depth 512 – 317 m, 03 June 2014.
31
Description. Erect, dichotomously branching form, with few scopiform flattened
processes. Length 3.1 cm, diameter 3.2 cm and thickness 0.4 cm. Surface irregular and finely
hispid (due to protruding spicules) with small circular oscules (<1 mm) scattered throughout.
Spicules protruding <1 mm from the surface, thus fuzzy to the touch. Texture soft and
spongy, compressible and easily torn. Colour in situ off-white, white in preservative.
Skeleton. Choanosomal skeleton consists of a dense interwoven mass of sinuous
styles cored in fascicles. All axial spicules are disposed longitudinally in a plumose fashion.
Spicule tracts are sometimes definable for only a very short distance before becoming
obscured in the general mass. The peripheral region is short and not well formed. Peripheral
spicules arranged individually, or multiple spicules branch tangentially to the axis in a
plumoreticulated fashion and ascend to, and usually protrude through, the ectosome. Styles in
the axial and peripheral skeleton do not appear to be differentiated, but are irregularly
arranged. Specialized ectosomal skeleton absent.
Spiculation. Megascleres. Styles, smooth, bent to sinuous, variable and hastate to
somewhat blunt distally, no easily discernible size classes (continuous): 462.9 – 1332.8 x
18.8 (15.4 – 22.5) µm, n = 10. Microscleres. Absent.
Substratum, Depth range and Ecology. Five specimens found on rocky substrata in
two sleds, predominantly composed of biogenic rubble, hydrozoans and rhodoliths. Depth
range: 103 – 512 m.
Geographic Distribution. Walters Shoal Seamount.
32
Remarks. The present material conforms to Ptilocaulis Carter, 1883 as diagnosed by
the presence of a vaguely reticulated axial skeleton and extra-axial skeleton formed by
fibrofascicles cored with styles and ending in surface or scopiform processes which are
distinctive for this genus (Alvarez & Hooper 2002). There are 11 currently accepted species
of Ptilocaulis worldwide (van Soest et al. 2015), of which only one, Ptilocaulis spiculifer
(Lamarck, 1814), occurs in the region of interest (Table 3).
Originally described by Lamarck in 1814, P. spiculifer has been recorded from Kenya
by Pulitzer-Finali (1993). The latter author notes curved styles of one size class with faintly
tylote bases (specimen one: 260 – 340 x 11.5 – 16 µm; specimen two: 230 – 290 x 9 –
14 µm). This was consistent with measurements from Ridley (1884): 350 x 19 µm and Dendy
(1922): 300 x 12.3 µm. However, both Ridley (1884) and Dendy (1922) record the
megascleres as having broadly rounded bases which is more consistent with the present
material. Nonetheless, all re-descriptions of this species depict a much smaller spicule size
range than that found for the present material, which thus likely constitutes a new species.
Order Haplosclerida Topsent, 1928
Family Callyspongiidae de Laubenfels, 1936
Genus Callyspongia Duchassaing & Michelotti, 1864
Subgenus Callyspongia (Callyspongia) Duchassaing & Michelotti, 1864
Callyspongia (Callyspongia) sp. • (Fig. 9 A – E, Table 8)
Material examined. TS 2330 (WSL-INV94(1)), TS 2341 (WSL-INV94(13)), TS
2353 (WSL-INV94(25)): Walters Shoal Seamount, Grid WSL044, Station ALG10976,
collected via sled (no 6) by the RV Algoa, (33°14.0' S; 43°55.5' E) - (33°13.7' S; 43°55.6' E),
33
duration 9 min, depth 28 – 25 m, 02 June 2014. TS 2369 (WSL-INV75(10)), TS 2370 (WSLINV75(11)), TS 2371 (WSL-INV75(12)): Walters Shoal Seamount, collected via SCUBA
(dive 1) by the RV Algoa, duration 35 min, depth 29 m, 30 May 2014. TS 2382 (WSLINV83(2)): Walters Shoal Seamount, collected via SCUBA (dive 2) by the RV Algoa,
duration 36 min, depth 29 m, 30 May 2014. TS 2479 (WSL-INV84(8)): Walters Shoal
Seamount, Grid WSL042, Station ALG10974, collected via sled (no 4) by the RV Algoa,
(33°11.2' S; 43°51.0' E) - (33°11.2' S; 43°50.7' E), duration 10 min, depth 34 – 28 m, 02 June
2014. TS 2537 (WSL-INV102(3)): Walters Shoal Seamount, Grid WSL045, Station
ALG10977, collected via sled (no 7) by the RV Algoa, (33°13.8' S; 43°56.1' E) - (33°14.2' S;
43°55.9' E), duration 16 min, depth 80 m, 02 June 2014.
Description. Massive, predominantly ramose but tubular, growing from a common
base, from which upright clusters of finger-like projections arise, interconnected laterally.
Tubes usually coalesced for a greater or lesser distance, occasionally united along entire
length. Length 6.0 cm, diameter 9.1 cm and thickness 1.4 cm. Surface smooth and velvety to
the touch. Oscules (3 – 9 mm diameter) present at the apex of the tubes, which become
fibrous at the tips. Transparent membrane covering exterior. Texture soft and spongy,
compressible and easily torn. Colour in situ bright blue, turning beige with purple tips above
water. In preservative, pale yellow.
Skeleton. Choanosome with a regularly rectangular-meshed skeleton formed by
multispicular primary spongin fibres ~30 µm wide, and ~71 – 430 µm apart, interconnected
often perpendicularly by secondary unispicular fibres ~20 µm thick, forming meshes ~110 –
250 µm wide. Unispicular tertiary fibres sometimes present, ~25 µm thick, forming meshes
~80 µm thick, which interconnect secondary fibres perpendicularly. Specialised ectosomal
34
skeleton absent, but primary fibres project as short, compact tufts of spicules beyond the
exopinacoderm.
Spiculation. Megascleres. Oxeas, short, smooth, straight to slightly curved medially,
hastate: 62.4 (56.6 – 68.8) x 3.0 (2.3 – 4.2) µm, n = 10. Microscleres. Absent
Substratum, Depth range and Ecology. Nine specimens found on rocky substrata in
three sleds and both dives, often with crinoids as epifauna. Depth range: 25 – 80 m.
Geographic Distribution. Walters Shoal Seamount.
Remarks. The present material conforms to the genus Callyspongia (Callyspongia)
Duchassaing & Michelotti, 1864 as diagnosed by a single ectosomal non-hispid layer,
multispicular well-defined choanosomal fibres with a distinct spongin sheath, forming a
rectangular mesh without free spicules, and a smooth surface (Desqueyroux-Faúndez &
Valentine 2002).
The present material does not seem conspecific with the three species of this genus
that have been recorded from the region of interest (Table 3): Callyspongia (Callyspongia)
differentiata (Dendy, 1922), Callyspongia (Callyspongia) reticutis (Dendy, 1905) and
Callyspongia (Callyspongia) tubulosa sensu (Esper, 1797), the latter of which was redescribed by Samaai & Gibbons in 2005. Both C. (C.) differentiata and C. (C.) reticutis have
slightly larger spicule sizes (80 x 3 µm and 72 x 2.6 µm respectively) but also differ to the
present material with regards to skeletal structure, with the former having secondary fibres
devoid of spicules and the latter having multispicular secondary fibres. Callyspongia (C.)
tubulosa is most similar morphologically to the present material, but has larger oxeas (110 –
35
140 x 13 µm) with multispicular secondary fibres. Thus, the present material likely
constitutes a new species.
Order Poecilosclerida Topsent, 1928
Family Coelosphaeridae Dendy, 1922
Genus Lissodendoryx Topsent, 1892
Subgenus Lissodendoryx (Lissodendoryx) Topsent, 1892
Lissodendoryx (Lissodendoryx) pygmaea (Burton, 1931) (Fig. 10 A – I, Table 9)
Synonymy
Myxilla pygmaea Burton, 1931: p. 342, Pl. XXIII, Fig. 1.
Material examined. TS 2364 (WSL-INV75(5)), TS 2365 (WSL-INV75(6)), TS 2366
(WSL-INV75(7)), TS 2367 (WSL-INV75(8)), TS 2368 (WSL-INV75(9)): Walters Shoal
Seamount, collected via SCUBA (dive 1) by the RV Algoa, (33°11.2' S; 43°50.7' E), duration
35 min, depth 29 m, 30 May 2014.
Description. Massive, ridge-shaped form. Length 13.0 cm, diameter 7.5 cm and
thickness 2.6 cm. Surface smooth and uneven, or markedly coarse with a thin, transparent
membrane covering the entire exterior. Oscules (up to 5 mm in size) present, scattered
randomly on top of the ridge. Texture firm but spongy, compressible and not easily torn.
Colour in situ light red to orange externally, pale yellow internally. In preservative, beige.
Skeleton. Choanosomal skeleton comprises a fairly tight-meshed regular reneroid
reticulation, with primary fibres ~40 µm across, running obliquely to the surface, composed
36
of spongin and cored by groups of 1 – 3 smooth styles, diverging in a plumoreticulate manner
to ectosome. Secondary fibres ~20 – 30 µm across, enter the primary fibres at an angle, cored
by single styles. Primary and secondary fibres with spongin, without a distinct sheath.
Ectosomal skeleton comprises a distinct and continuous palisade of tylotes, perpendicular to
and penetrating the surface, sometimes forming radiating bouquets ~150 – 200 µm deep.
Microscleres scattered throughout.
Spiculation. Megascleres. Ectosomal tylotes, smooth, straight shafted, with
elongated, well-developed heads: 181.5 (164.4 – 196.4) x 4.5 (3.5 – 5.1) µm, n = 10. Styles
smooth, slightly curved with pronounced shaft and hastate end: 125.6 (116.8 – 137.5) x 6.0
(4.4 – 6.8) µm, n = 10. Microscleres. Sigmas, both C- and S-shaped: 28.6 (26.4 – 31.1) µm,
n = 10. Arcuate isochelae in two size classes: I) 23.8 (22.4 – 25.0) µm, n = 10; II) 13.2 (12.1
– 14.1) µm, n = 10.
Substratum, Depth range and Ecology. Five specimens found on rocky substrata
during a dive, with crinoids as epifauna. Depth: 29 m.
Geographic Distribution. KwaZulu-Natal (South Africa), Walters Shoal Seamount.
Remarks. The present material conforms to Lissodendoryx (Lissodendoryx) pygmaea
(Burton, 1931), originally described as Myxilla pygmaea (Tylotes: 150 x 6 µm; Styles: 105 x
4 µm; Sigmas: 21 – 27 µm; Chelae: I) 21 – 27 µm, II) 12 µm). However, the present material
has slightly longer megascleres. Burton (1931) noted that the erected species may be allied
with, or identical to, Lissodendoryx (Lissodendoryx) isodictyalis (Carter, 1882), but
disregarded this notion due to the latter species’ distribution. Lissodendoryx (L.) pygmaea
37
was also noted by Lévi (1963, 1969) to resemble the type specimen of Lissodendoryx
(Lissodendoryx) ternatensis (Thiele, 1903) from Ternate, and a specimen from Vema
Seamount (off South Africa) which he described as the latter species. Hofman & van Soest
(1995) suggest that L. (L.) pygmaea may possibly be a closely related, but separate, species to
L. (L.) ternatensis, with the latter species having two classes of sigmas (as does L. (L.)
isodictyalis). However, both Lévi (1963, 1969) and Samaai & Gibbons (2005) note only one
category of sigmas for L. (L.) ternatensis.
Although the descriptions of L. (L.) ternatensis by Lévi (1963, 1969) conform well to
the present material, it has been placed conservatively here under L. (L.) pygmaea based on
the taxonomic ambiguities and disjunct distribution of L. (L.) ternatensis. Obviously, the
species that comprise Lissodendoryx (Lissodendoryx) in South Africa, and farther afield, are
in need of further investigation.
Family Dendoricellidae Hentschel, 1923
Genus Fibulia Carter, 1886
Fibulia ectofibrosa (Lévi, 1963) (Fig. 11 A – E, Table 10)
Synonymy
Desmacidon ectofibrosa Lévi, 1963: p. 26, Fig. 27, Pl. IV A, B.
Fibula ectyofibrosa Samaai & Gibbons, 2005: p. 57, Figs. 4G, 41 A – C.
Material examined. TS 2303 (WSL-INV55): Walters Shoal Seamount, Grid
WSL022, Station ALG10954, collected via sled (no 2) by the RV Algoa, (33°10.9' S;
43°48.6' E) - (33°11.2' S; 43°50.2' E), duration 41 min, depth 170 – 72 m, 29 May 2014. TS
2472 (WSL-INV84(1)), TS 2473 (WSL-INV84(2)), TS 2477 (WSL-INV84(6)), TS 2487
38
(WSL-INV84(16)), TS 2497 (WSL-INV84(26)), TS 2510 (WSL-INV84(39)): Walters Shoal
Seamount, Grid WSL042, Station ALG10974, collected via sled (no 4) by the RV Algoa,
(33°11.2' S; 43°51.0' E) - (33°11.2' S; 43°50.7' E), duration 10 min, depth 34 – 28 m, 02 June
2014.
Description. Thickly encrusting and amorphous form. Length 3.8 cm, diameter
3.0 cm and thickness 1.1 cm. Surface smooth and slippery, with ridges and randomly
scattered oscules. Oscules are non-circular, ~1 – 2 mm in diameter, often slightly indented.
Texture rubbery, firm and dense. Specimen not compressible, nor easily torn. In one
specimen (TS 2303), bright orange-red spherical eggs (~1 mm diameter) present. Colour in
situ dark red, light brown in preservative. Preservative becomes bright orange with time.
Most specimens leave a red-brown exudate on tissue paper.
Skeleton. Choanosome contains a multispicular reticulate skeleton, comprised of
robust fibres arranged somewhat radially, cored with oxeas. Fibres ~100 µm thick, sinuous,
running somewhat perpendicular to the surface, not differentiated into primary and secondary
tracts. Oxeas and arcuate chelae scattered throughout. Fibres penetrate ectosome, expanding
radially to form brushes. Ectosome contains erect, radial bouquets of oxeas that sometimes
pierce the surface, <200 µm thick.
Spiculation. Megascleres. Oxeas hastate, tornote-like, smooth, straight or slightly
curved: 316.2 (282.3 – 342.3) x 6.2 (4.2 – 7.7) µm, n = 10. Microscleres. Arcuate
unguiferous isochelae: 13.9 (12.4 – 15.4) µm, n = 10.
39
Substratum, Depth range and Ecology. Seven specimens found on rocky substrata
in two sleds, almost always in association with the same species of hydroid as epifauna.
Depth range: 28 – 170 m.
Geographic Distribution. South African Exclusive Economic Zone, Walters Shoal
Seamount.
Remarks. The present material conforms to both the original description of
Desmacidon ectofibrosa by Lévi (1963) and genus reassignment to Fibula ectyofibrosa
(Oxeas: 353 (318 – 382) x 14 (14) µm, n = 10; Chelae: 18 (18) µm, n = 10) by Samaai &
Gibbons (2005). However, in both cases, the material presented here was found to have a
slightly smaller megasclere width when compared to the descriptions above, and slightly
smaller chelae. Desmacidon ectofibrosa Lévi, 1963 was thought to be misplaced (and thus
reassigned) by Samaai & Gibbons (2005) based on the presence of arcuate chelae, which
would suggest the genus Fibulia, rather than Desmacidon, which has anchorate chelae (van
Soest 2002a). According to the World Porifera Database (May 2015, van Soest et al. 2015),
this species was reassigned to the genus Isodictya, but with no reference mentioned. As
Isodictya has palmate isochelae, this reassignment seems misplaced. Thus, the current
material is considered here a species of the genus Fibulia. According to Samaai & Gibbons
(2005), this species is common along the west coast of South Africa and exhibits a variety of
growth forms.
Family Latrunculiidae Topsent, 1922
Genus Latrunculia du Bocage, 1869
Subgenus Latrunculia (Biannulata) Samaai, Gibbons & Kelly, 2006
40
Latrunculia (Biannulata) sp. • (Fig. 12 A – F)
Material examined. TS 2563 (WSL-INV74(45)): Walters Shoal Seamount, Grid
WSL024, Station ALG10956, collected via sled (no 3) by the RV Algoa, (33°08.8' S;
43°49.1' E) - (33°09.0' S; 43°50.5' E), duration 33 min, depth 348 – 103 m, 29 May 2014.
Description. Thinly encrusting form on biogenic rubble. Length 2.0 cm, diameter
1.9 cm and thickness <0.1 cm. Surface slightly rough (probably due to the texture of the
rock), with very small areolate porefields, 0.2 mm in diameter. Not compressible, easily torn.
Colour in situ, and in preservative, black.
Skeleton. Choanosomal skeleton comprises an irregular polygonal reticulation
formed by wispy tracts of smooth styles. The tracts range in width from 60 – 90 µm and form
meshes that are 150 µm wide. Within the inner choanosome, tracts diverge towards the
surface. Interstitial spicules present. Ectosome comprises a palisade of densely packed,
interlocking anisodiscorhabds arranged vertically one spicule deep with their basal spinose
whorls buried in the outer ectosomal membrane.
Spiculation. Megascleres. Styles smooth, polytylote, straight or slightly sinuous with
elongated heads, often distally tornote: 263.5 (236.6 – 290.7) x 5.6 (4.7 – 6.6) µm, n = 10.
Microscleres. Anisodiscorhabds with well separated furcate whorls and hooked spines. Shaft
occasionally spined with undifferentiated basal whorl and manubrium: 39.6 (36.6 – 42.3) x
4.5 (3.6 – 5.0) µm, width including whorls: 22.3 (19.3 – 24.3) µm, n = 10.
41
Substratum, Depth range and Ecology. One specimen found on rocky substrata in a
single sled composed predominantly of dead clam shells and hydrozoans. Depth range: 103 –
348 m.
Geographic Distribution. Walters Shoal Seamount.
Remarks. The present material conforms to Latrunculia du Bocage, 1869 as
diagnosed by the presence of anisodiscorhabd microscleres (Samaai & Kelly 2002). In this
material, the microscleres have two whorls (mean and subsidiary) of spines on the shaft
which suggests placement in the subgenus Latrunculia (Biannulata) Samaai, Gibbons &
Kelly, 2006 (Samaai et al. 2006). This placement is supported by Dr Toufiek Samaai
(personal communication, April 1, 2015).
The present material is not conspecific with the five species of Latrunculia
(Biannulata) that have been recorded from the region of interest (Table 3): Latrunculia
(Biannulata) algoaensis Samaai, Janson & Kelly, 2012; Latrunculia (Biannulata) gotzi
Samaai, Janson & Kelly, 2012; Latrunculia (Biannulata) kerwathi Samaai, Janson & Kelly,
2012; Latrunculia (Biannulata) lunaviridis Samaai, Gibbons, Kelly & Davies-Coleman, 2003
and Latrunculia (Biannulata) microacanthoxea Samaai, Gibbons, Kelly & Davies-Coleman,
2003. Apart from L. (B.) kerwathi, which is thinly encrusting, all the other species have either
a thickly encrusting or massive semi-spherical form, often with volcano or cylindrical shaped
oscules. In addition, all the above species have microscleres that are visually distinct to those
found in the present material, which lacks a crown-like tuft of spines forming the apical
whorl and manubrium. Thus the present material likely constitutes a new species.
42
Family Microcionidae Carter, 1875
Subfamily Microcioninae Carter, 1875
Genus Clathria Schmidt, 1862
Subgenus Clathria (Clathria) Schmidt, 1862
Clathria (Clathria) sp. • (Fig. 13 A – K, Table 11)
Material examined. TS 2302 (WSL-INV54): Walters Shoal Seamount, Grid
WSL022, Station ALG10954, collected via sled (no 2) by the RV Algoa, (33°10.9' S;
43°48.6' E) - (33°11.2' S; 43°50.2' E), duration 41 min, depth 170 – 72 m, 29 May 2014. TS
2342 (WSL-INV94(14)), TS 2348 (WSL-INV94(20)), TS 2355 (WSL-INV94(27)): Walters
Shoal Seamount, Grid WSL044, Station ALG10976, collected via sled (no 6) by the RV
Algoa, (33°14.0' S; 43°55.5' E) - (33°13.7' S; 43°55.6' E), duration 9 min, depth 28 – 25 m,
02 June 2014. TS 2399 (WSL-INV92(10)), TS 2422 (WSL-INV92(11)): Walters Shoal
Seamount, Grid WSL043, Station ALG10975, collected via sled (no 5) by the RV Algoa,
(33°13.8' S; 43°55.5' E) - (33°13.1' S; 43°55.8' E), duration 11 min, depth 28 – 30 m, 02 June
2014. TS 2508 (WSL-INV84(37)), TS 2511 (WSL-INV84(40)): Walters Shoal Seamount,
Grid WSL042, Station ALG10974, collected via sled (no 4) by the RV Algoa, (33°11.2' S;
43°51.0' E) - (33°11.2' S; 43°50.7' E), duration 10 min, depth 34 – 28 m, 02 June 2014.
Description. Thickly encrusting, lobate form. Length 3.4 cm, diameter 2.5 cm and
thickness 1.3 cm. Surface undulating but smooth and velvety, with randomly scattered small,
round oscules (<1 mm in diameter), sunken with no distinct membranous lip. Texture soft
and spongy, compressible and easily torn. Colour in situ orange, beige in preservative.
43
Skeleton. Choanosomal skeleton regularly reticulate, forming irregular anastomoses
of differentiated primary and secondary fibres, diverging in plumoreticulate manner towards
ectosome. Fibres are differentiated into primary and secondary transverse components.
Primary fibres cored with principal styles, cemented by spongin that does not form a distinct
sheath around the fibre and echinated by acanthostyles. Secondary fibres with unispicular
tracts of principal styles. Ectosomal and subectosomal skeleton comprised of principal styles
and auxiliary subtylostyles, with the former arising from ascending choanosomal tracts being
slightly plumose and diverging into erect bundles which project obliquely through the
surface. The latter form compact diverging brushes at the ectosomal surface, barely
penetrating the subectosomal membrane. Microscleres scattered throughout choanosome.
Spiculation. Megascleres. Styles, smooth, curved, with well-rounded to almost
subtylote-like base, distally hastate: 234.3 (178.7 – 320.0) x 9.3 (7.9 – 11.5) µm, n = 10.
Subtylostyles, smooth, straight with terminally spined elongated base, distally fusiform:
211.4 (129.7 – 313.1) x 3.0 (2.4 – 3.8) µm, n = 10. Acanthostyles, straight to slightly bent,
with well-rounded to almost subtylote-like base, distally hastate: 138.0 (132.2 – 148.0) x 7.3
(5.6 – 9.7) µm, n = 10. Microscleres. Toxas, terminally spined, in two size classes: I) 146.1
(111.0 – 177.2) µm, n = 10; II) 45.1 (35.3 – 61.1) µm, n = 10. Palmate isochelae: 12.5 (11.2 –
14.2) µm, n = 10.
Substratum, Depth range and Ecology. Eight specimens found on rocky substrata
in four sleds. Depth range: 25 – 170 m.
Geographic Distribution. Walters Shoal Seamount.
44
Remarks. The present material conforms to Clathria (Clathria) Schmidt, 1862 as
diagnosed by a single category of auxiliary style and no marked difference between the axial
and extra-axial regions in the choanosomal skeleton (Hooper 2002). There are 26 species of
Clathria (Clathria) found within the region of interest (Table 3), none of which seem
conspecific with the material here.
Of these 26 species, three are similar to the present material, including Clathria
(Clathria) dayi Lévi, 1963; Clathria (Clathria) oculata Burton, 1933 and Clathria (Clathria)
inhacensis Thomas, 1979. Clathria (Clathria) dayi (Styles: 300 – 525 x 25 – 30 µm;
Auxiliary styles: 175 – 280 x 4 – 6 µm; Acanthostyles: 225 – 300 x 15 – 25 µm; Toxas: 175 –
300 x 2 – 5 µm; Chelae: 5 – 7 µ) is found within the southern African Exclusive Economic
Zone and has similar spicules to the present material. However, Lévi (1963) records spicules
which are larger and thicker than the material here, with unspined auxiliary styles and only
one class of toxa. Alternatively, C. (C.) oculata (Styles: 140 x 7µm; Auxiliary styles: 160 x
3 µm; Acanthostyles: 65 x 4µm; Toxas: 160 µm; Chelae: 6 µm) has slightly smaller and
narrower spicule sizes, but also has unspined auxiliary styles and only one class of smooth
toxa. This is also true for C. (C.) inhacensis (Styles: 121 – 172 (142) x 4 – 5 (4) µm;
Auxiliary styles: 124 – 181 (144) x 2 – 4 (3) µm; Acanthostyles: 41 – 58 (50) x 3 – 5 µm;
Toxas: 120 µm; Chelae: 8 – 10 µm), which has one class of hair-like toxa. Thus, the present
material likely constitutes a new species.
Order Suberitida Chombard & Boury-Esnault, 1999
Family Halichondriidae Gray, 1867
Genus Halichondria Fleming, 1828
Subgenus Halichondria (Halichondria) Fleming, 1828
45
Halichondria (Halichondria) sp. • (Fig. 14 A – F, Table 12)
Material examined. TS 2336 (WSL-INV94(7)), TS 2338 (WSL-INV94(10)), TS
2339 (WSL-INV94(11)), TS 2340 (WSL-INV94(12)), TS 2350 (WSL-INV94(22)): Walters
Shoal Seamount, Grid WSL044, Station ALG10976, collected via sled (no 6) by the RV
Algoa, (33°14.0' S; 43°55.5' E) - (33°13.7' S; 43°55.6' E), duration 9 min, depth 28 – 25 m,
02 June 2014. TS 2373 (WSL-INV75(14)), TS 2374 (WSL-INV75(15)), TS 2375 (WSLINV75(16)), TS 2377 (WSL-INV75(18)), TS 2378 (WSL-INV75(19)), TS 2379 (WSLINV75(20)), TS 2380 (WSL-INV75(21)): Walters Shoal Seamount, collected via SCUBA
(dive 1) by the RV Algoa, (33°11.2' S; 43°50.7' E), duration 35 min, depth 29 m, 30 May
2014. TS 2381 (WSL-INV83(1)), TS 2383 (WSL-INV83(3)), TS 2384 (WSL-INV83(4)), TS
2385 (WSL-INV83(5)), TS 2387 (WSL-INV83(7)): Walters Shoal Seamount, collected via
SCUBA (dive 2) by the RV Algoa, (33°10.6' S; 43°51.0' E), duration 36 min, depth 29 m, 30
May 2014. TS 2390 (WSL-INV92(1)), TS 2391 (WSL-INV92(2)), TS 2392 (WSLINV92(3)), TS 2393 (WSL-INV92(4)): Walters Shoal Seamount, Grid WSL043, Station
ALG10975, collected via sled (no 5) by the RV Algoa, (33°13.8' S; 43°55.5' E) - (33°13.1' S;
43°55.8' E), duration 11 min, depth 28 – 30 m, 02 June 2014. TS 2481 (WSL-INV84(10)),
TS 2482 (WSL-INV84(11)), TS 2483 (WSL-INV84(12)), TS 2484 (WSL-INV84(13)), TS
2485 (WSL-INV84(14)), TS 2486 (WSL-INV84(15)), TS 2490 (WSL-INV84(19)), TS 2492
(WSL-INV84(21)), TS 2499 (WSL-INV84(28)): Walters Shoal Seamount, Grid WSL042,
Station ALG10974, collected via sled (no 4) by the RV Algoa, (33°11.2' S; 43°51.0' E) (33°11.2' S; 43°50.7' E), duration 10 min, depth 34 – 28 m, 02 June 2014.
Description. Thickly encrusting, semi-spherical form. Length 5.0 cm, diameter
4.0 cm and thickness 2.5 cm. Surface smooth, uneven, with various ridge-like structures, and
46
covered with small volcano-shaped papillae. Oscules (1 – 2 mm) scattered randomly on the
upper surface, also occurring on the apex of volcano-shaped papillae (which become
depressed above water) in other specimens (e.g. TS 2338). Membrane present that covers the
exterior. Texture spongy and dense, of medium compressibility and easily torn. Colour in situ
dark brown, with very dark brown (almost black) regions, light brown with dark brown
regions in preservative. Specimen smells like soil and leaves a brown exudate on tissue paper.
Skeleton. Confused choanosomal skeleton, typically halichondrid, with oxeas of
variable length arranged in a disorderly fashion (spicules distributed randomly), showing
little tendency to form ascending tracts, and separated by well-developed subdermal spaces.
The ectosomal skeleton typically comprises a tangential spicule layer of varying thickness
(~100 – 300 µm), often becoming confused via intercrossing spicules. Spicules do not
penetrate the surface. Ectosome not readily detachable from choanosome.
Spiculation. Megascleres. Oxeas, smooth, straight to slightly curved, fusiform, in
three size classes: I) 403.3 (349.9 – 461.6) x 9.9 (6.2 – 13.6) µm, n = 10; II) 232.0 (208.0 –
288.4) x 7.7 (5.7 – 9.2) µm, n = 10; III) 145.3 (112.5 – 198.6) x 6.1 (5.0 – 7.4) µm, n = 10.
Microscleres. Absent.
Substratum, Depth range and Ecology. Thirty specimens found on rocky substrata
in three sleds and both dives, with ascidians, tube worms, coralline algae and/or hydroids as
epifauna. Depth range: 25 – 34 m.
Geographic Distribution. Walters Shoal Seamount.
47
Remarks. The present material has a typically halichondrid skeleton (Erpenbeck &
van Soest 2002), megascleres that are exclusively oxeas, as well as oscules on conical
elevations, and is thus placed in the genus Halichondria Fleming, 1828. It does not seem to
be conspecific with the seven species of Halichondria (Halichondria) Fleming, 1828 that
have been recorded from the region of interest (Table 3), based on morphology and the
presence of three size classes of oxeas.
Halichondria (Halichondria) capensis and Halichondria (Halichondria) gilvus, both
described by Samaai & Gibbons (2005) from the west coast of South Africa, and
Halichondria (Halichondria) prostrata Thiele, 1905 from Antarctica, have one size class of
oxeas (333 (319 – 355) x 12 (12) µm, n = 10; 391 (328 – 437) x 15 (9 – 18) µm, n = 10 and
300 – 320 x 9 µm respectively) with a relatively narrow size range. In addition, both H. (H.)
capensis and H. (H.) gilvus have conspicuous papillae, as opposed to the present material
which has irregular, spongy, easily deformed turrets. Within the Western Indian Ocean,
Halichondria (Halichondria) cartilaginea (Esper, 1794) and Halichondria (Halichondria)
lendenfeldi Lévi, 1961 also both have one size class of oxeas (185 – 203 x 3 – 4 µm; 400 –
600 x 11 – 13 µm respectively). These two species also differ from the present material
morphologically, with H. (H.) cartilaginea having a bushy form, with a slightly brittle
consistency and small (0.02 mm) pores, while H. (H.) lendenfeldi has a hispid, velvety
surface with many pores distributed over the entire surface. Halichondria (Halichondria)
aldabrensis Lévi, 1961 has two size classes of oxeas (I) 275 – 650 x 4 – 10 µm; II) 650 – 950
x 10 – 30 µm), which are larger than those found in the present material. Finally,
Halichondria (Halichondria) tenuiramosa Dendy, 1922 which occurs extensively in the
Indian Ocean, has one size class of very small oxeas (210 x 6 µm), with a creeping,
branching form. Thus, the present material likely constitutes a new species.
48
Family Suberitidae Schmidt, 1870
Genus Aaptos Gray, 1867
Aaptos sp. • (Fig. 15 A – H, Table 13)
Material examined. TS 2502 (WSL-INV84(31)), TS 2503 (WSL-INV84(32)):
Walters Shoal Seamount, Grid WSL042, Station ALG10974, collected via sled (no 4) by the
RV Algoa, (33°11.2' S; 43°51.0' E) - (33°11.2' S; 43°50.7' E), duration 10 min, depth 34 –
28 m, 02 June 2014.
Description. Thickly encrusting form. Length 1.9 cm, diameter 1.7 cm and thickness
0.7 cm. A dense array of spicules at the surface (~1 mm), arranged in a confused fashion
rendering the surface prickly to the touch. No visible oscules. Texture dense and firm, barely
compressible specimens tear so-so. Colour in situ dull black externally, almost appearing
grey due to visible spicules at the surface. Internal colour in situ beige. Colour in preservative
dull brown externally, internally grey-beige.
Skeleton. Choanosomal skeleton comprises dense tracts of megascleres (~230 –
290 µm wide) that arise from the base and radiate vertically through the choanosome, fanning
out and forming brushes into the ectosome. These brushes form a dense palisade at the
surface, with smaller spicules intermingled (often perpendicular to surface) between the
larger spicules. Subectosomal region consists of a layer of densely packed, tangentially
orientated megascleres. Ectosome consists of small styles and larger intermediate styles,
which form palisades of vertically arranged brushes. The distal ends of these megascleres
protrude through sponge surface.
49
Spiculation. Megascleres. Strongyloxeas, smooth, straight to slightly bent, thickest
centrally with reduced apices, distally fusiform: 954.4 (677.5 – 1284.6) x 14.1 (7.5 –
20.0) µm, n = 10. Styles, smooth, straight to slightly bent, often thickest centrally in largest
size class, distally fusiform, in three size classes: I) 875.8 (674.1 – 1252.4) x 27.4 (23.6 –
32.3) µm, n = 10; II) 446.0 (348.3 – 576.4) x 14.9 (8.9 – 19.7) µm, n = 10; III) 188.3 (127.5 –
291.1) x 5.0 (3.0 – 6.9) µm, n = 10. Microscleres. Absent.
Substratum, Depth range and Ecology. Two specimens found in one sled on rocky
substrate. Depth range: 28 – 34 m.
Geographic Distribution. Walters Shoal Seamount.
Remarks. The present material has a radiate skeleton of strongyloxeas, with a dense
ectosomal palisade and is thus placed in the genus Aaptos Gray, 1867 (van Soest 2002b). It
does not seem to be conspecific with the two species of Aaptos that have been recorded from
the region of interest (Table 3), including Aaptos alphiensis Samaai & Gibbons, 2005 and
Aaptos nuda (Kirkpatrick, 1903). Aaptos alphiensis was described by Samaai & Gibbons
(2005) from the west coast of South Africa, as having both primary and intermediate
subtylostyles, intermediate styles and dermal tylostyles, while Kirkpatrick (1903) notes the
presence of only oxeas in A. nuda.
There have been several records of Aaptos aaptos (Schmidt, 1864), which has both
strongyloxeas and styles, within the region of interest. However, the World Porifera Database
(van Soest et al. 2015) suggests these records are inaccurate due to the geographic
distribution of this species which has been reported from many areas around the world. Thus,
the present material likely constitutes a new species.
50
Order Tethyida Morrow & Cárdenas, 2015
Family Tethyidae Gray, 1848
Genus Tethya Lamarck, 1815
Tethya sp. • (Fig. 16 A – G, Table 14)
Material examined. TS 2311 (WSL-INV50(2)), TS 2327 (WSL-INV40): Walters
Shoal Seamount, Grid WSL022, Station ALG10954, collected via sled (no 2) by the RV
Algoa, (33°10.9' S; 43°48.6' E) - (33°11.2' S; 43°50.2' E), duration 41 min, depth 170 – 72 m,
29 May 2014. TS 2337 (WSL-INV94(8)), TS 2349 (WSL-INV94(21)), TS 2352 (WSLINV94(24)), TS 2358 (WSL-INV94(30)): Walters Shoal Seamount, Grid WSL044, Station
ALG10976, collected via sled (no 6) by the RV Algoa, (33°14.0' S; 43°55.5' E) - (33°13.7' S;
43°55.6' E), duration 9 min, depth 28 - 25m, 02 June 2014. TS 2362 (WSL-INV75(3)), TS
2363 (WSL-INV75(4)), TS 2376 (WSL-INV75(17)): Walters Shoal Seamount, collected via
dive (no 1) by the RV Algoa, (33°11.2' S; 43°50.7' E), duration 35 min, depth 29 m, 30 May
2014. TS 2420 (WSL-INV24(a)): Walters Shoal Seamount, Grid WSL021, Station
ALG10953, collected via sled (no 1) by the RV Algoa, (33°11.0' S; 43°53.9' E) - (33°11.0' S;
43°52.9' E), duration 40 min, depth 53 – 43 m, 29 May 2014. TS 2430 (WSL-INV119(4)):
Walters Shoal Seamount, collected via lobster trap by the RV Algoa, (33°11.6' S; 43°50.5' E),
duration 328 min, depth 39 m, 05 June 2014. TS 2474 (WSL-INV84(3)), TS 2489 (WSLINV84(18)), TS 2493 (WSL-INV84(22)), TS 2496 (WSL-INV84(25)), TS 2498 (WSLINV84(27)): Walters Shoal Seamount, Grid WSL042, Station ALG10974, collected via sled
(no 4) by the RV Algoa, (33°11.2' S; 43°51.0' E) - (33°11.2' S; 43°50.7' E), duration 10 min,
depth 34 – 28 m, 02 June 2014. TS 2538 (WSL-INV102(4)): Walters Shoal Seamount, Grid
51
WSL045, Station ALG10977, collected via sled (no 7) by the RV Algoa, (33°13.8' S;
43°56.1' E) - (33°14.2' S; 43°55.9' E), duration 16 min, depth 80 m, 02 June 2014.
Description. Spherical to semi-spherical form. Length 1.6 cm, diameter 1.4 cm and
thickness 1.4 cm. Surface rough and fuzzy, but undulating and smooth in a couple of
specimens. In other specimens, one (rarely two) oscules present on apex (~1 mm). Welldeveloped ectosome, ~1 – 2 mm thick, which is distinct but not separable from the
choanosome. Texture tough, firm and dense. Not compressible, nor easily torn. Colour in situ
pale beige (with brown tinge) externally, olive green internally, with a white centre. In
preservative, pale beige. Ectosome colour in situ, and in preservative, white. Slightly sticky
exudate present in a few specimens.
Skeleton. Choanosomal skeleton radial, comprising compact anisostrongyloxea and
(aniso)strongyle (rare) tracts (~200 µm across) radiating from the centre of the sponge, often
penetrating the ectosome as expanding dermal brushes, with megascleres piercing the sponge
surface. Somewhat confused interstitial anisostrongyloxeas fill the space among the main
megasclere bundles. Microscleres are common in the inner choanosome between the tracts.
Thick, discernible ectosome (>1000 µm) comprised of small radial bouquets (~400 – 600 µm
across) of megascleres embedded within this region, which pierce the sponge surface.
Megasters (represented by spherasters) and micrasters are densely packed in ectosome,
somewhat entering the upper regions of the choanosome, with the former decreasing in size
from the sponge surface, inwards.
Spiculation. Megascleres. Primary and auxiliary anisostrongyloxeas, smooth,
straight, thickest centrally, with reduced, somewhat elongate apices, often distally hastate,
52
with no easily discernible size classes (continuous) and large size range: 292.7 – 1280.1 x
10.5 (5.6 – 22.7) µm, n = 10. Strongyles to anisostrongyles, relatively rare, smooth, straight,
thickest centrally, often fusiform, with no easily discernible size class: 995.6 (595.6 – 1249.3)
x 19.2 (9.0 – 24.9) µm, n = 10. Microscleres. Megasters - Spherasters with ~15 rays: 37.0
(21.3 – 56.0) µm, n = 10. Micrasters - Tylasters with ~11 terminally spined rays: 12.6 (10.5 –
15.1) µm, n = 10; Spheroxyasters with ~8 rays: 6.2 (5.3 – 7.0) µm, n = 10.
Substratum, Depth range and Ecology. Seventeen specimens found on rocky
substratum in five sleds, in the lobster trap and during one dive. This species found in
association with tube worms, bivalves and algae (in the form of epifauna). Depth range: 25 –
170 m.
Geographic Distribution. Walters Shoal Seamount.
Remarks. The present material conforms well to Tethya Lamarck, 1815, as diagnosed
by a spherical form, well-developed, distinct ectosome and main skeleton formed by radiating
strongyloexa bundles (Sarà 2002). It does not seem to be conspecific with the nine species of
Tethya that have been recorded from the region of interest (Table 3): Tethya globostellata
Lendenfeld, 1897; Tethya japonica Sollas, 1888; Tethya magna Kirkpatrick, 1903; Tethya
parvistella (Baer, 1906); Tethya peracuta (Topsent, 1918); Tethya robusta (Bowerbank,
1873); Tethya rubra Samaai & Gibbons, 2005; Tethya seychellensis (Wright, 1881) and
Tethya stellagrandis (Dendy, 1916).
Tethya globostellata Lendenfeld, 1897 (Anisostrongyloxeas: 1000 – 2100 x 24 –
32 µm; Styles: 400 – 500 x 14 – 16 µm; Amphistrongyles: 1000 – 1500 x 33 µm;
Oxyasters: 60 – 100 µm; Strongylasters: 9 – 12 µm) and T. parvistella (Baer, 1906)
53
(Anisostrongyloxeas: 718 – 1342 x 3 – 18 µm; Amphistrongyles: 841 – 1100 x 14 – 18 µm;
Sphaerasters: 33 – 59 µm; Tylasters: I) 7 µm, II) 11 µm) somewhat resemble the present
material. However, all the above-mentioned species lack the smallest spheroxyasters. Thus,
the present material likely constitutes a new species.
Order Tetractinellida Marshall, 1876
Suborder Astrophorina Sollas, 1887
Family Ancorinidae Schmidt, 1870
Genus Ancorina Schmidt, 1862
Ancorina sp. • (Fig. 17 A – L, Table 15)
Material examined. TS 2475 (WSL-INV84(4)), TS 2476 (WSL-INV84(5)): Walters
Shoal Seamount, Grid WSL042, Station ALG10974, collected via sled (no 4) by the RV
Algoa (Voyage 208), (33°11.2' S; 43°51.0' E ) - (33°11.2' S; 43°50.7' E ), duration 10 min,
depth 34 – 28 m, 02 June 2014.
Description. Massive, amorphous form. Length 9.4 cm, diameter 5.6 cm and
thickness 3.4 cm. Surface microhispid, and thus prickly to the touch. A few oscules evident
on the ridge (0.5 – 1 mm) and several (1 – 2 mm) on the underside of specimen TS 2476.
Texture firm, dense and slightly rubbery. Barely compressible, not easily torn. Ectosome
(~2 mm) present, not separable from the choanosome and yellow in situ, white in
preservative. Colour in situ dark brown with yellowish tinge and darker brown, almost black
ridges externally and paler brown internally. In preservative, dark brown externally and paler
brown internally. Water retentive, leaving a brown exudate on tissue paper.
54
Skeleton. Choanosomal skeleton consists of radiating tracts of plagiotriaenes and
oxeas. Tracts of large oxeas occur between the plagiotriaenes in mid- and deep choanosomal
layers of the sponge. Oxyasters abundant and scattered throughout. Towards the surface, the
tracts become denser and are entirely composed of plagiotriaenes with overlapping cladi.
Ectosomal skeleton comprises a thick discernible layer (>1000 µm) with radiating
plagiotriaene tracts that pierce the surface, through a dense (up to ~100 µm thick) layer of
sanidasters.
Spiculation. Megascleres. Oxeas, smooth, straight to slightly bent, in two size
classes: I) 1748.4 (1276.7 – 2017.8) x 30.3 (16.9 – 36.8) µm, n = 10; II) 975.5 (727.5 –
1133.7) x 9.1 (6.3 – 12.5) µm, n = 10. Plagiotriaenes with short, stout cladi, rhabdome
straight to slightly bent, in three size classes: I) rhabdome 1759.8 (1550.3 – 2074.9) x 38.5
(33.4 – 46.5) µm, cladome 152.3 (130.3 – 175.0) µm, cladi 89.9 (75.1 – 116.9) µm, n = 10;
II) rhabdome 976.3 (924.1 – 1037.1) x 19.9 (16.6 – 23.8) µm, cladome 65.5 (51.5 – 84.4) µm,
cladi 29.8 (18.8 – 38.2) µm, n = 10; III) rhabdome 608.2 (457.8 – 766.9) x 10.8 (6.1 –
18.4) µm, cladome 31.8 (19.6 – 53.4) µm, cladi 13.9 (8.4 – 24.0) µm, n = 10. Microscleres.
Oxyasters with ~10 rays, smooth or with hooked spines: 10.9 (8.5 – 14.6) µm, n = 10.
Acanthoxyasters with 4 rays and hooked spines: 18.2 (15.7 – 22.1) µm, n = 10.
Acanthoxyasters, reduced tetracts with hooked spines, variable in form and spinosity: 19.2
(14.6 – 23.5) µm, n = 10. Sanidasters, acanthose, irregularly spined: 5.9 (5.2 – 6.8) µm, n =
10.
Substratum, Depth range and Ecology. Two specimens found in one sled on rocky
substrate in association with tube worms and algae. Depth range: 28 – 34 m.
55
Geographic Distribution. Walters Shoal Seamount.
Remarks. The present material conforms to Ancorina Schmidt, 1862 as diagnosed by
a conspicuous ectosome, the presence of oxeas and triaenes as megascleres and microscleres
comprising sanidasters and euasters (Uriz 2002). Two species are present in the region of
interest (Table 3): Ancorina corticata Lévi, 1964 and Ancorina nanosclera Lévi, 1967. The
present material is not conspecific with the latter species due to the presence of anatriaenes in
the material described by Lévi (1967) and seems more similar to A. corticata (Oxeas: 2000 –
2400 x 50 µm; Plagiotriaenes: rhabdome 1400 x 70 µm, cladi 130 – 150 x 50 µm; Oxyasters:
15 – 20 µm; Sanidasters: 6 µm), which lacks anatriaenes.
However, the present material differs by having two size classes of oxeas, three size
classes of plagiotriaenes and reduced tetract acanthoxyasters. Ancorina corticata was also redescribed by Samaai & Gibbons (2005), with scanning electron microscope images of the
sanidasters provided, which look vastly different to the sanidasters found here. Thus, the
present material likely constitutes a new species.
Genus Chelotropella Lendenfeld, 1907
Chelotropella sp. • (Fig. 18 A – L)
Material examined. TS 2310 (WSL-INV50(1)): Walters Shoal Seamount, Grid
WSL022, Station ALG10954, collected via sled (no 2) by the RV Algoa (33°10.9' S; 43°48.6'
E) - (33°11.2' S; 43°50.2' E), duration 41 min, depth 170 – 72 m, 29 May 2014.
56
Description. Spherical form. Length 1.9 cm, diameter 1.5 cm and thickness 1.7 cm.
Surface microhispid and prickly to the touch. One oscule (~3 mm) present at the top of the
specimen. Thin ectosome (~1 mm) present, separable from the choanosome. Texture firm and
dense, not compressible. Colour in situ dull dark brown externally, paler brown internally. In
preservative, colour light brown. Slightly sticky exudate.
Skeleton. Choanosomal skeleton comprises thick, radial tracts of oxeas and triaenes
(~200 – 400 µm across), forming two subdermal layers in the peripheral region. Calthrops
arranged in a somewhat disorganized fashion, occasionally congregating in horizontal
formations, parallel to sponge surface. Subectosomal skeleton comprises large subdermal
cavities, triaenes orientated radially, with cladomes forming two layers parallel to the surface.
Strongyloacanthasters concentrated in the ectosome (~300 – 500 µm), and scattered
throughout the peripheral region, including around subdermal spaces.
Spiculation. Megascleres. Oxeas, smooth, straight to slightly curved, distally
fusiform: 2148.9 (1097.4 – 3015.5) x 20.3 (10.9 – 30.9) µm, n = 10. Dichotriaenes, rare, in
two size classes: I) often broken, rhabdome 2812.1 (2615.1 – 3077.6) x 51.1 (49.1 –
52.4) µm, cladome 501.3 (360.5 – 573.0) µm, stout protoclads 135.4 (110.8 – 153.5) µm,
long deuteroclads terminating in somewhat blunt points 169.4 (89.3 – 221.8) µm, n = 3; II)
rhabdome 1142.7 (786.5 – 1501.2) x 28.6 (23.6 – 33.7) µm, cladome 288.4 (253.9 –
333.9) µm, stout protoclads 117.6 (103.2 – 134.9) µm, short deuteroclads terminating in
somewhat sharp points 38.8 (20.9 – 49.6) µm, n = 7. Anatriaenes: 1146.7 (806.5 – 1437.5) x
9.4 (8.1 – 11.1) µm, with cladome 71.8 (53.5 – 84.6) µm, n = 10. Plagiotriaenes, shortshafted, rare: 302.2 (111.0 – 694.4) x 15.5 (9.7 – 21.8) µm, with cladome 145.4 (62.9 –
259.9) µm, and cladi 77.2 (34.9 – 136.2) µm, n = 8. Calthrops, regular in shape, found in two
57
size classes (ray): I) 474.1 (403.9 – 595.3) x 52.1 (43.3 – 60.7) µm, n = 10; II) 190.4 (134.0 –
259.3) x 24.4 (14.9 – 33 .8) µm, n = 10. Microscleres. Strongyloacanthasters with ~10
terminally hook-spined rays: 18.8 (14.1 – 23.1) µm, n = 10.
Substratum, Depth range and Ecology. One specimen found in a single sled on
rocky substrate, which was host to many bivalves and sponges. Depth range: 72 – 170 m.
Geographic Distribution. Walters Shoal Seamount.
Remarks. The present material conforms to Chelotropella Lendenfeld, 1907 as
diagnosed by the presence of calthrops, oxeas and peripheral dichotriaenes which form a
radial skeleton and two subdermal layers in the peripheral region (van Soest & Hooper 2002).
Erected by Lendenfeld in 1907 for a single species, this genus comprises two described
species to date: Chelotropella sphaerica Lendenfeld, 1907 and Chelotropella neocaledonica
Lévi & Lévi, 1983, of which only the former occurs in the region of interest, with the latter
found in New Caledonia.
Although similar to C. sphaerica with regards to morphology (spherical sponge of
~1.8 cm with granular surface as described by Lendenfeld in 1907), the spicular component
of the present material differs. The material in this study has megascleres that are smaller and
narrower than those described by Lendenfeld (1907) (Oxeas: 3600 – 5600 x 50 – 80 µm;
Dichotriaenes: rhabdome 2800 – 4400 x 100 – 440 µm, cladome 650 – 1300 µm, clades 130
– 170 µm; Calthrops: I) 700 – 1050 x 85 – 120 µm, II) 170 – 700 x 20 – 85 µm), with his
species also lacking anatriaenes and plagiotriaenes (although intermediate forms between
calthrops and triaenes are noted), but including the presence of various euaster morphologies.
58
Pulitzer-Finali (1993) record this species from Kenya, also with various euaster
morphologies, but with megasclere size ranges more in accordance with the present material
(Oxeas: 3500 – 4500 x 27 – 45 µm; Dichotriaenes: rhabdome 2400 x 80 µm, cladome
1600 µm, protoclads 270 µm, deuteroclads 500 µm; Calthrops: 300 – 760 µm). These authors
also found long anatriaenes (rhabdome 4000 x 20 – 36 µm, cladome 150 – 170 µm), which
led van Soest & Hooper (2002) to suggest that their material may be a new species distinct
from C. sphaerica.
Thus, due to the presence of anatriaenes (in a much smaller size range than recorded
by Pulitzer-Finali in 1993) and plagiotriaenes, as well as the lack of diverse euaster
morphologies (only one type found), the present material likely constitute a new species.
Family Geodiidae Gray, 1867
Subfamily Erylinae Sollas, 1888
Genus Penares Gray, 1867
Penares intermedia (Dendy, 1905) (Fig. 19 A – J, Table 16)
Synonymy
Plakinastrella intermedia Dendy, 1905: p. 67, Pl. I, Fig. 4, Pl. II, Fig. 2.
Material examined. TS 2300 (WSL-INV58), TS 2307 (WSL-INV57(2)), TS 2314
(WSL-INV51): Walters Shoal Seamount, Grid WSL022, Station ALG10954, collected via
sled (no 2) by the RV Algoa (Voyage 208), (33°10.9' S; 43°48.6' E) - (33°11.2' S; 43°50.2'
E), duration 41 min, depth 170 – 72 m, 29 May 2014. TS 2445 (WSL-INV74(11)), TS 2446
(WSL-INV74(12)), TS 2447 (WSL-INV74(13)), TS 2451 (WSL-INV74(17)), TS 2454
(WSL-INV74(20)), TS 2548 (WSL-INV74(30)), TS 2555 (WSL-INV74(37)): Walters Shoal
59
Seamount, Grid WSL024, Station ALG10956, collected via sled (no 3) by the RV Algoa
(Voyage 208), (33°08.8' S; 43°49.1' E) - (33°09.0' S; 43°50.5' E), duration 33 min, depth 348
– 103 m, 29 May 2014.
Description. Thickly encrusting form. Length 1.5 cm, diameter 1.7 cm and thickness
0.3 cm. Surface undulating but smooth with oscules (<1 mm) scattered randomly over the
surface. Thin ectosome (<1 mm) present, separable from the choanosome. Texture firm,
tough, dense and leathery. Specimen not compressible, easily torn. Colour in situ dull orangebrown externally and internally, pale olive green in preservative.
Skeleton. Confused choanosomal skeleton, comprising of oxeas and microxeas
arranged in a disorderly fashion (spicules distributed randomly), showing little tendency to
form tracts. Larger oxeas sometimes aggregating in loose (somewhat radial) slanting bundles
(~60 – 140 µm across). Dichotriaenes form subdermal skeleton, with cladome at surface and
rhabdome inwards. Oxyasters abundant and scattered throughout. Ectosomal skeleton
comprised of small oxeas, lying tangentially over dichotriaene clads, forming dense dermal
crust ~200 – 300 µm thick.
Spiculation. Megascleres. Oxeas, slightly curved, in three size classes: I) 840.4
(703.0 – 999.1) x 27.4 (21.8 – 36.8) µm, n = 10; II) 408.6 (318.2 – 505.7) x 18.7 (14.9 –
21.9) µm, n = 10; III) 140.7 (117.9 – 164.3) x 9.8 (7.1 – 12.3) µm, n = 10. Dichotriaenes,
with short rhabdomes, in two size classes: I) rhabdome not seen (~half the size of the
cladome), cladome 487.1 (380.8 – 578.8) µm, stout protoclads 90.6 (68.4 – 113.3) x 37.8
(27.5 – 48.6) µm, deuteroclads terminating in somewhat blunt points, often irregular at tips
145.3 (115.9 – 178.2) x 30.2 (20.1 – 38.7) µm, n = 10; II) rhabdome not seen (~half the size
60
of the cladome), cladome 325.1 (226.3 – 478.4) µm, thin protoclads 89.0 (74.5 – 102.8) x
21.0 (14.2 – 26.9) µm, deuteroclads terminating in sharp points 69.0 (28.3 – 122.8) x 15.1
(7.2 – 18.5) µm, n = 10. Microscleres. Microxeas, curved: 75.1 (62.6 – 92.1) x 6.0 (5.2 –
7.0) µm, n = 10. Acanthoxyasters with ~16 slender rays, hooked spines and sharply pointed
tips: 9.3 (7.7 – 12.3) µm, n = 10.
Substratum, Depth range and Ecology. Ten specimens found on rocky substrata in
two sleds, one consisting of predominantly bivalves and sponges, the other of biogenic debris
and hydrozoans. Depth range: 72 – 348 m.
Geographic Distribution. Sri Lanka (Holotype), Zanzibar, North Kenya Banks,
Walters Shoal Seamount.
Remarks. The present material conforms to Penares intermedia (Dendy, 1905)
originally described as Plakinastrella intermedia (Oxeas: I) 1200 x 37 µm, II) 180 x 10 µm;
Dichotriaenes: rhabdome 370 x 55 µm, with protoclads 92 x 55 µm; Oxyasters: 25 µm) and
further records of this species by Pulitzer-Finali (1993) (Oxeas: I) 1000 – 1500 x 33 –
62 µm, II) Oxeas: 75 – 410 x 5.5 – 22 µm; Dichotriaenes: rhabdome 190 µm, protoclads
95 µm, deuteroclads 160 µm; Oxyasters: 12 – 23 µm) and Thomas (1984) (Oxeas: I) 790 x
30 µm, II) 190 x 6 – 12 µm; Dichotriaenes: protoclads 80 x 50 µm, deuteroclads 280 x 5 µm;
Oxyasters: 18 µm).
Although Dendy (1905) only described one size class of dichotriaenes for P.
intermedia, he does make note of ‘slenderer’ forms which he suggests are not fully
developed. In addition, while providing two size classes of oxeas, he notes a large size range.
The present material definitely has spined oxyasters, but the spines are only visible through
61
the use of a scanning electron microscope, which explains the (slightly larger) ‘smooth’
oxyasters given in the original description. When viewed under a light microscope, the
oxyasters of the present material also appear smooth. Thomas (1984) noted minutely spined
oxyasters in his material.
Burton (1959) suggested that a similar species described by Dendy (1905),
Plakinastrella (now Penares) schulzei (Dendy, 1905), is conspecific with P. intermedia,
based on both the similarities in the figures drawn and a re-examination of the types. This
suggestion was followed by Thomas (1984), but neglected by Pulitzer-Finali (1993). To date,
P. intermedia and P. schulzei remain separate on the World Porifera Database (May 2015,
van Soest et al. 2015) and are thus considered distinct here.
3.3 Location and depth affiliations
Location
Fifty-five and 39 sponge species were collected from the western and eastern flank of Walters
Shoal Seamount respectively. Twenty-one new species were found on the western flank, with
11 of these restricted to this location, while 15 new species were found on the eastern flank,
with five of these restricted to this location (Table 17).
There was no clear pattern in the distribution of sponge assemblages on Walters Shoal
Seamount with regards to location (western vs. eastern flank; ANOSIM, R = -0.296, p =
0.839), with this finding illustrated in Fig. 20. Although SIMPER results indicate an average
dissimilarity of ~68% between the western and eastern side of the seamount (Table 18),
Table 17 documents several species that are shared by both sides (e.g. Halichondria
(Halichondria) sp. and Eurypon sp. 1).
62
This finding is further supported in that sampling locations on the western side of the
seamount had an average low sponge faunal similarity of ~35% (SIMPER), with the species
contributing to 90% of this similarity consisting of Callyspongia (Callyspongia) sp.
(26.25%), Halichondria (Halichondria) sp. (26.25%), Stelletta
purpurea Ridley, 1884
(26.25%) and Tethya sp. (11.41%). On the eastern side of Walters Shoal Seamount, sampling
locations had an overall lower sponge faunal similarity of ~19% (SIMPER), with the species
contributing to 90% of this similarity consisting of Clathria (Clathria) sp. (29.55%),
Halichondria (Halichondria) sp. (29.55%), Rhabderemia sp. (20.45%) and Protosuberites sp.
3 (20.45%).
Depth
The shallow and mesophotic depth zones of Walters Shoal Seamount had a similar number of
species present (27 and 28 respectively), with the submesophotic depth zone having the most
number of species present at 40. Species that are likely new were found predominantly in the
submesophotic depth zone (17), followed by the shallow depth zone (eight) and finally the
mesophotic depth zone (six). Fifteen new species were found exclusively in the
submesophotic depth zone, followed by five in the shallow depth zone, and only one in the
mesophotic depth zone (Table 19).
There was a clear pattern in the depth distribution of sponge assemblages on Walters Shoal
(shallow, mesophotic, submesophotic; ANOSIM, R = 0.609, p = 0.018), with this finding
illustrated in Fig. 21. Each depth zone had a distinct sponge assemblage, with the species
contributing to 90% (100% in the submesophotic zone) of sampling location similarity in
each depth zone provided in Table 20. The percent contribution of families and genera per
depth zone are given in Table 21, indicating that the family Ancorinidae was well represented
63
throughout, with Axinellidae the predominant family in the submesophotic zone. The genus
Stelletta was well represented in all depth zones, Callyspongia in both the shallow and
mesophotic zones and finally Phakellia and Protosuberites in the deepest zone.
The mesophotic zone acts as a transition between the shallow and submesophotic zones,
sharing eight and nine species with these zones respectively. The sponge fauna inhabiting the
shallow and submesophotic zones of the seamount were the most dissimilar, with only five
shared families (Table 22), three shared genera (Table 23) and three shared species
throughout, including Callyspongia (Toxochalina) cf. robusta (Ridley, 1884), Stelletta
purpurea Ridley, 1884 and M1. Further SIMPER results quantifying the percentage
difference between depth zones, and the species contributing to at least 60% of this
difference, are provided in Table 24.
3.4 Biogeographical affiliations
According to the 23 known sponge species recorded from Walters Shoal in this study, the
seamount demonstrates a relatively low similarity to surrounding regions. The highest
affinities were with the Western Indo-Pacific (21.8% shared species) and Temperate Southern
African (10.3% shared species) realms. No affiliations were found with Vema Seamount, the
Temperate South American or Southern Ocean realms. At the province level, Walters Shoal
Seamount demonstrates the most affiliation with the Western Indian Ocean (21.8% shared
species), Agulhas (9.0% shared species) and Benguela (5.1% shared species) provinces.
Within these provinces, the sponge fauna was most similar to that found in the East African
Coral Coast Ecoregion (12.8% shared species), followed by the Seychelles as well as the
Western and Northern Madagascar (both 10.3% shared species) ecoregions. Affiliations with
64
the remaining ecoregions in the Western Indian Ocean (excluding Southeast Madagascar) and
Temperate Southern African (excluding Amsterdam-St Paul) provinces were approximately 1
– 5% shared species (see Table 25).
At higher taxonomic levels (including all OTU’s) the sponge fauna of Walters Shoal was
comprised predominantly of species in the Ancorinidae (12.7%), Halichondriidae (10.9%),
Axinellidae (9.1%) and Suberitidae (9.1%) families. This was consistent with the surrounding
regions, with more than half of the ecoregions having a large representation of the family
Ancorinidae (see Table 26). The Northern Monsoon Current Coast, Seychelles, Delagoa and
Natal ecoregions have a fauna dominated by this family. Alternatively, Halichondriidae was
only relatively well represented in the East African Coral Coast and Seychelles ecoregions,
Axinellidae in the East African Coral Coast, Cargados Carajos/Tromelin Island and Delagoa
ecoregions and Suberitidae only at Walters Shoal Seamount. Stelletta, Phakellia and
Protosuberites were the most represented genera at Walters Shoal at 7.8%, 5.9% and 5.9%
respectively. Phakellia and Protosuberites were not well represented in the other ecoregions,
while Stelletta was relatively well represented in the Mascarene Islands, Delagoa and Natal
ecoregions.
Thirty-nine percent of the known sponge species found at Walters Shoal Seamount are
widely distributed in the Indian Ocean (Fig. 22). Of these, five species – Callyspongia
(Toxochalina) robusta (Ridley, 1884), Chondrosia debilis Thiele, 1900, Discodermia
panoplia Sollas, 1888, Stelletta purpurea Ridley, 1884 and Zyzzya fuliginosa (Carter, 1879) –
have distributions that also extend into the Pacific Ocean. A similar number of species (35%)
are found exclusively within the Western Indian Ocean region, with this study representing
the southernmost distribution record for several of these (e.g. Amorphinopsis fistulosa
(Vacelet, Vasseur & Lévi, 1976) and Axinyssa aplysinoides (Dendy, 1922)). Twenty-six
65
percent of the known species recorded from this study have a restricted distribution around
South Africa.
66
Chapter 4 – Discussion
This thesis constitutes the only study dedicated exclusively to the diversity, distribution and
biogeographical affiliations of the sponge fauna of Walters Shoal Seamount and augments the
current knowledge of sponges in the very data-sparse Western Indian Ocean region, including
the little-known seamount habitat.
4.1 Diversity
A total of 255 sponge specimens were collected from Walters Shoal Seamount, comprising
78 operational taxonomic units (OTU’s) or putative species. Twenty-three of these are
known, 26 likely constitute species new to science and potential endemics, 16 could only be
identified to higher taxonomic levels and 13 could only be designated as morphospecies due
to a lack of diagnostic material. A large proportion (~80%) of the OTU’s were assigned to the
class Demospongiae, which includes about 80% of all described sponge species worldwide
(Hooper & van Soest 2002, van Soest et al. 2012).
This study represents one of the highest records of sponge faunal diversity from seamount
studies thus far, with other works recording less than 40 species (Lévi 1969, SchlacherHoenlinger et al. 2005, Xavier & van Soest 2007). This could possibly be attributed to
limited sampling in previous studies (as noted by Schlacher-Hoenlinger et al. 2005 and
Xavier & van Soest 2007), as well as the inclusion of deeper specimens in the current study.
It could also reflect biogeographical affinities of Walters Shoal with the highly diverse
Western Indo-Pacific Realm (Roberts et al. 2002) and/or global patterns of sponge diversity,
with higher numbers in the tropics (van Soest et al. 2012).Walters Shoal is also somewhat
67
isolated (Groeneveld et al. 2006, Gopal 2007), possibly leading to diversification (Kadmon &
Allouche 2007). In addition, the region is subject to a wide range of biogeographic and/or
oceanographic features, as suggested by Laptikhovsky et al. (2015) to explain the high
diversity of cephalopod fauna from both the Southwest Indian Ocean and Madagascar Ridge
(sampled just northwest of Walters Shoal). Finally, the coralligenous-like substrate may
generate small-scale spatial complexity and allow for the formation of heterogeneous
microhabitats (Bertolino et al. 2013). This in turn might enable diversification, especially
with regards to small cryptic species, with many of the sponges documented from Walters
Shoal Seamount, especially in the deeper regions, being morphologically similar to those
recorded for Mediterranean coralligenous accretions by Bertolino et al. (2013).
In contrast, Collette & Parin (1991) recorded a relatively depauperate shallow-water fish
community of 20 species from the seamount, similar to a temperate rocky fish community,
although with less diversity. This is possibly due to the absence of (larger scale) structural
complexity, as well as limited food resources, with the maximum accumulation of vertically
migrating zooplankton occurring just below the photic zone, and the supply declining over
very shallow structures that occur within this layer (Genin 2004, Genin & Dower 2007). The
discrepancy between seamount ichthyofauna and benthic communities has been recorded
previously, with mobile plankton and pelagic fish species often similar to (or the same as)
those from nearby oceanic pelagic communities, while sessile invertebrates often differ more
from the surrounding seafloor and/or continental margins (Stocks & Hart 2007, Shank 2010).
New species
Previous studies on Walters Shoal Seamount have led to the discovery of several new and
endemic invertebrate (Kensley 1969, Clark 1972, Kensley 1975, Kensley 1981, Groeneveld
68
et al. 2006) and fish species (Poss & Collette 1990, Collette et al. 1991, Iwamoto et al. 2004).
This study found a relatively high number (~33.3%) of putative new sponge species probably
attributed to the undersampled and underworked state of this group from the Western Indian
Ocean region (Kelly-Borges 1997, Richmond 2001), other seamounts in the Indian Ocean
(Sautya et al. 2011) and Walters Shoal. This is further demonstrated by the fact that many of
the new species include some of the most abundant (Halichondria (Halichondria) sp.,
Rhabderemia sp., Tethya sp.), accessible, and conspicuous (Callyspongia (Callyspongia) sp.,
Clathria (Clathria) sp.) specimens collected during this study. Moreover, this number could
increase following further investigation of those specimens currently only identified to higher
taxonomic levels and/or designated as morphospecies.
Interesting specimens include Latrunculia (Biannulata) sp., Hymerhabdia sp., Chelotropella
sp. and Thrombus sp. The genus Latrunculia is found predominantly in Southern Ocean
waters (Samaai & Kelly 2002) and contains biologically active compounds (e.g. Capon et al.
1987, Duckworth & Battershill 2001), while there are eight species of the genus
Hymerhabdia worldwide, with the species found in this study representing the first record of
this genus in the Indian Ocean (van Soest et al. 2015). The Chelotropella species represents
the third species of this genus documented globally (van Soest et al. 2015), while there are
five species documented from the monogeneric Thrombus, with the current material being the
second species documented from the Indian Ocean (van Soest et al. 2015).
4.2 Location and depth affiliations
As found in other sessile benthic assemblages on seamounts (e.g. Bo et al. 2011, Sautya et al.
2011, Thresher et al. 2014, McClain & Lundsten 2015), the sponge fauna inhabiting these
features often demonstrate significant differences with position on the seamount and depth,
69
often according to local geomorphology and hydrodynamic conditions (Bo et al. 2011).
Examples include studies by Henrich et al. (1992, Vesterisbanken Seamount), Pereira et al.
(2015, Condor Seamount) and Xavier et al. (2015, Schultz Seamount).
Location
The sponge fauna of Walters Shoal Seamount demonstrated no clear patterns in distribution
with regards to location (western vs. eastern flank) with several species shared by both sides.
This can also be seen in other benthic invertebrates, especially in the shallow regions of the
seamount, including the crinoid Comanthus wahlbergi. This is possibly due to the flat (see
Fig. 2), generally homogenous topography of this seamount (Fig. 23), characterised by its
relatively small size, shallow nature, low growth profile and the absence of structural
complexity as noted by Collette & Parin (1991), in addition to local oceanographic
conditions.
Both Nesis (1994) and Gopal (2007) suggest some form of isolation and larvae retainment of
the waters above the seamount, which may possibly be a horizontal tidal current as recorded
by Collette & Parin (1991). Previous reports of upwelling at Walters Shoal (Collette & Parin
1991) were thus not supported by the sponge distributions found in this study. However,
Read & Pollard (2015) found high blocking factors for shallow seamounts within the
Southwest Indian Ocean, which is conducive to Taylor cap formation, whereby water is
trapped over the crest of the seamount. Thus, further sampling on the northern and southern
flank of Walters Shoal is necessary to rule this process out.
70
Depth
The structure and composition of seamount benthic communities is often influenced by
depth, according to environmental gradients (such as temperature and oxygen concentration)
that are associated with this factor (Stocks & Hart 2007, Clark et al. 2010, Consalvey et al.
2010). As expected, sponge assemblages on Walters Shoal Seamount demonstrated a clear
pattern with regards to depth distribution, with each depth zone (shallow: 15 – 30 m,
mesophotic: 31 – 150 m, submesophotic: >150 m) harbouring a distinct sponge assemblage.
Sponge fauna similarities according to the sampling locations in each depth zone, decreased
from the shallow (~35%) to the mesophotic (~21%) and submesophotic (~15%) zones,
probably due to the decreasing number of sampling locations per zone (shallow: five,
mesophotic: four and submesophotic: three). The increasing area and depth range possibly
also plays a role, with the submesophotic zone incorporating all specimens from ~150 –
500 m, while the shallow zone only incorporates those from ~20 – 30 m. In addition, depth
ranges may be species- (or higher taxonomic level) specific, with certain families and genera
dominating a particular depth zone, or well represented throughout.
Species richness and the number of putative new species was highest in the submesophotic
depth zone (approximately double that found in the shallow and mesophotic depth zones),
with 15 putative new species found exclusively in this zone. This is inconsistent with findings
by Samaai et al. (2010), who found sponge species richness to decline with depth, but is
attributed here to the larger area and depth range incorporated in the submesophotic depth
zone as discussed above. Additionally, the lack of work done on the sponge fauna of the
Western Indian Ocean, especially in deeper regions (Kelly-Borges 1997, Richmond 2001)
could explain the higher number of putative new species in the deepest zone.
71
4.3 Biogeographical affiliations
The 26 species that are likely new to science are also possibly endemic to Walters Shoal
Seamount, and thus demonstrate a relatively high level (~33.3%) of endemism. This finding
is consistent with other studies on seamount sponges: Lévi (1969) recorded 53%, Xavier &
van Soest (2007) recorded 28% and Schlacher-Hoenlinger et al. (2005) noted a fauna
dominated by ‘spot endemics’ (species restricted to a single site) from South Pacific
seamounts. As sessile organisms, with larvae that have limited swimming capabilities,
occasional asexual propagation and a relatively short planktonic life (Maldonado 2006,
Mariani et al. 2006), most sponges are found in local or regional areas of endemism (van
Soest et al. 2012), with shallow seamounts possibly constituting centres of endemism for
shallow-water sponges as suggested by Xavier & van Soest (2007), and supported by Lévi’s
(1969) study. This high level of potential endemism could be further attributed to the
somewhat isolated nature of the feature (Groeneveld et al. 2006, Gopal 2007) and the
retentive oceanographic processes found above Walters Shoal (Nesis 1994, Gopal 2007).
Then again, the degree of seamount endemicity has been called into question, with too little
work done on these features and the fauna they support, to use this term with confidence
(McClain 2007). Therefore, the high level of sponge fauna endemism reported here is more
likely indicative of the low sampling effort in this region as mentioned previously, and within
deeper ocean realms as suggested by Samadi et al. (2007).
Collette & Parin (1991) recorded a high level of endemism in the shallow-water fish fauna of
Walters Shoal, with 30 – 40% endemic to some part of the chain of islands and seamounts
within their defined West Wind Drift Islands Province (WWDIP), including Tristan da
Cunha, Gough Island, Vema Seamount, Walters Shoal, UN-2 (unnamed seamount south of
Madagascar) and the St Paul and Amsterdam islands (Nesis 2003). Fewer species (~two) are
endemic to Walters Shoal Seamount alone. This higher level of endemism for benthic
72
seamount invertebrates is consistent with findings from Wilson & Kaufmann (1987), Stocks
& Hart (2007), Xavier & van Soest (2007) and Shank (2010), and is probably due to the
generally more advanced biogeographic and taxonomic knowledge of fish as well as their
mobility, which enables genetic mixing with non-seamount populations (Stocks & Hart
2007).
Based on the 23 known sponge species recorded in this study, Walters Shoal Seamount has
affinities with the Western Indo-Pacific and Temperate Southern African realms and is
comprised of almost equally represented provincial (Western Indian Ocean excluding South
Africa; 35%) and widespread to cosmopolitan (Indian Ocean; 39%) species. These
affiliations, in addition to the range extensions found for several species in this study, indicate
that there is some means of larval dispersal within this region.
There is a deep oceanic trench between Walters Shoal Seamount and the African shelf
(Romanov 2003, Gopal 2007), with sponge larvae probably dispersed via local
oceanographic mechanisms, including currents and eddies. The circulation of the Southwest
Indian Ocean is dominated by the combined eastward flow of the Agulhas Return Current
(ARC) and Subtropical Front (Read & Pollard 2015). However, Walters Shoal lies in a
subtropical gyre north of these flows, in a region of slow mean westward flow between the
southern tip of Madagascar and the ARC and is close to the path of eddies that propagate
southwest from the east of Madagascar (Pollard & Read 2015, Read & Pollard 2015).
Consequently, although previously included in the WWDIP, Walters Shoal is not located
within the West Wind Drift and is bathed by warmer, south-to-southwestwardly flowing
waters from the subtropical branch of the South Equatorial Current (Iwamoto et al. 2004),
demonstrated by the warm surface waters (19 – 23⁰C) recorded by Collette & Parin in 1991.
73
The affiliation of Walters Shoal Seamount with the Western-Indo Pacific (especially the East
African Coral Coast, Seychelles as well as the Western and Northern Madagascar ecoregions)
is probably driven by the train of large anti-cyclonic eddies within the Mozambique Channel,
that transport entrained larvae south (Ridderinkhof et al. 2001, de Ruijter et al. 2002). Larvae
may be further entrained into the Agulhas Current, possibly explaining faunal similarities
with South Africa. Additionally, sponge larvae may be entrained and transported to South
Africa via eddies propagating southwest from the east of Madagascar (Pollard & Read 2015,
Read & Pollard 2015). Eddies have previously been shown to act as strong retention
mesoscale structures that transport larvae and connect marine populations (Landeira et al.
2010).
Overall, Walters Shoal sponge fauna demonstrated a relatively low similarity to surrounding
regions, with no species found to be common to both the seamount and other ecoregions
within the WWDIP, as found for the fish fauna. This may be due to the use of the incomplete
World Porifera Database (van Soest et al. 2015), which is biased according to collection and
taxonomy efforts (van Soest et al. 2012). For example, the database only records 21 sponge
species for the Tristan Gough Ecoregion, 13 for Vema Seamount and eight for the
Amsterdam-St Paul Ecoregion. This is further supported by the finding that within the
Western Indian Ocean Province, Walters Shoal sponge faunal similarities increased
according to the number of species recorded for that ecoregion (i.e. the East African Coral
Coast Ecoregion had the highest sponge faunal affinities with Walters Shoal as well as the
highest number of sponge species recorded at 172).
Thus, the findings of this study regarding the biogeographical affiliations and high potential
endemism of the sponge fauna found on Walters Shoal Seamount should be considered with
caution. Although current, and updated regularly, the World Porifera Database (van Soest et
al. 2015), on which these findings were based, is still incomplete and lacking data, with the
74
retrievable distribution data a minimum of what is known about current species distributions
(R. van Soest, personal communication, February 13, 2015).
The use of this database does provide some insight though, with findings somewhat
consistent with previous work on the fish and cephalopod fauna of this seamount. In addition,
at higher taxonomic levels (including all OTU’s) the sponge fauna was comprised
predominantly of species in the Ancorinidae family, which was consistent with the
surrounding regions that have affiliations with this seamount, with more than half of the
ecoregions having a large representation of this family. Hence, although not conclusive, this
study, in conjunction with the previous work done on the seamount, could act as a basis for
future work, leading to a more thorough understanding of the biogeographical affiliations of
this shallow seamount.
4.4 Study limitations and future work
Key limitations found during this study include the ambiguous definition of Walters Shoal in
the literature, with researchers citing the seamount in some form, but providing different
coordinates as well as the inaccessibility of essential papers. The latter includes Parin et al.
(1993) and Detinova & Sagaidachny (1994), who documented distribution patterns of both
the benthic and water-column fauna of Walters Shoal. As of September (2015),
communications are still underway with T.N. Molodtsova of the P.P. Shirshov Institute of
Oceanology to try gain access to this literature, after correspondence with various other
researchers has been unsuccessful.
Samples were obtained from relatively few sites (including nine epibenthic sleds, two
SCUBA dives and a lobster trap) and further sampling, especially on the northern and
75
southern flank (which were largely neglected), could reveal an even higher diversity of
sponge fauna, or further elucidate sponge assemblage location and depth distributions.
Another issue faced was the relatively small size of sponge specimens obtained, which often
led to difficulties in obtaining enough material for adequate identification and descriptions. In
addition, the lack of work carried out on the sponge fauna of the Western Indian Ocean, and
the resultant state of the largely outdated taxonomic literature, which is in need of extensive
revision (Kelly-Borges 1997, Richmond 2001), often hampered the ability to identify and
describe specimens confidently. As these records are the basis for the (incomplete) World
Porifera Database (van Soest et al. 2015), bias according to collection and taxonomy efforts
(van Soest et al. 2012) was also evident when using these data to further elucidate the
biogeographical affiliations of Walters Shoal. Finally, the lack of work on seamountinhabiting sponges made comparisons of the sponges in this study, and those documented
from other seamount studies, tenuous.
Future work regarding the sponges collected from Walters Shoal Seamount in this study
includes the publication of new species descriptions, with samples from most of the 255
specimens collected for genetic work, in order to confirm current identifications. In addition,
larger scale genetic work needs to be conducted on both the invertebrate and fish fauna of
Walters Shoal, and surrounding non-seamount populations in order to further understand the
biogeography of this seamount, and the role currents and eddies possibly play in larval
dispersal and connectivity. This, in conjunction with further work on the sponge fauna and
oceanographic processes of the Western Indian Ocean region, may also clarify the possibility
or role of this seamount in acting as a stepping stone for species along the Madagascar Ridge,
or further eastwards into the central Indian Ocean. As suggested by van Soest et al. (2012), a
regional approach in the attempt to document sponge fauna and expose distribution patterns
is needed in the Western Indian Ocean region. As such, the taxonomic literature in this region
76
is in need of extensive revision, with the aid of new technologies and accessible resources
(e.g. online sponge identification website and guidebooks). This will also enable a more
robust database (World Porifera Database, van Soest et al. 2015) for use in future work.
4.5 Conclusion
This study has substantially contributed to the knowledge of the sponge fauna from
seamounts within the Indian Ocean, but more specifically, Walters Shoal Seamount. Prior to
this study, Sautya et al. (2011) suggested that there were only reports on ‘Porifera’ and
‘Hexactinellida’ from two Indian Ocean seamounts each in the literature.
Nonetheless, this is only one element of the multidisciplinary cruise launched in May (2014)
as a component of the third phase of the African Coelacanth Ecosystem Programme (ACEP
III). Once additional data from the cruise has been processed, including information on the
invertebrate and fish fauna, as well as the physical and chemical environment of the shoal,
the findings of this study will hopefully contribute to a better understanding of the Walters
Shoal Seamount ecosystem.
77
Figures
Fig. 1: Map showing the location of Walters Shoal Seamount (red star) within the
bathymetric context of the Western Indian Ocean region (generated using QGIS v.2.6.1;
available: qgis.osgeo.org/en/site/).
78
Fig. 2: Bathymetric map of Walters Shoal Seamount (generated using Surfer 9; Golden
Software, available: www.goldensoftware.com).
79
A
B
Fig. 3: Sponge specimen sampling strategies included SCUBA dives (A, © Toufiek Samaai)
and the use of a roughed epibenthic sled, here shown being deployed by the RV Algoa crew
(B, © Robyn Payne).
80
Fig. 4: Bathymetric map of Walters Shoal Seamount (generated using Surfer 9; Golden
Software, available: www.goldensoftware.com), with sled (S), dive (D) and lobster trap (LT)
sampling locations.
81
SAMPLE#
ORGANISM
sponge
ascidian
other
INSTANT ID
IDENTIFICATION
DATE
COLLECTOR
LOCATION
DEPTH
HABITAT
SUBSTRATE
DIMENSIONS thickness
FORM
frilly
colonial
other:
length
thinly encrusting
vase
solitary
thickly encrusting
tube
social
COLOUR exterior
pattern?
fingery projections
branching
stalked
interior
TEXTURE/CONSISTENCY soft
dense
crisp
brittle
other:
COMPRESSIBLE very
TEARS easily so so
SPICULES? no
diameter
change?
medium
barely
hard
BREAKS easily
FIBERS? no
undulating but smooth
slippery
bumpy
no
yes
size
EPI/ENDOFAUNA?
no
yes
describe
MUCOUS/EXUDATE? no
yes
sticky slimey describe
yes
ABUNDANCE rare
in situ
so so
hard
yes
pitted
rough
fuzzy
conulose
distribution
describe
occasional
SAMPLE SIZE
PHOTO
firm
not
OSCULES/SIPHONS
SMELL? no
fingers
bushy
spongy
fibrous
tough
rubbery
cheesy
stony
crunchy
stringy
sandy
falls apart
yes
SURFACE smooth
prickly
sandy
massive
spherical
common abundant
kg
above water
Fig. 5: Sheet completed per sponge specimen to denote macroscopical features (note: this
sheet was filled in as far as possible per specimen, but often several fields were omitted).
82
Fig. 6: Map showing the ecoregions, as defined by Spalding et al. (2007), surrounding
Walters Shoal Seamount (red star) that were included in the biogeographical analyses.
Ecoregions 101 (Bight of Sofala/Swamp Coast) and 217 (Bouvet Island) were excluded as
they had one and zero sponge species recorded by the World Porifera Database (van Soest et
al. 2015) respectively. Vema Seamount is also included for comparison (blue star), due to its
associations with the West Wind Drift Islands Province. Figure generated using QGIS
v.2.6.1, available: qgis.osgeo.org/en/site/.
83
Fig. 7: A – Agelas ceylonica Dendy, 1905. B, C, D – Skeletal architecture. E – Verticillate
acanthostyle I. F – Verticillate acanthostyle II.
84
Fig. 8: A – Ptilocaulis sp. • B, C, D – Skeletal architecture. E – Styles.
85
Fig. 9: A, B – Callyspongia (Callyspongia) sp. • (© Toufiek Samaai). C, D – Skeletal
architecture. E – Oxea.
86
Fig. 10: A, B – Lissodendoryx (Lissodendoryx) pygmaea (Burton, 1931) (© Stephen
Kirkman). C, D – Skeletal architecture. E – Tylote. F – Style. G – Sigma (C-shaped). H –
Isochela I. I – Isochela II.
87
Fig. 11: A – Fibulia ectofibrosa (Lévi, 1963). B, C – Skeletal architecture. D – Oxea. E –
Isochela.
88
Fig. 12: A, B – Latrunculia (Biannulata) sp. • C, D – Skeletal architecture. E – Style. F –
Anisodiscorhabds.
89
Fig. 13: A – Clathria (Clathria) sp. • B, C – Skeletal architecture. D – Style. E, F –
Terminally spined subtylostyle. G – Acanthostyle. H – Toxa I. I, J – Terminally spined Toxa
II. K – Isochela.
90
Fig. 14: A – Halichondria (Halichondria) sp. • B, C – Skeletal architecture. D – Oxea I. E –
Oxea II. F – Oxea III.
91
Fig. 15: A – Aaptos sp. • B, C, D – Skeletal architecture. E – Stronglyoxea. F – Style I. G –
Style II. H – Style III.
92
Fig. 16: A – Tethya sp. • B – Skeletal architecture. C – Anisostrongyloxea. D – Strongyle. E
– Spherasters. F – Tylaster. G – Spheroxyasters.
93
Fig. 17: A – Ancorina sp. • B – Skeletal architecture. C – Oxea I. D – Oxea II. E –
Plagiotriaene I. F – Plagiotriaene II. G – Plagiotriaene III. H – Plagiotriaene extremities. I –
Oxyasters. J – Acanthoxyaster. K – Acanthoxyaster (reduced tetract). L – Sanidaster.
94
Fig. 18: A – Chelotropella sp. • B, C, D – Skeletal architecture. E – Oxea. F – Dichotriaene I.
G – Dichotriaene II. H – Anatriaene. I – Plagiotriaene. J – Calthrop I. K – Calthrop II. L –
Strongyloacanthaster.
95
Fig. 19: A – Penares intermedia (Dendy, 1905). B, C – Skeletal architecture. D – Oxea I. E –
Oxea II. F – Oxea III. G – Dichotriaene I. H – Dichotriaene II. I – Microxea. J –
Acanthoxyaster.
96
Fig. 20: Non-metric MDS ordination of sampling locations according to location (western vs.
eastern flank) on Walters Shoal Seamount, where S = Sled, D = Dive and LT = Lobster Trap.
97
Fig. 21: Non-metric MDS ordination of sampling locations according to depth (Shallow: 15 –
30 m, Mesophotic: 31 – 150 m, Submesophotic: >150 m) on Walters Shoal Seamount, where
S = Sled, D = Dive and LT = Lobster Trap.
98
South Africa
26%
Indian Ocean
39%
Western Indian Ocean
35%
Fig. 22: Biogeographical affinities of the 23 known sponge species recorded from Walters
Shoal Seamount.
99
Fig. 23: Image depicting the low spatial complexity and growth profile of Walters Shoal
Seamount (© Imtiyaaz Malick).
100
Tables
Table 1: Invertebrate (including sponge) collection sampling strategy, where depth zone is denoted
by the symbols S (Shallow: 15 – 30 m), M (Mesophotic: 31 – 150 m) and SM (Submesophotic:
>150 m) according to Lesser et al. (2009).
Date
29/05
30/05
02/06
Grid no.
Station no.
Position
(start)
Position
(end)
Method
(# sponges)
Station
(open/closed)
Duration (min)
Depth (m)
Zone
WSL021
ALG10953
33°11.0' S
43°53.9' E
33°11.0' S
43°52.9' E
Sled 1
4
08:48/09:28
40
53 – 43
M
WSL022
ALG10954
33°10.9' S
43°48.6' E
33°11.2' S
43°50.2' E
Sled 2
31
10:59/11:40
41
170 – 72
M
WSL024
ALG10956
33°08.8' S
43°49.1' E
33°11.2' S
43°50.7' E
33°09.0' S
43°50.5' E
–
Sled 3
55
Dive 12
21
17:05/17:38
33
11:44/12:19
35
348 – 103
SM
29
S
–
–
–
–
33°10.6' S
43°51.0' E
–
Dive 2
9
13:24/14:00
36
29
S
WSL042
ALG10974
WSL043
ALG10975
33°11.2' S
43°51.0' E
33°13.8' S
43°55.5' E
33°11.2' S
43°50.7' E
33°13.1' S
43°55.8' E
Sled 4
40
Sled 5
13
13:15/13:25
10
14:24/14:35
11
34 – 28
S
28 – 30
S
WSL044
ALG10976
33°14.0' S
43°55.5' E
33°13.7' S
43°55.6' E
Sled 6
30
15:05/15:14
9
28 – 25
S
WSL045
ALG10977
WSL046
ALG10978
33°13.8' S
43°56.1' E
33°09.8' S
43°56.6' E
33°14.2' S
43°55.9' E
33°09.8' S
43°56.2' E
Sled 7
7
Sled 8
23
15:34/15:50
16
10:52/11:17
25
80
M
240 – 120
SM
WSL047
ALG10979
–
–
33°09.7' S
43°58.4' E
33°11.6' S
43°50.5' E
33°09.8' S
43°57.0' E
–
Sled 9
14
Lobster Trap
8
11:44/12:34
50
09:40/15:08
328
512 – 317
SM
39
M
03/06
05/06
2
Four divers were present in each dive. The second dive was more focused on the fish fauna of the
seamount.
101
Table 2: Microwave 5mm/2 layer method for sponge specimen histology processing.
Step
Dehydrate
Clean
Dry
Harden for wax
1. Fixation
2. Flushing
3. Rinsing
4. Ethanol
5. Xylene
6. Isopropanol
7. Vaporization
8. Wax Impregnation
Time
(min)
105
2
30
45
90
20
1.5
140
Temperature
(°C)
50
37
45
55
50
60
70
Pressure
(mBar)
600
995 – 150
Agent
70% ethanol
60% ethanol
Absolute alcohol
Absolute alcohol
Xylene
Isopropanol
N/A
N/A
102
Table 3: Ecoregions included in the biogeographical analyses. Categorisation follows Spalding et
al. (2007), with numbers in brackets indicating the number of sponge species recorded in each
ecoregion, compiled from the World Porifera Database (van Soest et al. 2015). Ecoregions 101 and
217 were excluded as they had one and zero sponge species recorded respectively. Vema Seamount
is also included (affiliated with West Wind Drift Islands Province). Last updated May 2015.
Western Indo-Pacific Realm
20. Western Indian Ocean Province
94. Northern Monsoon Current Coast Ecoregion (44)
95. East African Coral Coast Ecoregion (172)
96. Seychelles Ecoregion (147)
97. Cargados Carajos/Tromelin Island Ecoregion (27)
98. Mascarene Islands Ecoregion (35)
99. Southeast Madagascar Ecoregion (4)
100. Western and Northern Madagascar Ecoregion (150)
101. Bight of Sofala/Swamp Coast Ecoregion (1) (excluded)
102. Delagoa Ecoregion (34)
Temperate South America Realm
49. Tristan Gough Province
189. Tristan Gough Ecoregion (21)
Temperate Southern Africa Realm
50. Benguela Province
190. Namib Ecoregion (excluded)
191. Namaqua Ecoregion (138)
51. Agulhas Province
192. Agulhas Bank Ecoregion (131)
193. Natal Ecoregion (101)
52. Amsterdam–St Paul Province
194. Amsterdam-St Paul Ecoregion (8)
Southern Ocean Realm
59. Subantarctic Islands Province
212. Macquarie Island Ecoregion (excluded)
213. Heard and Macdonald Islands Ecoregion (7)
214. Kerguelen Islands Ecoregion (63)
215. Crozet Islands Ecoregion (8)
216. Prince Edward Islands Ecoregion (18)
217. Bouvet Island Ecoregion (0) (excluded)
218. Peter the First Island Ecoregion (excluded)
61. Continental High Antarctic Province
224. East Antarctic Wilkes Land Ecoregion (174)
225. East Antarctic Enderby Land Ecoregion (8)
226. East Antarctic Dronning Maud Land Ecoregion (7)
227. Weddell Sea Ecoregion (71)
228. Amundsen/Bellingshausen Sea Ecoregion (excluded)
229. Ross Sea Ecoregion (excluded)
Other
Vema Seamount (13)
103
Table 4: Sponge species documented from Walters Shoal Seamount per sampling location (S =
Sled, D = Dive, LT = Lobster Trap), where (X) indicates presence and (–) indicates absence. The
symbol (•) denotes all species that are likely new to science.
Species
Aaptos sp. •
Agelas ceylonica
Amorphinopsis (?) sp.
Amorphinopsis cf. fistulosa
Ancorina sp. •
Axinellidae sp.
Axinyssa cf. aplysinoides
Biemna bihamigera
Brachiaster (?) sp.
Bubaridae sp.
Callyspongia (Toxochalina) cf.
robusta
Callyspongia (Callyspongia) sp. •
Chelotropella sp. •
Chondrosia cf. debilis
Clathria (Clathria) sp. •
Clathrinida sp. 1
Clathrinida sp. 2
Desmanthus sp. •
Dictyoceratida sp.
Dictyodendrilla cf. pallasi
Discodermia panoplia
Eurypon sp. 1 •
Eurypon sp. 2 •
Fibulia ectofibrosa
Halichondria (Halichondria) sp. •
Haplosclerida sp. 1
Haplosclerida sp. 2
Haplosclerida sp. 3
Haplosclerida sp. 4
Haplosclerida sp. 5
Hymedesmia (Hymedesmia) sp. •
Hymeniacidon sp. •
Hymerhabdia sp. •
Latrunculia (Biannulata) sp. •
Lissodendoryx (Lissodendoryx)
pygmaea
Microcionidae sp.
Paradesmanthus sp. •
Penares intermedia
S1
–
–
–
–
–
–
–
–
–
–
S2
–
X
–
–
–
–
–
X
–
–
S3
–
X
X
–
–
–
–
X
–
X
S4
X
–
–
X
X
–
–
–
–
–
S5
–
–
–
–
–
–
–
–
–
–
S6
–
–
–
X
–
–
–
–
–
–
S7
–
–
–
–
–
–
–
–
–
–
S8
–
–
–
–
–
–
X
–
–
–
S9
–
–
–
–
–
X
–
–
X
–
D1
–
–
–
–
–
–
–
–
–
–
D2
–
–
–
–
–
–
–
–
–
–
LT
–
–
–
X
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
X
X
X
X
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
X
X
X
X
X
–
–
X
–
–
–
–
–
–
X
–
X
X
X
–
–
–
–
X
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
X
–
X
X
–
–
–
–
–
–
X
X
–
–
X
–
X
–
–
X
–
X
–
–
X
–
–
–
X
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
X
X
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
X
–
X
–
–
–
–
–
–
–
–
–
–
–
104
Species
Phakellia sp. 1 •
Phakellia sp. 2 •
Phakellia sp. 3 •
Phorbas cf. frutex
Poecillastra compressa
Poecilosclerida sp.
Protosuberites sp. 1 •
Protosuberites sp. 2 •
Protosuberites sp. 3 •
Ptilocaulis sp. •
Raspailiidae sp.
Rhabderemia sp. •
Spongosorites sp. •
Stelletta agulhana
Stelletta cf. cylindrica
Stelletta purpurea
Stelletta tulearensis
Stryphnus progressus
Tedania (Tedania) sansibarensis
Tedania (Tedania) tubulifera
Terpios cruciata
Tethya sp. •
Thrombus sp. •
Timea cf. spherastraea
Verongiida sp.
Vulcanella sp. •
Zyzzya fuliginosa
S1
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
S2
–
–
–
X
X
–
–
–
–
–
X
–
X
–
–
–
X
–
–
X
–
X
–
–
X
–
–
S3
X
X
X
–
X
–
X
X
–
X
–
X
–
–
–
–
–
X
–
–
–
–
X
–
–
–
X
S4
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
X
X
–
–
–
–
–
S5
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
S6
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
X
–
–
–
–
–
S7
X
–
–
–
–
–
–
–
–
–
–
–
X
–
–
X
X
–
–
–
–
X
–
–
–
–
–
S8
–
X
–
–
–
X
–
–
X
–
–
X
X
–
X
X
–
–
–
–
–
–
–
X
–
–
–
S9
X
–
–
–
–
–
–
–
X
X
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
D1
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
X
–
–
–
–
–
X
–
–
–
–
–
D2
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
X
–
–
–
–
–
–
–
–
LT
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
X
–
–
–
–
–
Unknowns:
M1
M2
M3
M4
M5
M6
M7
M8
M9
M10
M11
M12
M13
–
–
–
–
–
–
–
–
–
–
–
–
–
X
X
–
–
X
X
–
–
–
X
X
–
X
X
–
–
–
–
–
–
–
–
–
–
–
–
X
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
X
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
–
–
–
105
Table 5: Sponge species documented from Walters Shoal Seamount. Depth range is indicative of sled location depths and thus may not represent the
true species depth range. Dive location depths are indicative of true species depth range. The symbol (•) denotes all species that are likely new to
science. Specimens that could only be identified to a higher taxonomic level (i.e. order, family) are denoted as sp. Unknowns represent morphospecies
that were included in the depth and location analyses but require further investigation for identification. Classification and distribution records follow
the World Porifera Database (van Soest et al. 2015).
No. of specimens
Location
Depth Range (m)
Distribution
1
1
S2
S6
72 – 170
25 – 28
Walters Shoal Seamount
Walters Shoal Seamount
9
S2,S3
72 – 348
Found extensively
throughout the Indian
Ocean
Kingdom Animalia
Phylum Porifera Grant, 1836
Class Calcarea Bowerbank, 1862
Subclass Calcinea Bidder, 1898
Order Clathrinida Hartman, 1958
1. Clathrinida sp. 1
2. Clathrinida sp. 2
Class Demospongiae Sollas, 1885
Subclass Heteroscleromorpha Cárdenas, Perez & Boury–Esnault, 2012
Order Agelasida Hartman, 1980
Family Agelasidae Verrill, 1907
3. Agelas ceylonica Dendy, 1905
Family Hymerhabdiidae Morrow, Picton, Erpenbeck, Boury–Esnault, Maggs & Allcock, 2012
4. Hymerhabdia sp. •
2
S3
103 – 348
Walters Shoal Seamount
Order Axinellida Lévi, 1953
Family Axinellidae Carter, 1875
106
5. Axinellidae sp.
6. Phakellia sp. 1 •
7. Phakellia sp. 2 •
8. Phakellia sp. 3 •
9. Ptilocaulis sp. •
Family Raspailiidae Nardo, 1833
10. Raspailiidae sp.
Subfamily Raspailiinae Nardo, 1833
11. Eurypon sp. 1 •
12. Eurypon sp. 2 •
1
4
4
1
5
S9
S3,S7,S9
S3,S8
S3
S3,S9
317 – 512
80 – 512
103 – 348
103 – 348
103 – 512
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
2
S2
72 – 170
Walters Shoal Seamount
2
1
S4,S6
S9
25 – 34
317 – 512
Walters Shoal Seamount
Walters Shoal Seamount
S2,S3
72 – 348
20
S3,S8,S9
103 – 512
Walters Shoal Seamount
1
S3
103 – 348
Walters Shoal Seamount
2
3
S8
S9
120 – 240
317 – 512
Walters Shoal Seamount
Walters Shoal Seamount
2
2
1
3
1
S4
S5
S5
S6
S8
28 – 34
28 – 30
28 – 30
25 – 28
120 – 240
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
9
10
S4,S6,S7,D1,D2
S1,S2,S4,S6,S8,LT
25 – 80
25 – 240
Walters Shoal Seamount
Manning-Hawkesbury
(Holotype), New
Order Biemnida Morrow, Redmond, Picton, Thacker, Collins, Maggs, Sigwart, Allcock, 2013
Family Biemnidae Hentschel, 1923
13. Biemna bihamigera (Dendy, 1922)
2
Family Rhabderemiidae Topsent, 1928
14. Rhabderemia sp. •
Order Bubarida Morrow & Cárdenas, 2015
Family Bubaridae Topsent, 1894b
15. Bubaridae sp.
Family Desmanthidae Topsent, 1894a
16. Desmanthus sp. •
17. Paradesmanthus sp.•
Order Haplosclerida Topsent, 1928
18. Haplosclerida sp. 1
19. Haplosclerida sp. 2
20. Haplosclerida sp. 3
21. Haplosclerida sp. 4
22. Haplosclerida sp. 5
Family Callyspongiidae de Laubenfels, 1936
23. Callyspongia (Callyspongia) sp. •
24. Callyspongia (Toxochalina) cf. robusta (Ridley, 1884)
Found extensively
throughout the Indian
Ocean
107
Zealand, Chatham
Island, Australia,
Indonesia, Philippines,
Natal, Madagascar,
Kenya
Order Poecilosclerida Topsent, 1928
25. Poecilosclerida sp.
Family Acarnidae Dendy, 1922
26. Zyzzya fuliginosa (Carter, 1879)
Family Coelosphaeridae Dendy, 1922
27. Lissodendoryx (Lissodendoryx) pygmaea (Burton, 1931)
Family Dendoricellidae Hentschel, 1923
28. Fibulia ectofibrosa (Lévi, 1963)
Family Hymedesmiidae Topsent, 1928
29. Hymedesmia (Hymedesmia) sp. •
30. Phorbas cf. frutex Pulitzer–Finali, 1993
Family Latrunculiidae Topsent, 1922
31. Latrunculia (Biannulata) sp. •
Family Microcionidae Carter, 1875
32. Microcionidae sp.
Subfamily Microcioninae Carter, 1875
33. Clathria (Clathria) sp. •
Family Tedaniidae Ridley & Dendy, 1886
34. Tedania (Tedania) sansibarensis Baer, 1906
35. Tedania (Tedania) tubulifera Lévi, 1963
Order Suberitida Chombard & Boury–Esnault, 1999
Family Halichondriidae Gray, 1867
36. Amorphinopsis (?) sp.
37. Amorphinopsis cf. fistulosa (Vacelet, Vasseur & Lévi, 1976)
38. Axinyssa cf. aplysinoides (Dendy, 1922)
1
S8
120 – 240
Walters Shoal Seamount
1
S3
103 – 348
Found extensively
throughout the Indian
Ocean
5
D1
29
KwaZulu-Natal; South
Africa
7
S2,S4
28 – 170
South Africa, Namaqua
1
1
S4
S2
28 – 34
72 – 170
Walters Shoal Seamount
East African Coral
Coast, Kenya
1
S3
103 – 348
Walters Shoal Seamount
1
S8
120 – 240
Walters Shoal Seamount
8
S2,S4,S5,S6
25 – 170
Walters Shoal Seamount
1
1
D2
S2
29
72 – 170
Zanzibar, Tanzania
South Africa, Namaqua
1
4
1
S3
S4,S6,LT
S8
103 – 348
25 – 39
120 – 240
Walters Shoal Seamount
Madagascar
Found extensively
108
39. Halichondria (Halichondria) sp.•
40. Hymeniacidon sp.
41. Spongosorites sp.•
Family Suberitidae Schmidt, 1870
42. Aaptos sp.•
43. Protosuberites sp. 1•
44. Protosuberites sp. 2•
45. Protosuberites sp. 3•
46. Terpios cruciata (Dendy, 1905)
throughout Western
Indian Ocean
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
30
4
3
S4,S5,S6,D1,D2
S3
S2,S7,S8
25 – 34
103 – 348
72 – 240
2
2
1
3
2
S4
S3
S3
S8,S9
S4
28 – 34
103 – 348
103 – 348
120 – 512
28 – 34
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Found extensively
throughout the Indian
Ocean
17
S1,S2,S4,S6,S7,D1,LT
25 – 170
Walters Shoal Seamount
1
S8
120 – 240
East African Coral Coast
Order Tetractinellida Marshall, 1876
Suborder Astrophorina Sollas, 1887
Family Ancorinidae Schmidt, 1870
49. Ancorina sp. •
50. Chelotropella sp. •
51. Stelletta agulhana Lendenfeld, 1907
52. Stelletta cf. cylindrica Thomas, 1973
53. Stelletta purpurea Ridley, 1884
2
1
1
1
17
S4
S2
D1
S8
S4,S6,S7,S8,D1,D2,LT
28 – 34
72 – 170
29
120 – 240
25 – 240
54. Stelletta tulearensis Vacelet, Vasseur & Lévi, 1976
2
S2,S7
72 – 170
55. Stryphnus progressus (Lendenfeld, 1907)
Family Geodiidae Gray, 1867
Subfamily Erylinae Sollas, 1888
56. Penares intermedia (Dendy, 1905)
1
S3
103 – 348
Walters Shoal Seamount
Walters Shoal Seamount
South Africa, Namaqua
Seychelles
Found extensively
throughout Pacific and
Indian Ocean
Madagascar (Holotype),
Kenya
South Africa
10
S2,23
72 – 348
Order Tethyida Morrow & Cárdenas, 2015
Family Tethyidae Gray, 1848
47. Tethya sp. •
Family Timeidae Topsent, 1928
48. Timea cf. spherastraea Burton, 1959
Sri Lanka (Holotype),
109
Zanzibar, Kenya
Family Pachastrellidae Carter, 1875
57. Brachiaster (?) sp.
Family Theonellidae Lendenfeld, 1903
58. Discodermia panoplia Sollas, 1888
1
S9
317 – 512
Walters Shoal Seamount
1
S3
103 – 348
Indonesia (Kai Islands:
Holotype), Madagascar
1
S3
103 – 348
Walters Shoal Seamount
4
S2,S3
72 – 348
1
S9
317 – 512
North Atlantic, West
Africa, South Africa
Walters Shoal Seamount
Order Dendroceratida Minchin, 1900
Family Dictyodendrillidae Bergquist, 1980
62. Dictyodendrilla cf. pallasi (Ridley, 1884)
1
S6
25 – 28
Order Dictyoceratida Minchin, 1900
63. Dictyoceratida sp.
1
S8
120 – 240
Family Thrombidae Sollas, 1888
59. Thrombus sp. •
Family Vulcanellidae Cárdenas, Xavier, Reveillaud, Schander & Rapp, 2011
60. Poecillastra compressa (Bowerbank, 1866)
61. Vulcanella sp. •
Subclass Keratosa Grant, 1861
Seychelles (Holotype),
Falkland Islands,
Antarctic Ocean
Walters Shoal Seamount
Subclass Verongimorpha Erpenbeck, Sutcliffe, De Cook, Dietzel, Maldonado, Van Soest, Hooper, Wörheide, 2012
Order Chondrosiida Boury–Esnault & Lopes, 1985
Family Chondrosiidae Schulze, 1877
64. Chondrosia cf. debilis Thiele, 1900
1
S2
72 – 170
Indonesia, Saudi Arabia,
southern Red Sea,
Madagascar
Order Verongiida Bergquist, 1978
65. Verongiida sp.
1
S2
72 – 170
Walters Shoal Seamount
Unknowns
110
66. M1
67. M2
68. M3
69. M4
70. M5
71. M6
72. M7
73. M8
74. M9
75. M10
76. M11
77. M12
78. M13
6
1
1
1
1
1
2
1
1
1
1
2
1
S2,S3,S4,S7,S8,LT
S2
S4
S8
S2
S2
S5,D2
S9
S5
S2
S2
S5
S2
28 – 348
72 – 170
28 – 34
120 – 240
72 – 170
72 – 170
28 – 30
317 – 512
28 – 30
72 – 170
72 – 170
28 – 30
72 – 170
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
Walters Shoal Seamount
111
Table 6: Spicule dimensions of four Agelas ceylonica Dendy, 1905 specimens from this study, n = 10.
Specimen
TS 2309
Verticillate Acanthostyle I (µm)
190.9 (153.6 – 259.8) x 5.9 (4.3 – 8.3), 13 – 23 whorls
Verticillate Acanthostyle II (µm)
116.9 (84.0 – 140.2) x 5.7 (4.7 – 7.1), 11 – 16 whorls
TS 2313
191.5 (163.9 – 216.5) x 9.0 (6.3 – 11.0), 15 – 22 whorls
115.6 (89.6 – 148.0) x 4.3 (3.1 – 5.5), 12 – 17 whorls
TS 2317
192.5 (159.6 – 259.3) x 7.4 (6.4 – 9.2), 12 – 22 whorls
121.7 (90.2 – 141.5) x 4.9 (4.0 – 5.7), 11 – 16 whorls
TS 2441
229.3 (163.5 – 302.5) x 7.6 (6.1 – 10.1), 14 – 20 whorls
132.7 (117.9 – 146.9) x 5.7 (4.6 – 6.4), 11 – 16 whorls
Table 7: Spicule dimensions of four Ptilocaulis sp. • specimens from this study, n = 10.
Specimen no.
TS 2440
TS 2448
TS 2458
TS 2546
TS 2570
Style (µm)
462.9 – 1332.8 x 18.8 (15.4 – 22.5)
399.1 – 1197.0 x 15.8 (9.4 – 19.0)
364.1 – 1571.8 x 17.6 (13.6 – 22.4)
390.5 – 1285.1 x 17.6 (11.8 – 22.7)
497.6 – 1413.6 x 18.1 (14.2 – 26.9)
Table 8: Spicule dimensions of four Callyspongia (Callyspongia) sp. • specimens from this study, n = 10.
Specimen
TS 2330
TS 2341
TS 2353
TS 2369
Oxea (µm)
69.2 (61.7 – 75.9) x 3.5 (3.0 – 4.2)
64.5 (55.9 – 68.9) x 3.4 (2.6 – 4.0)
66.0 (58.3 – 74.9) x 3.0 (2.4 – 3.7)
62.4 (56.6 – 68.8) x 3.0 (2.3 – 4.2)
112
Table 9: Spicule dimensions of four Lissodendoryx (Lissodendoryx) pygmaea (Burton, 1931) specimens from this study, n = 10.
Specimen
TS 2364
TS 2365
TS 2366
TS 2367
Tylote (µm)
181.5 (164.4 – 196.4) x
Style (µm)
125.6 (116.8 – 137.5) x
Sigma (µm)
28.6
Chela I (µm)
23.8
Chela II (µm)
13.2
4.5 (3.5 – 5.1)
6.0 (4.4 – 6.8)
(26.4 – 31.1)
(22.4 – 25.0)
(12.1 – 14.1)
202.8 (185.0 – 214.3) x
138.3 (131.2 – 143.2) x
28.5
25.0
12.9
4.7 (3.3 – 6.1)
6.2 (5.9 – 6.7)
(23.4 – 32.2)
(22.9 – 26.0)
(11.2 – 14.6)
201.0 (190.2 – 206.9) x
135.5 (128.7 – 141.8) x
28.6
23.8
13.5
4.8 (3.6 – 5.6)
5.3 (4.7 – 6.1)
(27.1 – 30.3)
(20.2 – 25.3)
(12.7 – 14.4)
196.6 (189.3 – 203.4) x
135.4 (125.0 – 140.7) x
29.2
23.5
12.7
4.2 (3.2 – 5.0)
5.6 (4.8 – 6.0)
(26.2 – 32.2)
(22.0 – 25.0)
(11.1 – 13.9)
Table 10: Spicule dimensions of four Fibulia ectofibrosa (Lévi, 1963) specimens from this study, n = 10.
Specimen
TS 2303
Oxea (µm)
336.8 (307.0 – 389.3) x 7.6 (5.4 – 9.8)
Chela (µm)
13.9 (12.7 – 15.4)
TS 2472
316.2 (282.3 – 342.3) x 6.2 (4.2 – 7.7)
13.9 (12.4 – 15.4)
TS 2473
334.1 (281.3 – 386.0) x 6.0 (3.6 – 7.7)
13.5 (12.5 – 14.5)
TS 2477
363.1 (303.1 – 412.8) x 6.8 (3.6 – 9.3)
13.4 (12.3 – 14.5)
113
Table 11: Spicule dimensions of four Clathria (Clathria) sp. • specimens from this study, n = 10.
Specimen
TS 2302
Style (µm)
248.4 (204.2 – 307.1) x
10.9 (8.5 – 13.1)
Subtylostyle (µm)
217.7 (138.9 – 319.1) x
3.3 (2.8 – 3.8)
Acanthostyle (µm)
145.3 (134.8 – 153.6) x
8.2 (4.8 – 9.3)
Toxa I (µm)
144.7
(121.4 – 160.4)
Toxa II (µm)
50.7
(35.0 – 84.3)
Chela (µm)
13.5
(11.8 – 15.0)
TS 2342
234.3 (178.7 – 320.0) x
9.3 (7.9 – 11.5)
211.4 (129.7 – 313.1) x
3.0 (2.4 – 3.8)
138.0 (132.2 – 148.0) x
7.3 (5.6 – 9.7)
146.1
(111.0 – 177.2)
45.1
(35.3 – 61.1)
12.5
(11.2 – 14.2)
TS 2348
247.1 (194.7 – 354.4) x
9.9 (8.3 – 11.8)
218.5 (119.2 – 309.8) x
2.8 (2.0 – 3.8)
139.0 (131.4 – 150.1) x
7.7 (5.3 – 8.9)
143.7
(113.8 – 177.3)
41.3
(27.2 – 55.4)
12.8
(11.6 – 14.1)
TS 2355
228.5 (181.8 – 318.7) x
7.7 (6.1 – 10.0)
230.2 (145.0 – 311.4) x
2.4 (2.1 – 3.0)
139.6 (123.4 – 155.1) x
6.7 (5.3 – 8.4)
113.6
(72.6 – 163.0)
45.7
(33.9 – 62.0)
13.8
(12.9 – 15.1)
Table 12: Spicule dimensions of four Halichondria (Halichondria) sp. • specimens from this study, n = 10.
Specimen
TS 2336
Oxea I (µm)
415.5 (315.6 – 469.1) x
7.9 (6.7 – 9.4)
Oxea II (µm)
239.8 (203.3 – 284.5) x
6.6 (4.4 – 9.6)
Oxea III (µm)
155.0 (113.4 – 173.1) x
6.0 (4.2 – 7.9)
TS 2338
435.5 (377.5 – 513.7) x
11.6 (8.5 – 13.6)
255.7 (209.5 – 287.3) x
8.1 (6.7 – 9.6)
136.8 (114.8 – 161.7) x
5.7 (4.8 – 6.5)
TS 2339
403.3 (349.9 – 461.6) x
9.9 (6.2 – 13.6)
232.0 (208.0 – 288.4) x
7.7 (5.7 – 9.2)
145.3 (112.5 – 198.6) x
6.1 (5.0 – 7.4)
TS 2340
444.2 (418.3 – 474.1) x
10.8 (8.5 – 13.3)
238.4 (211.3 – 274.8) x
7.2 (5.7 – 8.9)
127.2 (120.1 – 142.2) x
5.2 (4.4 – 6.2)
114
Table 13: Spicule dimensions of Aaptos sp. • specimens from this study, n = 10.
Specimen
TS 2502
TS 2503
Stronglyoxea (µm)
954.4 (677.5 – 1284.6) x
Style I (µm)
875.8 (674.1 – 1252.4) x
Style II (µm)
446.0 (348.3 – 576.4) x
Style III (µm)
188.3 (127.5 – 291.1) x
14.1 (7.5 – 20.0)
27.4 (23.6 – 32.3)
14.9 (8.9 – 19.7)
5.0 (3.0 – 6.9)
981.2 (682.8 – 1253.7) x
939.0 (600.9 – 1282.3) x
417.5 (340.1 – 496.4) x
201.5 (117.1 – 291.2) x
21.3 (16.1 – 28.4)
24.9 (17.4 – 35.5)
11.1 (8.1 – 16.1)
5.6 (3.3 – 9.0)
Table 14: Spicule dimensions of four Tethya sp. • specimens from this study, n = 10.
Specimen
TS 2311
Anisostrongyloxea (µm)
285.3 – 1225.0 x
11.1 (3.0 – 20.1)
Strongyle (µm)
733.0 (355.5 – 1124.0) x
14.4 (7.4 – 21.8)
Spheraster (µm)
45.7 (27.5 – 58.5)
Tylaster (µm)
13.9 (12.4 – 15.1)
Spheroxyaster (µm)
6.2 (4.3 – 7.7)
TS 2327
339.1 – 1455.3 x
12.3 (4.2 – 24.3)
1021.3 (522.2 – 1461.7) x
17.4 (12.3 – 23.6)
57.3 (37.8 – 74.8)
11.9 (10.4 – 13.9)
6.7 (4.8 – 10.3)
TS 2337
235.5 – 1306.3 x
10.3 (5.0 – 21.8)
1063.8 (758.1 – 1476.3) x
16.4 (8.3 – 21.4)
51.7 (43.0 – 62.1)
12.5 (10.3 – 14.8)
8.7 (3.7 – 12.2)
TS 2358
292.7 – 1280.1 x
10.5 (5.6 – 22.7)
995.6 (595.6 – 1249.3) x
19.2 (9.0 – 24.9)
37.0 (21.3 – 56.0)
12.6 (10.5 – 15.1)
6.2 (5.3 – 7.0)
115
Table 15: Spicule dimensions of Ancorina sp. • specimens from this study, n = 10.
Plagiotriaene I
Spicule Type
Oxea I (µm)
TS 2475
1748.4 (1276.7 – 2017.8) x 30.3 (16.9 – 36.8)
TS 2476
1764.9 (1521.0 – 2063.3) x 29.9 (20.6 – 39.2)
Oxea II (µm)
975.5 (727.5 – 1133.7) x 9.1 (6.3 – 12.5)
933.6 (801.1 – 1090.6) x 8.3 (5.5 – 11.8)
1759.8 (1550.3 – 2074.9) x 38.5 (33.4 – 46.5)
1668.9 (1215.6 – 1965.4) x 35.9 (25.0 – 46.7)
152.3 (130.3 – 175.0)
140.0 (84.5 – 190.0)
89.9 (75.1 – 116.9)
85.6 (54.4 – 113.3)
976.3 (924.1 – 1037.1) x 19.9 (16.6 – 23.8)
973.4 (849.5 – 1070.9) x 19.0 (13.5 – 26.0)
Cladome (µm)
65.5 (51.5 – 84.4)
57.7 (38.1 – 78.3)
Cladi (µm)
29.8 (18.8 – 38.2)
28.7 (14.9 – 61.4)
608.2 (457.8 – 766.9) x 10.8 (6.1 – 18.4)
564.3 (419.6 – 630.5) x 8.0 (5.7 – 15.2)
Cladome (µm)
31.8 (19.6 – 53.4)
23.3 (16.3 – 28.4)
Cladi (µm)
13.9 (8.4 – 24.0)
12.4 (6.7 – 14.9)
Oxyaster (µm)
10.9 (8.5 – 14.6)
11.8 (9.2 – 14.2)
Acanthoxyaster I (µm)
18.2 (15.7 – 22.1)
21.3 (17.1 – 27.9)
Acanthoxyaster II (µm)
19.2 (14.6 – 23.5)
22.5 (15.0 – 27.8)
5.9 (5.2 – 6.8)
5.7 (5.0 – 6.5)
Rhabdome (µm)
Cladome (µm)
Cladi (µm)
Plagiotriaene II
Plagiotriaene III
Rhabdome (µm)
Rhabdome (µm)
Sanidaster (µm)
116
Table 16: Spicule dimensions of four Penares intermedia (Dendy, 1905) specimens from this study, n = 10 unless otherwise stated.
Dichotriaene I
Dichotriaene II
Spicule Type
Oxea I (µm)
TS 2300
697.8 (603.9 – 758.5) x
16.9 (11.8 – 21.3)
TS 2307
727.3 (637.9 – 806.7) x
17.0 (14.2 – 20.6)
TS 2445
699.6 (613.8 – 925.5) x
18.4 (13.2 – 27.2)
TS 2447
840.4 (703.0 – 999.1) x
27.4 (21.8 – 36.8)
Oxea II (µm)
359.3 (266.6 – 587.2) x
14.0 (10.4 – 20.8)
354.9 (223.0 – 546.1) x
13.9 (9.3 – 16.7)
311.4 (271.4 – 414.5) x
16.8 (14.7 – 20.2)
408.6 (318.2 – 505.7) x
18.7 (14.9 – 21.9)
Oxea III (µm)
134.8 (103.4 – 197.2) x
9.3 (6.6 – 13.2)
139.8 (108.1 – 185.7) x
9.1 (6.7 – 11.4)
142.7 (102.9 – 195.4) x
10.8 (8.1 – 13.3)
140.7 (117.9 – 164.3) x
9.8 (7.1 – 12.3)
193.2 x 30.0, n = 1
133.5 (49.1 – 200.9) x
29.0 (20.1 – 38.8), n = 6
194.5 x 23.9, n = 1
None seen
Cladome (µm)
462.3 (460.7 – 463.9), n = 2
457.8 (338.7 – 540.4)
484.8 (422.3 – 528.2), n = 6
487.1 (380.8 – 578.8)
Protoclad (µm)
103.6 (101.1 – 106.1) x
25.9 (24.5 – 27.2), n = 2
114.5 (97.1 – 139.3) x
30.9 (25.5 – 36.7)
102.3 (91.5 – 119.2) x
36.2 (33.5 – 40.1), n = 6
90.6 (68.4 – 113.3) x
37.8 (27.5 – 48.6)
Deuteroclad (µm)
144.8 (138.8 – 150.7) x
25.1 (24.6 – 25.6), n = 2
121.9 (88.1 – 151.6) x
25.1 (20.7 – 31.0)
153.2 (119.4 – 196.7) x
29.2 (24.1 – 33.7), n = 6
145.3 (115.9 – 178.2) x
30.2 (20.1 – 38.7)
Rhabdome (µm)
82.3 (74.7 – 89.9) x
25.0 (19.1 – 30.8), n = 2
88.8 (39.6 – 132.8) x
23.4 (13.9 – 31.4), n = 7
66.7 (30.7 – 122.1) x
20.6 (14.4 – 30.0), n = 5
None seen
Cladome (µm)
341.8 (316.5 – 384.0), n = 6
349.9 (258.5 – 481.0)
329.8 (281.8 – 380.7), n = 7
325.1 (226.3 – 478.4)
Protoclad (µm)
103.5 (78.3 – 124.8) x
19.6 (15.6 – 27.5), n = 6
119.3 (92.5 – 138.2) x
17.9 (10.4 – 25.6)
94.1 (79.4 – 108.3) x
19.6 (15.2 – 24.6), n = 7
89.0 (74.5 – 102.8) x
21.0 (14.2 – 26.9)
Deuteroclad (µm)
76.0 (46.7 – 87.2) x
14.8 (10.4 – 18.7), n = 6
65.1 (31.7 – 101.4) x
13.7 (7.5 – 18.9)
80.9 (43.4 – 121.8) x
14.7 (11.9 – 17.8), n = 7
69.0 (28.3 – 122.8) x
15.1 (7.2 – 18.5)
76.1 (63.1 – 86.6) x
6.6 (5.3 – 8.0)
79.6 (62.7 – 99.1) x
5.8 (5.2 – 6.6)
83.1 (73.8 – 99.0) x
7.4 (5.9 – 9.3)
75.1 (62.6 – 92.1) x
6.0 (5.2 – 7.0)
Rhabdome (µm)
Microxea (µm)
117
Table 17: Walters Shoal Seamount sponge species list per location. The symbol (•) denotes all species that are likely new to science. When orange,
this symbol denotes the new species found only at that respective location.
Western Flank
Aaptos sp. •
Agelas ceylonica
Amorphinopsis (?) sp.
Amorphinopsis cf. fistulosa
Ancorina sp. •
Biemna bihamigera
Bubaridae sp.
Callyspongia (Toxochalina) cf. robusta
Callyspongia (Callyspongia) sp. •
Chelotropella sp. •
Chondrosia cf. debilis
Clathria (Clathria) sp. •
Clathrinida sp. 1
Discodermia panoplia
Eurypon sp. 1 •
Fibulia ectofibrosa
Halichondria (Halichondria) sp. •
Haplosclerida sp. 1
Hymedesmia (Hymedesmia) sp. •
Hymeniacidon sp. •
Hymerhabdia sp. •
Latrunculia (Biannulata) sp. •
Lissodendoryx (Lissodendoryx) pygmaea
Penares intermedia
Phakellia sp. 1 •
Phakellia sp. 2 •
Middle
Callyspongia (Toxochalina) cf. robusta
Tethya sp. •
Eastern Flank
Amorphinopsis cf. fistulosa
Axinellidae sp.
Axinyssa cf. aplysinoides
Brachiaster (?) sp.
Callyspongia (Toxochalina) cf. robusta
Callyspongia (Callyspongia) sp. •
Clathria (Clathria) sp. •
Clathrinida sp. 2
Desmanthus sp. •
Dictyoceratida sp.
Dictyodendrilla cf. pallasi
Eurypon sp. 1 •
Eurypon sp. 2 •
Halichondria (Halichondria) sp. •
Haplosclerida sp. 2
Haplosclerida sp. 3
Haplosclerida sp. 4
Haplosclerida sp. 5
Microcionidae sp.
Paradesmanthus sp. •
Phakellia sp. 1 •
Phakellia sp. 2 •
Poecilosclerida sp.
Protosuberites sp. 3 •
Ptilocaulis sp. •
Rhabderemia sp. •
118
Phakellia sp. 3 •
Phorbas cf. frutex
Poecillastra compressa
Protosuberites sp. 1 •
Protosuberites sp. 2 •
Ptilocaulis sp. •
Raspailiidae sp.
Rhabderemia sp. •
Spongosorites sp. •
Stelletta agulhana
Stelletta purpurea
Stelletta tulearensis
Stryphnus progressus
Tedania (Tedania) sansibarensis
Tedania (Tedania) tubulifera
Terpios cruciata
Tethya sp. •
Thrombus sp. •
Verongiida sp.
Zyzzya fuliginosa
M1, M2, M3, M5, M6, M7, M10, M11, M13
# species: 55
# new species: 21
# new species only found at this location: 11
Spongosorites sp. •
Stelletta cf. cylindrica
Stelletta purpurea
Stelletta tulearensis
Tethya sp. •
Timea cf. spherastraea
Vulcanella sp. •
M1, M4, M7, M8, M9, M12
# species: 2
# new species: 1
# new species only found at this location: 0
# species: 39
# new species: 15
# new species only found at this location: 5
119
Table 18: SIMPER results – percentage contribution of each species that overall contribute to at
least 60% of the difference between the western and eastern flank of Walters Shoal Seamount.
Average dissimilarity between the western and eastern flank of the seamount is ~68%.
Species
Callyspongia (Callyspongia) sp.
Clathria (Clathria) sp.
Callyspongia (Toxochalina) cf. robusta (Ridley, 1884)
Stelletta purpurea Ridley, 1884
Amorphinopsis cf. fistulosa (Vacelet, Vasseur & Lévi, 1976)
M9
Haplosclerida sp. 2
Haplosclerida sp. 3
M12
Tethya sp.
M7
Eurypon sp. 1
Clathrinida sp. 2
Dictyodendrilla cf. pallasi (Ridley, 1884)
Haplosclerida sp. 4
Tedania (Tedania) sansibarensis Baer, 1906
Phakellia sp. 1
Lissodendoryx (Lissodendoryx) pygmaea (Burton, 1931)
Stelletta agulhana Lendenfeld, 1907
Fibulia ectofibrosa (Lévi, 1963)
M1
Spongosorites sp.
% Contribution
4,7
4,62
4,53
3,88
3,66
3
3
3
3
2,69
2,69
2,43
2,33
2,33
2,33
2,15
2,1
2
2
1,66
1,66
1,63
120
Table 19: Walters Shoal Seamount sponge species list per depth zone. The symbol (•) denotes all species that are likely new to science. When orange,
this symbol denotes the new species found only in that respective depth zone.
Shallow (15 – 30 m)
Aaptos sp. •
Amorphinopsis cf. fistulosa
Ancorina sp. •
Callyspongia (Toxochalina) cf. robusta
Callyspongia (Callyspongia) sp. •
Clathria (Clathria) sp. •
Clathrinida sp. 2
Dictyodendrilla cf. pallasi
Eurypon sp. 1 •
Fibulia ectofibrosa
Halichondria (Halichondria) sp. •
Haplosclerida sp. 1
Haplosclerida sp. 2
Haplosclerida sp. 3
Haplosclerida sp. 4
Hymedesmia (Hymedesmia) sp. •
Lissodendoryx (Lissodendoryx) pygmaea
Stelletta agulhana
Stelletta purpurea
Tedania (Tedania) sansibarensis
Terpios cruciata
Tethya sp. •
M1, M3, M7, M9, M12
Mesophotic (31 – 150 m)
Agelas ceylonica
Amorphinopsis cf. fistulosa
Biemna bihamigera
Callyspongia (Toxochalina) cf. robusta
Callyspongia (Callyspongia) sp. •
Chelotropella sp. •
Chondrosia cf. debilis
Clathria (Clathria) sp. •
Clathrinida sp. 1
Fibulia ectofibrosa
Penares intermedia
Phakellia sp. 1 •
Phorbas cf. frutex
Poecillastra compressa
Raspailiidae sp.
Spongosorites sp. •
Stelletta purpurea
Stelletta tulearensis
Tedania (Tedania) tubulifera
Tethya sp. •
Verongiida sp.
M1, M2, M5, M6, M10, M11, M13
Submesophotic (>150 m)
Agelas ceylonica
Amorphinopsis (?) sp.
Axinellidae sp.
Axinyssa cf. aplysinoides
Biemna bihamigera
Brachiaster (?) sp.
Bubaridae sp.
Callyspongia (Toxochalina) cf. robusta
Desmanthus sp. •
Dictyoceratida sp.
Discodermia panoplia
Eurypon sp. 2 •
Haplosclerida sp. 5
Hymeniacidon sp. •
Hymerhabdia sp. •
Latrunculia (Biannulata) sp. •
Microcionidae sp.
Paradesmanthus sp. •
Penares intermedia
Phakellia sp. 1 •
Phakellia sp. 2 •
Phakellia sp. 3 •
Poecillastra compressa
Poecilosclerida sp.
Protosuberites sp. 1 •
Protosuberites sp. 2 •
121
Protosuberites sp. 3 •
Ptilocaulis sp. •
Rhabderemia sp. •
Spongosorites sp. •
Stelletta cf. cylindrica
Stelletta purpurea
Stryphnus progressus
Thrombus sp. •
Timea cf. spherastraea
Vulcanella sp. •
Zyzzya fuliginosa
M1, M4, M8
# species: 27
#new species: 8
#new species only found in this depth zone: 5
# species: 28
#new species: 6
#new species only found in this depth zone: 1
#species: 40
#new species: 17
#new species only found in this depth zone: 15
122
Table 20: SIMPER results – species that contribute to 90% (100% in submesophotic zone) of
sampling location similarity in each depth zone (Shallow: 15 – 30 m, Mesophotic: 31 – 150 m,
Submesophotic: >150 m). Average sponge faunal similarity of each depth zone is given in brackets.
Shallow
Mesophotic
Submesophotic
Species
(~35%)
(~21%)
(~15%)
Halichondria (Halichondria) sp.
33.81%
Callyspongia (Callyspongia) sp.
25.97%
Stelletta purpurea Ridley, 1884
25.97 %
Clathria (Clathria) sp.
7.84%
Callyspongia (Toxochalina) cf. robusta (Ridley, 1884)
33.33%
Tethya sp.
33.33%
M1
33.33%
Rhabderemia sp.
50.00%
Protosuberites sp. 3
50.00%
123
Table 21: Walters Shoal Seamount sponges – percent contribution of higher taxonomic levels
(families and genera) per depth zone.
Depth Zone
Shallow
(15 – 30 m)
Families
Ancorinidae (17.6%), Callyspongiidae (11.8%),
Halichondriidae (11.8%), Suberitidae (11.8%),
Coelosphaeridae (5.9%), Dictyodendrillidae
(5.9%), Hymedesmiidae (5.9%), Isodictyidae
(5.9%), Microcionidae (5.9%), Raspailiidae
(5.9%), Tedaniidae (5.9%), Tethyidae (5.9%)
Mesophotic
(31 – 150 m)
Ancorinidae (15.8%), Callyspongiidae (10.5%),
Halichondriidae (10.5%), Agelasidae (5.3%),
Axinellidae (5.3%), Biemnidae (5.3%),
Chondrosiidae (5.3%), Geodiidae (5.3%),
Hymedesmiidae (5.3%), Isodictyidae (5.3%),
Microcionidae (5.3%), Raspailiidae (5.3%),
Tedaniidae (5.3%), Tethyidae (5.3%),
Vulcanellidae (5.3%)
Submesophotic
( > 150 m)
Axinellidae (14.7%), Halichondriidae (11.8%),
Ancorinidae (8.8%), Suberitidae (8.8%),
Desmanthidae (5.9%), Vulcanellidae (5.9%),
Acarnidae (2.9%), Agelasidae (2.9%),
Biemnidae (2.9%), Bubaridae (2.9%),
Callyspongiidae (2.9%), Geodiidae (2.9%),
Hymerhabdiidae (2.9%), Latrunculiidae (2.9%),
Microcionidae (2.9%), Pachastrellidae (2.9%),
Raspailiidae (2.9%), Rhabderemiidae (2.9%),
Theonellidae (2.9%), Thrombidae (2.9%),
Timeidae (2.9%)
Genera
Callyspongia (11.8%), Stelletta (11.8%),
Aaptos (5.9%), Amorphinopsis (5.9%),
Ancorina (5.9%), Clathria (5.9%),
Dictyodendrilla (5.9%), Eurypon
(5.9%), Fibulia (5.9%), Halichondria
(5.9%), Hymedesmia (5.9%),
Lissodendoryx (5.9%), Tedania (5.9%),
Terpios (5.9%), Tethya (5.9%)
Callyspongia (11.1%), Stelletta (11.1%),
Agelas (5.6%), Amorphinopsis (5.6%),
Biemna (5.6%), Chelotropella (5.6%),
Chondrosia (5.6%), Clathria (5.6%),
Fibulia (5.6%), Penares (5.6%),
Phakellia (5.6%), Phorbas (5.6%),
Poecillastra (5.6%), Spongosorites
(5.6%), Tedania (5.6%), Tethya (5.6%)
Phakellia (9.7%), Protosuberites
(9.7%), Stelletta (6.5%), Agelas (3.2%),
Amorphinopsis (3.2%), Axinyssa (3.2%),
Biemna (3.2%), Brachiaster (3.2%),
Callyspongia (3.2%), Desmanthus
(3.2%), Discodermia (3.2%), Eurypon
(3.2%), Hymeniacidon (3.2%),
Hymerhabdia (3.2%), Latrunculia
(3.2%), Paradesmanthus (3.2%),
Penares (3.2%), Poecillastra (3.2%),
Ptilocaulis (3.2%), Rhabderemia
(3.2%), Spongosorites (3.2%),
Stryphnus (3.2%), Thrombus (3.2%),
Timea (3.2%), Vulcanella (3.2%),
Zyzzya (3.2%)
124
Table 22: Walters Shoal Seamount – sponge families per depth zone (Shallow: 15 – 30 m,
Mesophotic: 31 – 150 m, Submesophotic: >150 m), where (X) indicates presence and (–) indicates
absence.
Family
Acarnidae
Agelasidae
Ancorinidae
Axinellidae
Biemnidae
Bubaridae
Callyspongiidae
Chondrosiidae
Coelosphaeridae
Desmanthidae
Dictyodendrillidae
Geodiidae
Halichondriidae
Hymedesmiidae
Hymerhabdiidae
Isodictyidae
Latrunculiidae
Microcionidae
Pachastrellidae
Raspailiidae
Rhabderemiidae
Suberitidae
Tedaniidae
Tethyidae
Theonellidae
Thrombidae
Timeidae
Vulcanellidae
Shallow
–
–
X
–
–
–
X
–
X
–
X
–
X
X
–
X
–
X
–
X
–
X
X
X
–
–
–
–
Mesophotic
–
X
X
X
X
–
X
X
–
–
–
X
X
X
–
X
–
X
–
X
–
–
X
X
–
–
–
X
Submesophotic
X
X
X
X
X
X
X
–
–
X
–
X
X
–
X
–
X
X
X
X
X
X
–
–
X
X
X
X
125
Table 23: Walters Shoal Seamount – sponge genera per depth zone (Shallow: 15 – 30 m,
Mesophotic: 31 – 150 m, Submesophotic: >150 m), where (X) indicates presence and (–) indicates
absence.
Genus
Aaptos
Agelas
Amorphinopsis
Ancorina
Axinyssa
Biemna
Brachiaster
Callyspongia
Chelotropella
Chondrosia
Clathria
Desmanthus
Dictyodendrilla
Discodermia
Eurypon
Fibulia
Halichondria
Hymedesmia
Hymeniacidon
Hymerhabdia
Latrunculia
Lissodendoryx
Paradesmanthus
Penares
Phakellia
Phorbas
Poecillastra
Protosuberites
Ptilocaulis
Rhabderemia
Spongosorites
Stelletta
Stryphnus
Tedania
Terpios
Tethya
Thrombus
Timea
Vulcanella
Zyzzya
Shallow
X
–
X
X
–
–
–
X
–
–
X
–
X
–
X
X
X
X
–
–
–
X
–
–
–
–
–
–
–
–
–
X
–
X
X
X
–
–
–
–
Mesophotic
–
X
X
–
–
X
–
X
X
X
X
–
–
–
–
X
–
–
–
–
–
–
–
X
X
X
X
–
–
–
X
X
–
X
–
X
–
–
–
–
Submesophotic
–
X
X
–
X
X
X
X
–
–
–
X
–
X
X
–
–
–
X
X
X
–
X
X
X
–
X
X
X
X
X
X
X
–
–
–
X
X
X
X
126
Table 24: SIMPER results – percentage contribution (bold) of each species that overall contribute
to at least 60% of the difference between depth zones (Shallow: 15 – 30 m, Mesophotic: 31 –
150 m, Submesophotic: >150 m). Average dissimilarities between depth zones given in brackets.
Shallow & Mesophotic (~79%)
Mesophotic & Submesophotic (~83%)
Shallow & Submesophotic (~97%)
Halichondria (Halichondria) sp.
7,26
Callyspongia (Callyspongia) sp.
6,38
M1
6,11
Callyspongia (Toxochalina) cf. robusta
4,98
Amorphinopsis cf. fistulosa
4,3
Clathria (Clathria) sp.
3,84
Spongosorites sp.
3,48
Stelletta tulearensis
3,48
Tethya sp.
3,26
M7
3,26
Stelletta purpurea
2,6
Tedania (Tedania) sansibarensis
2,13
Rhabderemia sp.
4,91
Tethya sp.
4,91
Stelletta tulearensis
3,75
Ptilocaulis sp.
3,6
Callyspongia (Toxochalina) cf. robusta
3,14
Phakellia sp. 1
3,14
Phakellia sp. 2
3,14
Eurypon sp. 1
2,03
Phakellia sp. 1
2,01
Lissodendoryx (Lissodendoryx)
pygmaea
1,97
Stelletta agulhana
1,97
Fibulia ectofibrosa
1,83
Discodermia panoplia
1,83
Hymeniacidon sp.
1,83
Halichondria (Halichondria) sp.
4,26
Rhabderemia sp.
4,26
Clathria (Clathria) sp.
3,15
Phakellia sp. 1
3,07
Ptilocaulis sp.
3,07
Stelletta purpurea
2,85
Callyspongia (Callyspongia) sp.
2,75
Protosuberites sp. 3
2,75
Phakellia sp. 2
2,69
M1
2,3
Tethya sp.
2,19
M7
2,07
Callyspongia (Toxochalina) cf.
robusta
1,74
Amorphinopsis cf. fistulosa
1,64
Hymerhabdia sp.
1,83
Latrunculia (Biannulata) sp.
1,83
Phakellia sp. 3
1,83
Protosuberites sp. 1
1,83
Protosuberites sp. 2
1,83
Stryphnus progressus
1,83
Thrombus sp.
1,83
Zyzzya fuliginosa
1,83
Eurypon sp. 1
1,64
Brachiaster (?) sp.
1,57
Eurypon sp. 2
1,57
Paradesmanthus sp.
1,57
Vulcanella sp.
1,57
M8
1,57
Axinellidae sp.
1,57
Agelas ceylonica
1,51
Callyspongia (Callyspongia) sp. 3,08
Protosuberites sp. 3
3,08
Stelletta purpurea
2,93
Spongosorites sp.
2,44
Bubaridae sp.
1,83
127
Amorphinopsis (?) sp.
1,83
Biemna bihamigera
1,51
Bubaridae sp.
1,51
Discodermia panoplia
1,51
Hymeniacidon sp.
1,51
Hymerhabdia sp.
1,51
Latrunculia (Biannulata) sp.
1,51
128
Table 25: Biogeographical affinities of the Walters Shoal Seamount sponge fauna based on the 23 known species from this study. Categorisation follows
Spalding et al. (2007). Abbreviations: IO = Indian Ocean, WIO = Western Indian Ocean, SA = South Africa, NMCC = Northern Monsoon Current Coast,
EACC = East African Coral Coast, SEY = Seychelles, CCTI = Cargados Carajos/Tromelin Island, MAS = Mascarene Islands, WANM = Western and Northern
Madagascar, DEL = Delagoa, NAM = Namaqua, AGU = Agulhas Bank and NAT = Natal. The symbol (X) indicates presence, while (–) indicates absence.
Species
Agelas ceylonica Dendy, 1905
Amorphinopsis fistulosa (Vacelet, Vasseur & Lévi,
1976)
Axinyssa aplysinoides (Dendy, 1922)
Biemna bihamigera (Dendy, 1922)
Callyspongia (Toxochalina) robusta (Ridley, 1884)
Chondrosia debilis Thiele, 1900
Dictyodendrilla pallasi (Ridley, 1884)
Discodermia panoplia Sollas, 1888
Fibulia ectofibrosa (Lévi, 1963)
Lissodendoryx (Lissodendoryx) pygmaea (Burton, 1931)
Penares intermedia (Dendy, 1905)
Phorbas frutex Pulitzer–Finali, 1993
Poecillastra compressa (Bowerbank, 1866)
Stelletta agulhana Lendenfeld, 1907
Stelletta cylindrica Thomas, 1973
Stelletta purpurea Ridley, 1884
Stelletta tulearensis Vacelet, Vasseur & Lévi, 1976
Stryphnus progressus (Lendenfeld, 1907)
Tedania (Tedania) sansibarensis Baer, 1906
Tedania (Tedania) tubulifera Lévi, 1963
Terpios cruciata (Dendy, 1905)
Timea spherastraea Burton, 1959
Zyzzya fuliginosa (Carter, 1879)
Similarity (shared species)
Absolute number
Percentage (%)
Western Indo-Pacific Realm
Western Indian Ocean Province
Affinity NMCC EACC SEY CCTI MAS WANM DEL
–
–
X
–
–
–
–
IO
–
–
–
–
–
X
–
WIO
–
X
X
X
–
X
–
WIO
–
X
X
–
–
X
–
IO
–
X
–
–
–
X
–
IO
–
–
–
–
–
X
–
IO
–
–
X
–
–
–
–
WIO
–
–
–
–
–
X
–
IO
–
–
–
–
–
–
–
SA
–
–
–
–
–
–
–
SA
X
X
–
–
–
–
–
IO
–
X
–
–
–
–
–
WIO
–
–
–
–
–
–
–
SA
–
–
–
–
–
–
–
SA
–
–
X
–
–
–
–
WIO
X
X
X
–
X
–
X
IO
–
X
–
–
–
X
–
WIO
–
–
–
–
–
–
–
SA
–
X
–
–
–
–
–
WIO
–
–
–
–
–
–
–
SA
–
–
X
–
–
–
X
IO
–
X
–
–
–
–
–
WIO
–
X
X
–
–
X
–
IO
2
2,6
10
12,8
8
10,3
1
1,3
1
1,3
8
10,3
2
2,6
Temperate Southern Africa Realm
Benguela Province
Agulhas Province
NAM
AGU
NAT
–
–
–
–
–
–
–
–
–
–
–
–
–
–
X
–
–
–
–
–
–
–
–
–
X
X
–
–
–
X
–
–
–
–
–
–
X
–
–
X
X
X
–
–
–
–
–
X
–
–
–
–
X
–
–
–
–
X
X
–
–
–
–
–
–
–
–
–
–
4
5,1
4
5,1
4
5,1
129
Table 26: The most represented sponge families and genera per ecoregion that was found to have
biogeographical affiliations with Walters Shoal Seamount. Categorisation follows Spalding et al.
(2007), with numbers in brackets indicating the number of sponge species recorded in each
ecoregion, compiled from the World Porifera Database (van Soest et al. 2015). Ecoregion 101 was
excluded as it had only one sponge species recorded. Last updated May 2015.
Western Indo-Pacific Realm
20. Western Indian Ocean Province
94. Northern Monsoon Current Coast Ecoregion
(44)
95. East African Coral Coast Ecoregion
(172)
96. Seychelles Ecoregion
(147)
97. Cargados Carajos/Tromelin Island Ecoregion
(27)
98. Mascarene Islands Ecoregion
(35)
99. Southeast Madagascar Ecoregion
(4)
100. Western and Northern Madagascar Ecoregion
(150)
101. Bight of Sofala/Swamp Coast Ecoregion
(1)
102. Delagoa Ecoregion
(34)
Temperate Southern Africa Realm
50. Benguela Province
190. Namib Ecoregion
191. Namaqua Ecoregion
(138)
51. Agulhas Province
192. Agulhas Bank Ecoregion
(131)
193. Natal Ecoregion
Families
Ancorinidae (13.6%),
Phloeodictyidae (11.4%),
Raspailiidae (11.4%)
Chalinidae (8.1%),
Halichondriidae (7.0%),
Ancorinidae (4.7%),
Axinellidae (4.7%),
Callyspongiidae (4.7%)
Ancorinidae (8.2%),
Acarnidae (5.4%),
Halichondriidae (4.8%),
Microcionidae (4.8%)
Axinellidae (11.1%),
Microcionidae (11.1%)
Raspailiidae (11.4%),
Spongiidae (11.4%),
Grantiidae (8.6%)
Geodiidae (75.0%),
Spongiidae (25.0%)
Chalinidae (6.0%),
Microcionidae (5.3%),
Ancorinidae (4.7%),
Mycalidae (4.7%)
Excluded
Ancorinidae (17.6%),
Axinellidae (11.8%),
Microcionidae (11.8)
Excluded
Microcionidae (15.2%),
Mycalidae (8.7%)
Geodiidae (7.6%),
Grantiidae (7.6%),
Latrunculiidae (6.9%)
Ancorinidae (13.9%),
Genera
Hemiasterella (6.8%),
Higginsia (6.8%),
Oceanapia (6.8%),
Xestospongia (6.8%)
Haliclona (7.6%),
Callyspongia (4.7%),
Biemna (3.5%),
Mycale (3.5%)
Clathria (4.8%),
Biemna (3.4%),
Rhabdastrella (3.4%),
Tethya (3.4%)
Clathria (11.1%),
Dragmacidon (7.4%)
Leucandra (8.6%),
Dysidea (5.7%),
Spongia (5.7%),
Stelletta (5.7%)
Geodia (75.0%),
Spongia (25.0%)
Haliclona (6.0%),
Clathria (5.3%),
Mycale (4.7%)
Clathria (8.8%),
Stelletta (8.8%)
Clathria (12.3%),
Mycale (8.7%),
Haliclona (5.8%),
Isodictya (5.8%)
Clathria (5.3%),
Isodictya (5.3%),
Leucandra (5.3%)
Clathria (8.9%),
130
(101)
Other
Walters Shoal Seamount
Geodiidae (9.9%),
Microcionidae (8.9%)
Geodia (6.9%),
Stelletta (6.9%)
Ancorinidae (12.7%),
Halichondriidae (10.9%),
Axinellidae (9.1%),
Suberitidae (9.1%)
Stelletta (7.8%),
Phakellia (5.9%),
Protosuberites (5.9%)
131
References
ALVAREZ, B. & HOOPER, J.N.A., 2002. Family Axinellidae Carter, 1875. In: J.N.A.
HOOPER & R.W.M. VAN SOEST, eds, Systema Porifera: a guide to the
classification of sponges. First edn. New York, Boston, Dordrecht, London, Moscow:
Kluwer Academic/Plenum Publishers, pp. 724-747.
BAER, L., 1906. Silicispongien von Sansibar, Kapstadt und Papeete. Archiv für
Naturgeschichte, 72(1), pp. 1-32.
BARNES, D.K. & BELL, J.J., 2002. Coastal sponge communities of the West Indian Ocean:
taxonomic affinities, richness and diversity. African Journal of Ecology, 40(4), pp.
337-349.
BATSON, P., 2003. Deep New Zealand: blue water, black abyss. Christchurch, New
Zealand: Canterbury University.
BELL, J.J. & CARBALLO, J.L., 2008. Patterns of sponge biodiversity and abundance across
different biogeographic regions. Marine Biology, 155(6), pp. 563-570.
BERGQUIST, P.R., 1978. Sponges. London: Hutchinson & Co.
BERGQUIST, P.R., 1980. A revision of the supraspecific classification of the orders
Dictyoceratida, Dendroceratida and Verongida (class Demospongiae). New Zealand
Journal of Zoology, 7(4), pp. 443-503.
BERTOLINO, M., CERRANO, C., BAVESTRELLO, G., CARELLA, M., PANSINI, M. &
CALCINAI, B., 2013. Diversity of Porifera in the Mediterranean coralligenous
accretions, with description of a new species. ZooKeys, 336, pp. 1-37.
BIDDER, G.P., 1898. The skeleton and classification of calcareous sponges. Proceedings
of the Royal Society, 64, pp. 61-76.
132
BLAUSTEIN, R.J., 2010. High-seas biodiversity and genetic resources: science and policy
questions. Bioscience, 60(6), pp. 408-413.
BO, M., BERTOLINO, M., BORGHINI, M., CASTELLANO, M., HARRIAGUE, A.C., DI
CAMILLO, C.G., GASPARINI, G., MISIC, C., POVERO, P., PUSCEDDU, A.,
SCHROEDER, K. & BAVESTRELLO, G., 2011. Characteristics of the mesophotic
megabenthic assemblages of the Vercelli Seamount (North Tyrrhenian Sea). PLoS
ONE, 6(2), pp. e16357.
BOCAGE, J.V. BARBOZA, DU., 1869. Eponges siliceuses nouvelles du Portugal et de l’île
Saint-Iago (Archipel de Cap-Vert). Jornal de Sciencias Mathematicas, Physicas E
Naturaes, 2, pp. 159-162.
BOURY-ESNAULT, N. & LOPES, M.T., 1985. Les Démosponges littorales de l’Archipel
des Açores. Annales de l’Institut Océanographique, 61(2), pp. 149-225.
BOURY-ESNAULT, N. & RUTZLER, K., 1997. Thesaurus of sponge morphology.
Smithsonian Contributions to Zoology, 596, pp. 1-55.
BOWERBANK, J.S., 1862. On the anatomy and physiology of the Spongiadae. Part II.
Philosophical Transactions of the Royal Society of London, 152(2), pp. 747-836.
BOWERBANK, J.S., 1866. A monograph of the British Spongiadae. Volume 2. London:
Ray Society.
BOWERBANK, J.S., 1873. Contributions to a general history of the Spongiadae. Part IV.
Proceedings of the Zoological Society of London, pp. 3-25.
BRANCH, G.M. & BRANCH, M.L., 1981. The living shores of southern Africa. First edn.
Cape Town: Struik Publishers.
BREWIN, P.E., STOCKS, K.I. & MENEZES, G., 2007. A history of seamount research. In:
T.J. PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N. HAGGAN & R.S.
133
SANTOS, eds, Seamounts: ecology, fisheries & conservation. First edn. Oxford,
United Kingdom: Blackwell Publishing, pp. 41-61.
BRIGGS, J.C., 1974. Marine zoogeography. New York: McGraw-Hill.
BURTON, M., 1931. On a collection of marine sponges mostly from the Natal coast. Annals
of the Natal Museum, 6(3), pp. 337-358.
BURTON, M., 1933. Four new marine sponges from Natal. Annals of the Natal Museum,
7(2), pp. 249-254.
BURTON, M., 1959. Sponges. Scientific Reports. John Murray Expedition 1933-34. London:
British Museum (Natural History), pp. 151-281.
CAPON, R.J., MACLEOD, J.K. & WILLIS, A.C., 1987. Trunculins A and B,
norsesterterpene cyclic peroxides from a marine sponge, Latrunculia brevis. The
Journal of Organic Chemistry, 52(3), pp. 339-342.
CÁRDENAS, P., PÉREZ, T. & BOURY-ESNAULT, N., 2012. Sponge systematics facing
new challenges. Advances in Marine Biology, 61, pp. 79-209.
CÁRDENAS, P., XAVIER, J.R., REVEILLAUD, J., SCHANDER, C. & RAPP, H.T., 2011.
Molecular phylogeny of the Astrophorida (Porifera, Demospongiae) reveals an
unexpected high level of spicule homoplasy. PLoS ONE, 6(4), pp. e18318.
CARTER, H.J., 1875. Notes introductory to the study and classification of the Spongida.
Part II. Proposed classification of the Spongida. Annals and Magazine of Natural
History, 16(92), pp. 126-145.
CARTER, H.J., 1879. Contributions to our knowledge of the Spongida. Annals and
Magazine of Natural History, 3, pp. 284-304
CARTER, H.J., 1882. Some sponges from the West Indies and Acapulco in the Liverpool
Free Museum described, with general and classificatory remarks. Annals and
Magazine of Natural History, 9(52), pp. 266-301.
134
CARTER, H.J., 1883. Contributions to our knowledge of the Spongida. Annals and
Magazine of Natural History, 12(71), pp. 308-329.
CARTER, H.J., 1886. Descriptions of sponges from the neighbourhood of Port Phillip Heads,
South Australia, continued. Annals and Magazine of Natural History, 5(17), pp. 4053.
CHOMBARD, C. & BOURY-ESNAULT, N., 1999. Good congruence between morphology
and molecular phylogeny of Hadromerida, or how to bother sponge taxonomists.
Memoirs of the Queensland Museum, 44, pp. 100-100.
CLARK, A.M., 1972. Some crinoids from the Indian Ocean. Bulletin of the British Museum
(Natural History), 24(2), pp. 73-156.
CLARK, M.R., 2001. Are deepwater fisheries sustainable? - The example of orange roughy
(Hoplostethus atlanticus) in New Zealand. Fisheries Research, 51(2), pp. 123-135.
CLARK, M.R., ALTHAUS, F., SCHLACHER, T.A., WILLIAMS, A., BOWDEN, D.A. &
ROWDEN, A.A., 2015. The impacts of deep-sea fisheries on benthic communities: a
review. ICES Journal of Marine Science, fsv123, pp. 1-19.
CLARK, M.R. & KOSLOW, J.A., 2007. Impacts of fisheries on seamounts. In: T.J.
PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N. HAGGAN & R.S.
SANTOS, eds, Seamounts: ecology, fisheries & conservation. First edn. Oxford,
United Kingdom: Blackwell Publishing, pp. 413-441.
CLARK, M.R., ROWDEN, A.A., SCHLACHER, T., WILLIAMS, A., CONSALVEY, M.,
STOCKS, K.I., ROGERS, A.D., O'HARA, T.D., WHITE, M., SHANK, T.M. &
HALL-SPENCER, J.M., 2010. The ecology of seamounts: structure, function, and
human impacts. Annual Review of Marine Science, 2(1), pp. 253-278.
CLARK, M.R., ROWDEN, A.A. & STOCKS, K.I., 2004. CenSeam: a global census of
marine life on seamounts. A proposal for a new CoML field project.
135
CLARK, M.R., VINNICHENKO, V.I., GORDON, J.D.M., BECK-BULAT, G.Z.,
KUKHAREV, N.N. & KAKORA, A.F., 2007. Large-scale distant-water trawl
fisheries on seamounts. In: T.J. PITCHER, T. MORATO, P.J.B. HART, M.R.
CLARK, N. HAGGAN & R.S. SANTOS, eds, Seamounts: ecology, fisheries &
conservation. First edn. Oxford, United Kingdom: Blackwell Publishing, pp. 361-399.
CLARK, M.R., WATLING, L., ROWDEN, A.A., GUINOTTE, J.M. & SMITH, C.R., 2011.
A global seamount classification to aid the scientific design of marine protected area
networks. Ocean & Coastal Management, 54(1), pp. 19-36.
CLARKE, K.R. & GORLEY, R.N., 2006. PRIMER v6: user manual/tutorial. Plymouth:
PRIMER-E.
COLLETTE, B.B. & PARIN, N.V., 1991. Shallow-water fishes of Walters Shoals,
Madagascar Ridge. Bulletin of Marine Science, 48(1), pp. 1-22.
COLLETTE, B.B., SMITH, D.G. & BÖHLKE, E.B., 1991. Gymnothorax parini, a new
species of moray eel (Teleostei: Muraenidae) from Walters Shoals, Madagascar
Ridge. Proceedings of the Biological Society of Washington, 104(2), pp. 344-350.
CONSALVEY, M., CLARK, M.R., ROWDEN, A.A. & STOCKS, K.I., 2010. Life on
seamounts. In: A.D. MCINTYRE, ed, Life in the world’s oceans: diversity,
distribution, and abundance. First edn. Oxford: Wiley-Blackwell, pp. 123-138.
COSTELLO, M.J., COLL, M., DANOVARO, R., HALPIN, P., OJAVEER, H. &
MILOSLAVICH, P., 2010. A census of marine biodiversity knowledge, resources,
and future challenges. PloS ONE, 5(8), pp. e12110.
CRISTOBO, J., RIOS, P., POMPONI, S.A. & XAVIER, J.R., 2015. A new carnivorous
sponge, Chondrocladia robertballardi sp. nov. (Porifera: Cladorhizidae) from two
North-East Atlantic seamounts. Journal of the Marine Biological Association of the
United Kingdom, 95(7), pp. 1345-1352.
136
DAVIES, A.J., ROBERTS, J.M. & HALL-SPENCER, J., 2007. Preserving deep-sea natural
heritage: emerging issues in offshore conservation and management. Biological
Conservation, 138(3), pp. 299-312.
DE FORGES, B.R., KOSLOW, J.A. & POORE, G.C.B., 2000. Diversity and endemism of
the benthic seamount fauna in the Southwest Pacific. Nature, 405(6789), pp. 944-947.
DE LAUBENFELS, M.W.,1936. A discussion of the sponge fauna of the Dry Tortugas in
particular, and the West Indies in general, with material for a revision of the families
and orders of the Porifera. Carnegie Institute of Washington (Tortugas Laboratory
Paper No. 467), 30, pp. 1-225.
DE RUIJTER, W.P.M., RIDDERINKHOF, H., LUTJEHARMS, J.R.E., SCHOUTEN, M.W.
& VETH, C., 2002. Observations of the flow in the Mozambique Channel.
Geophysical Research Letters, 29(10), pp. 140-142.
DENDY, A., 1905. Report on the sponges collected by Professor Herdman at Ceylon in
1902. In: W. HERDMAN, ed, Report to the Government of Ceylon on the pearl
oyster fisheries of the Gulf of Manaar, Volume 3. London: Royal Society, pp. 57246.
DENDY, A., 1916. Report on the Homosclerophora and Astrotetraxonida collected by
H.M.S. ‘Sealark’ in the Indian Ocean. Reports of the Percy Sladen Trust Expedition to
the Indian Ocean in 1905, Volume 6. London: Transactions of the Linnean Society of
London, pp. 225-271.
DENDY, A., 1922. Report on the Sigmatotetraxonida collected by H.M.S. ‘Sealark’ in the
Indian Ocean. Reports of the Percy Sladen Trust Expedition to the Indian Ocean in
1905, Volume 7. London: Transactions of the Linnean Society of London, pp. 1-164.
DESQUEYROUX-FAÚNDEZ, R. & VALENTINE, C., 2002. Family Callyspongiidae de
Laubenfels, 1936. In: J.N.A. HOOPER & R.W.M. VAN SOEST, eds, Systema
137
Porifera: a guide to the classification of sponges. First edn. New York, Boston,
Dordrecht, London, Moscow: Kluwer Academic/Plenum Publishers, pp. 835-851.
DETINOVA, N.N. & SAGAIDACHNY, A.Y., 1994. Vertical distribution of bottom fauna
on the slope of the Walters Shoal (Madagascar Ridge). Transactions of the P.P.
Shirshov Institute of Oceanology [Trudy Instituta Okeanologii], 129, pp. 17-30.
DIAZ, M.C. & RÜTZLER, K., 2009. Biodiversity and abundance of sponges in Caribbean
mangrove: indicators of environmental quality. Smithsonian Contributions to the
Marine Sciences, 38, pp. 151-172.
DUCHASSAING DE FONBRESSIN, P. & MICHELOTTI, G., 1864. Spongiaires de la mer
Caraibe. Natuurkundige verhandelingen van de Hollandsche maatschappij der
wetenschappen te Haarlem, 21(2), pp. 1-124.
DUCKWORTH, A.R. & BATTERSHILL, C.N., 2001. Population dynamics and chemical
ecology of New Zealand Demospongiae Latrunculia sp. nov. and Polymastia croceus
(Poecilosclerida: Latrunculiidae: Polymastiidae). New Zealand Journal of Marine and
Freshwater Research, 35(5), pp. 935-949.
ERPENBECK, D., SUTCLIFFE, P., COOK, S.D.C., DIETZEL, A., MALDONADO, M.,
VAN
SOEST,
R.W.M.,
HOOPER,
J.N.A.
&
WÖRHEIDE,
G.,
2012.
Horny sponges and their affairs: on the phylogenetic relationships of keratose
sponges. Molecular Phylogenetics and Evolution, 63(3), pp. 809-816.
ERPENBECK, D. & VAN SOEST, R.W.M., 2002. Family Halichondriidae Gray, 1867. In:
J.N.A. HOOPER & R.W.M. VAN SOEST, eds, Systema Porifera: a guide to the
classification of sponges. First edn. New York, Boston, Dordrecht, London,
Moscow: Kluwer Academic/Plenum Publishers, pp. 787-815.
ESPER, E.J.C., 1794. Die Pflanzenthiere in Abbildungen nach der Natur mit Farben
erleuchtet, nebst Beschreibungen. Zweyter Theil. Nürnberg: Raspe.
138
ESPER, E.J.C., 1797. Fortsetzungen der Pflanzenthiere in Abbildungennach der Natur mit
Farben erleuchtet nebst Beschreibungen. Nürnberg: Erster Theil.
FLEMING, J., 1828. A history of British animals, exhibiting the descriptive characters and
systematical arrangement of the genera and species of quadrupeds, birds, reptiles,
fishes, mollusca, and radiata of the United Kingdom; including the indigenous,
extirpated, and extinct kinds, together with periodical and occasional visitants.
Edinburgh: Bell and Bradfute.
FULTON, B., MORATO, T. & PITCHER, T.J., 2007. Modelling seamount ecosystems and
their fisheries. In: T.J. PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N.
HAGGAN & R.S. SANTOS, eds, Seamounts: ecology, fisheries & conservation.
First edn. Oxford, United Kingdom: Blackwell Publishing, pp. 296-332.
GAGE, J.D. & TYLER, P.A., 1991. Deep-sea biology: a natural history of organisms at the
deep-sea floor. First edn. Cambridge: Cambridge University Press.
GALTSOFF, P.S., 1960. Sponges. Fishery Leaflet 490. Washington, D.C.: United States
Fish and Wildlife Service.
GENIN, A., 2004. Bio-physical coupling in the formation of zooplankton and fish
aggregations over abrupt topographies. Journal of Marine Systems, 50(1), pp. 3-20.
GENIN, A. & DOWER, J.F., 2007. Seamount plankton dynamics. In: T.J. PITCHER, T.
MORATO, P.J.B. HART, M.R. CLARK, N. HAGGAN & R.S. SANTOS, eds,
Seamounts: ecology, fisheries & conservation. First edn. Oxford, United Kingdom:
Blackwell Publishing, pp. 85-100.
GLOVER, A.G. & SMITH, C.R., 2003. The deep-sea floor ecosystem: current status and
prospects of anthropogenic change by the year 2025. Environmental Conservation,
30(3), pp. 219-241.
139
GOPAL, K., 2007. Genetic population structure of spiny lobster Palinurus delagoae in the
South-Western Indian Ocean, and the evolutionary history of Palinurus. MSc thesis.
Stellenbosch: University of Stellenbosch.
GRANT, R.E., 1836. Animal kingdom. In: R.B. TODD, ed, The cyclopaedia of anatomy and
physiology. Volume 1. London: Sherwood, Gilbert and Piper, pp. 107-118.
GRANT, R.E., 1861. Tabular view of the primary divisions of the animal kingdom,
intended to serve as an outline of an elementary course of recent zoology. London:
Walton & Maberly.
GRAY, J.E., 1848. List of the specimens of British sponges in the collection of the British
Museum (London). British Museum Publication, 8, pp. 1-24.
GRAY, J.E., 1867. Notes on the arrangement of sponges, with the descriptions of some new
genera. Proceedings of the Zoological Society of London, 2, pp. 492-558.
GROENEVELD, J.C., GRIFFITHS, C.L. & VAN DALSEN, A.P., 2006. A new species of
spiny lobster, Palinurus barbarae (Decapoda, Palinuridae) from Walters Shoals on
the Madagascar Ridge. Crustaceana, 79(7), pp. 821-833.
GUINOTTE, J.M., ORR, J., CAIRNS, S., FREIWALD, A., MORGAN, L. & GEORGE, R.,
2006. Will human-induced changes in seawater chemistry alter the distribution of
deep-sea scleractinian corals? Frontiers in Ecology and the Environment, 4(3), pp.
141-146.
HARTMAN, W.D., 1958. Re-examination of Bidder's classification of the Calcarea.
Systematic Zoology, 7(3), pp. 55-110.
HARTMAN, W.D., 1980. Systematics of the Porifera. In: W.D. HARTMAN, J.W. WENDT
& F. WIEDENMAYER, eds, Living and fossil sponges, notes for a short course.
Sedimenta 8. Miami: Rosenstiel School of Marine and Atmospheric Science, pp. 2451.
140
HENRICH, R., HARTMANN, M., REITNER, J., SCHÄFER, P., FREIWALD, A.,
STEINMETZ, S., DIETRICH, P. & THIEDE, J., 1992. Facies belts and communities
of the Arctic Vesterisbanken Seamount (Central Greenland Sea). Facies, 27(1), pp.
71-104.
HENTSCHEL, E., 1923. Erste Unterabteilung der Metazoa: Parazoa, Porifera-Schwämme.
In: W. KÜKENTHAL & T. KRUMBACH, eds, Handbuch der Zoologie. Eine
Naturgeschichteder Stämme des Tierreiches. Vol. 1, Protozoa, Porifera, Coelenterata,
Mesozoa. Berlin, Leipzi: Walter de Gruyter und Co, pp. 307-418.
HILLIER, J.K. & WATTS, A.B., 2007. Global distribution of seamounts from ship‐track
bathymetry data. Geophysical Research Letters, 34(13), pp. 1-5.
HOFMAN, C.C. & VAN SOEST, R.W.M., 1995. Lissodendoryx species of the IndoMalayan Archipelago (Demospongiae: Poecilosclerida). Beaufortia, 45(6), pp. 77103.
HOLLAND, K.N. & GRUBBS, R.D., 2007. Fish visitors to seamounts: tunas and billfish at
seamounts. In: T.J. PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N.
HAGGAN & R.S. SANTOS, eds, Seamounts: ecology, fisheries & conservation.
First edn. Oxford, United Kingdom: Blackwell Publishing, pp. 189-201.
HOLMES, K.E., 2000. Effects of eutrophication on bioeroding sponge communities with the
description of new West Indian sponges, Cliona spp. (Porifera: Hadromerida:
Clionidae). Invertebrate Biology, 119(2), pp. 125-138.
HOOPER, J.N.A., 1996. Guide to sponge collection and identification. Australia: Queensland
Museum.
HOOPER, J.N.A., 2002. Family Microcionidae Carter, 1875. In: J.N.A. HOOPER & R.W.M.
VAN SOEST, eds, Systema Porifera: a guide to the classification of sponges. First
141
edn. New York, Boston, Dordrecht, London, Moscow: Kluwer Academic/Plenum
Publishers, pp. 432-468.
HOOPER, J.N.A., 2003. 'Sponguide'. Guide to sponge collection and identification.
Australia: Queensland Museum.
HOOPER, J.N.A. & VAN SOEST, R.W.M., 2002. Systema Porifera: a guide to the
classification of sponges. First edn. New York, Boston, Dordrecht, London, Moscow:
Kluwer Academic/ Plenum Publishers.
HUBBS, C.L., 1959. Initial discoveries of fish faunas on seamounts and offshore banks in the
eastern Pacific. Pacific Science, XIII, pp. 311-316.
INGOLE, B. & KOSLOW, J.A., 2005. Deep-sea ecosystems of the Indian Ocean. Indian
Journal of Marine Sciences, 34(1), pp. 27-34.
IWAMOTO, T., SHCHERBACHEV, Y.N. & MARQUARDT, B., 2004. Grenadiers
(Gadiformes, Teleostei) of Walters Shoals, Southwestern Indian Ocean, with
description of a new "West-Wind Drift" species. Proceedings of the California
Academy of Sciences, 55(10), pp. 190-207.
JONES, D.O.B. & GATES, A.R., 2010. Deep-sea life of Scotland and Norway. United
Kingdom: Ophiura Publishing.
KADMON, R. & ALLOUCHE, O., 2007. Integrating the effects of area, isolation, and
habitat heterogeneity on species diversity: a unification of island biogeography and
niche theory. The American Naturalist, 170(3), pp. 443-454.
KASCHNER, K., 2007. Air-breathing visitors to seamounts: marine mammals. In: T.J.
PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N. HAGGAN & R.S.
SANTOS, eds, Seamounts: ecology, fisheries & conservation. First edn. Oxford,
United Kingdom: Blackwell Publishing, pp. 230-238.
142
KELLY, M., ERPENBECK, D., MORROW, C. & SOEST, R.V., 2015. First record of a
living species of the genus Janulum (Class Demospongiae) in the Southern
Hemisphere. Zootaxa, 3980(2), pp. 255-266.
KELLY-BORGES, M., 1997. Porifera - sponges. In: M.D. RICHMOND, ed, A guide to the
seashores of eastern Africa and the Western Indian Ocean islands. Stockholm,
Sweden: Swedish International Development Cooperation Agency (SIDA), SAREC.,
pp. 106-117.
KENSLEY, B.F., 1969. Decapod Crustacea from the South-West Indian Ocean. Annals of the
South African Museum, 52(7), pp. 149-181.
KENSLEY, B.F., 1975. Five species of Jaeropsis from the southern Indian Ocean
[Crustacea, Isopoda, Asellota]. Annals of the South African Museum, 67(10), pp. 367380.
KENSLEY, B.F., 1981. On the zoogeography of southern African decapod Crustacea, with a
distributional checklist of the species. Smithsonian Contributions to Zoology, 338, pp.
1-64.
KIRKPATRICK, R., 1903. Descriptions of South African sponges. Part III. Marine
Investigations in South Africa, 2(14), pp. 233-264.
KITCHINGMAN, A. & LAI, S., 2004. Inferences on potential seamount locations from midresolution bathymetric data. In: T. MORATO & D. PAULY, eds, Seamounts:
biodiversity and fisheries. Canada: Fisheries Centre, University of British Columbia,
pp. 7-12.
KVILE, K.Ø., TARANTO, G.H., PITCHER, T.J. & MORATO, T., 2014. A global
assessment of seamount ecosystems knowledge using an ecosystem evaluation
framework. Biological Conservation, 173, pp. 108-120.
143
LAMARCK J.B.P.A. & DE MONET, C.D., 1813 -1814. Sur les polypiers empâtés. Suite du
mémoire intitulé: Sur les polypiers empâtés. Suite des éponges. Annales du Muséum
National d'Histoire Naturelle, Paris, 20(6), pp. 294-458.
LAMARCK, J.B.P.A. & DE MONET, C.D., 1815. Suite des polypiers empâtés. Mémoires du
Muséum d’Histoire Naturelle, Paris, 1, pp. 69-340.
LANDEIRA, J.M., LOZANO-SOLDEVILLA, F., HERNÁNDEZ-LEÓN, S. & BARTON,
E.D., 2010. Spatial variability of planktonic invertebrate larvae in the Canary Islands
area. Journal of the Marine Biological Association of the United Kingdom, 90(6), pp.
1217-1225.
LAPTIKHOVSKY, V., BOERSCH-SUPAN, P., BOLSTAD, K., KEMP, K., LETESSIER, T.
& ROGERS, A.D., 2015. Cephalopods of the Southwest Indian Ocean Ridge: a
hotspot of biological diversity and absence of endemism. Deep Sea Research Part II:
Topical Studies in Oceanography, Advance online publication. Available from:
http://www.sciencedirect.com/science/article/pii/S0967064515002283.
LE CORRE, M., JAEGER, A., PINET, P., KAPPES, M.A., WEIMERSKIRCH, H., CATRY,
T., RAMOS, J.A., RUSSELL, J.C., SHAH, N. & JAQUEMET, S., 2012. Tracking
seabirds to identify potential marine protected areas in the tropical Western Indian
Ocean. Biological Conservation, 156, pp. 83-93.
LENDENFELD, R.V., 1897. Spongien von Sansibar. Abhandlungen herausgegeben von der
Senckenbergischen Naturforschenden Gesellschaft, 21, pp. 93-133.
LENDENFELD, R.V., 1903. Porifera. Tetraxonia. In: F.E. SCHULZE, ed, Das Tierreich.
Berlin: Friedländer, pp. 1-168.
LENDENFELD, R.V., 1907. Die Tetraxonia. Wissenschaftliche Ergebnisse der Deutschen
Tiefsee-Expedition auf der Dampfer Valdivia 1898-1899, 11(1-2), pp. 59-374.
144
LESSER, M.P., SLATTERY, M. & LEICHTER, J.J., 2009. Ecology of mesophotic coral
reefs. Journal of Experimental Marine Biology and Ecology, 375(1), pp. 1-8.
LETESSIER, T.B., DE GRAVE, S., BOERSCH-SUPAN, P.H., KEMP, K.M., BRIERLEY,
A.S. & ROGERS, A.D., 2015. Seamount influences on mid-water shrimps
(Decapoda) and gnathophausiids (Lophogastridea) of the South-West Indian Ridge.
Deep Sea Research Part II: Topical Studies in Oceanography, Advance online
publication. Available from:
http://www.sciencedirect.com/science/article/pii/S0967064515001757.
LÉVI, C., 1953. Sur une nouvelle classification des Démosponges. Comptes Rendus de
l'Académie des Sciences, 236(8), pp. 853-855
LÉVI, C., 1958. Résultats scientifiques des Campagnes de la ‘Calypso’. Campagne 19511952 en Mer Rouge (suite). 11. Spongiaires de Mer Rouge recueillis par la ‘Calypso’
(1951-1952). Annales de l’Institut Océanographique, 34(3), pp. 3-46.
LÉVI, C., 1961. Résultats scientifiques des Campagnes de la ‘Calypso’. Campagne 1954
dans l’Océan Indien (suite). 2. Les spongiaires de l’Ile Aldabra. Annales de l’Institut
Océanographique, 39(1), pp. 3-32.
LÉVI, C., 1963. Spongiaires d’Afrique du Sud. (1) Poecilosclérides. Transactions of the
Royal Society of South Africa, 37(1), pp. 1-72.
LÉVI, C., 1964. Spongiaires du Canal de Mozambique. Bulletin du Muséum National
d'Histoire Naturelle, 36(3), pp. 384-395
LÉVI, C., 1967. Spongiaires d'Afrique du Sud. (3) Tétractinellides. Transactions of the Royal
Society of South Africa, 37, pp. 227-256.
LÉVI, C., 1969. Spongiaires du Vema Seamount (Atlantique Sud). Bulletin du Muséum
National d'Histoire Naturelle, 41(4), pp. 952-973.
145
LÉVI, C. & LÉVI, P., 1983. Eponges Tétractinellides et Lithistides bathyales de NouvelleCalédonie. Bulletin du Muséum National d'Histoire Naturelle, 5(1), pp. 101-168.
LITVINOV, F., 2007. Fish visitors to seamounts: aggregations of large pelagic sharks above
seamounts. In: T.J. PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N.
HAGGAN & R.S. SANTOS, eds, Seamounts: ecology, fisheries & conservation.
First edn. Oxford, United Kingdom: Blackwell Publishing, pp. 202-206.
MALDONADO, M., 2006. The ecology of the sponge larva. Canadian Journal of Zoology,
84(2), pp. 175-194.
MARIANI, S., URIZ, M., TURON, X. & ALCOVERRO, T., 2006. Dispersal strategies in
sponge larvae: integrating the life history of larvae and the hydrologic component.
Oecologia, 149(1), pp. 174-184.
MARSHALL, N.B., 1979. Developments in deep-sea biology. Dorset: Blandford Press.
MARSHALL, W., 1876. Ideen über die Verwandtschaftsverhältnisse der Hexactinelliden.
Zeitschrift für wissenschaftliche Zoologie, 27(1), pp. 113-136.
MCCLAIN, C.R., 2007. Seamounts: identity crisis or split personality? Journal of
Biogeography, 34(12), pp. 2001-2008.
MCCLAIN, C.R. & LUNDSTEN, L., 2015. Assemblage structure is related to slope and
depth on a deep offshore Pacific seamount chain. Marine Ecology, 36(2), pp. 210220.
MENARD, H.W., 1964. Marine geology of the Pacific. First edn. New York: McGraw-Hill.
MINCHIN, E.A., 1900. Chapter III. Sponges. In: E.R. LANKESTER, ed, A treatise on
zoology. Part II. The Porifera and Coelenterata. London: Adam & Charles Black, pp.
1-178.
MLADENOV, P.V., 2013. Marine biology - a very short introduction. United Kingdom:
Oxford University Press.
146
MORATO, T., CHEUNG, W.W.L. & PITCHER, T.J., 2004. Vulnerability of seamount fish
to fishing: fuzzy analysis of life-history attributes. In: T. MORATO & D. PAULY,
eds, Seamounts: biodiversity and fisheries. Canada: Fisheries Centre, University of
British Columbia, pp. 51-60.
MORATO, T. & CLARK, M.R., 2007. Seamount fishes: ecology and life histories. In: T.J.
PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N. HAGGAN & R.S.
SANTOS, eds, Seamounts: ecology, fisheries & conservation. First edn. Oxford,
United Kingdom: Blackwell Publishing, pp. 170-188.
MORATO, T., VARKEY, D.A., DAMASO, C., MACHETE, M., SANTOS, M., PRIETO,
R., SANTOS, R.S. & PITCHER, T.J., 2008. Evidence of a seamount effect on
aggregating visitors. Marine Ecology Progress Series, 357, pp. 23-32.
MORROW, C.C. & CÁRDENAS, P., 2015. Proposal for a revised classification of the
Demospongiae (Porifera). Frontiers in Zoology, 12(7), pp. 1-27.
MORROW, C.C., PICTON, B.E., ERPENBECK, D., BOURY-ESNAULT, N., MAGGS,
C.A. & ALLCOCK, A.L., 2012. Congruence between nuclear and mitochondrial
genes in Demospongiae: a new hypothesis for relationships within the G4 clade
(Porifera: Demospongiae). Molecular Phylogenetics and Evolution, 62(1), pp. 174190.
MORROW, C.C., REDMOND, N.E., PICTON, B.E., THACKER, R.W., COLLINS, A.G.,
MAGGS, C.A., SIGWART, J.D. & ALLCOCK, A.L., 2013. Molecular phylogenies
support homoplasy of multiple morphological characters used in the taxonomy of
Heteroscleromorpha (Porifera: Demospongiae). Integrative and Comparative Biology,
53(3), pp. 428-446.
147
NARDO, G.D., 1833. Auszug aus einem neuen System der Spongiarien, wonach bereits die
Aufstellung in der Universitäts-Sammlung zu Padua gemacht ist. Isis, oder
Encyclopädische Zeitung Coll. Jena: Oken, pp. 519-523.
NESIS, K.N., 1994. Teuthofauna of Walters Shoals, a seamount in the Southwestern Indian
Ocean. Ruthenica, 4(1), pp. 67-77.
NESIS, K.N., 2003. Distribution of recent Cephalopoda and implications for Plio-Pleistocene
events. Berliner Paläobiologische Abhandlungen, 3, pp. 199-224.
O’HARA, T.D., 2007. Seamounts: centres of endemism or species richness for ophiuroids?
Global Ecology and Biogeography, 16(6), pp. 720-732.
PARIN, N.V., NESIS, K.N., SAGAIDACHNY, A.Y. & SHCHERBACHEV, Y.N., 1993.
Fauna of Walters Shoals, a seamount in the Southwestern Indian Ocean. Transactions
of the P.P. Shirshov Institute of Oceanology [Trudy Instituta Okeanologii], 128, pp.
199-216.
PECHENIK, J.A., 2009. Biology of the invertebrates. Sixth edn. New York: McGraw-Hill
Higher Education.
PEREIRA, R., GOMES PEREIRA, J.N., TEMPERA, F., PORTEIRO, F. & XAVIER, J.R.,
2015. Sponge assemblages of the Condor Seamount (Azores) characterized from
underwater imagery, 14th Deep-Sea Biology Symposium: abstract book, 31 August - 4
September 2015, UA Editora, pp. 258.
PITCHER, T.J., MORATO, T., HART, P.J.B., CLARK, M.R., HAGGAN, N. & SANTOS,
R.S., 2007. Preface. In: T.J. PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK,
N. HAGGAN & R.S. SANTOS, eds, Seamounts: ecology, fisheries & conservation.
First edn. Oxford, United Kingdom: Blackwell Publishing, pp. xvii-xxi.
POLLARD, R. & READ, J., 2015. Circulation, stratification and seamounts in the Southwest
Indian Ocean. Deep Sea Research Part II: Topical Studies in Oceanography,
148
Advance online publication. Available from:
http://www.sciencedirect.com/science/article/pii/S0967064515000478.
POSS, S.G. & COLLETTE, B.B., 1990. Scorpaenodes immaculatus, a new species of
scorpionfish (Osteichthyes: Scorpaenida) from Walters Shoals, Madagascar Ridge.
Proceedings of the Biological Society of Washington, 103(3), pp. 543-549.
PULITZER-FINALI, G., 1993. A collection of marine sponges from East Africa. Annales
Museo Civico Storia Naturale "Giacomo Doria", 89, pp. 247-350.
RAMIREZ-LLODRA, E., BRANDT, A., DANOVARO, R., DE MOL, B., ESCOBAR, E.,
GERMAN, C.R., LEVIN, L.A., ARBIZU, P.M., MENOT, L., BUHL-MORTENSEN,
P., NARAYANASWAMY, B.E., SMITH, C.R., TITTENSOR, D.P., TYLER, P.A.,
VANREUSEL, A. & VECCHIONE, M., 2010. Deep, diverse and definitely different:
unique attributes of the world's largest ecosystem. Biogeosciences, 7, pp. 2851-2899.
RAMIREZ-LLODRA, E., TYLER, P.A., BAKER, M.C., BERGSTAD, O.A., CLARK,
M.R., ESCOBAR, E., LEVIN, L.A., MENOT, L., ROWDEN, A.A., SMITH, C.R. &
VAN DOVER, C.L., 2011. Man and the last great wilderness: human impact on the
deep sea. PLoS ONE, 6(8), pp. e22588.
READ, J. & POLLARD, R., 2015. An introduction to the physical oceanography of six
seamounts in the Southwest Indian Ocean. Deep Sea Research Part II: Topical
Studies
in
Oceanography,
Advance
online
publication.
Available
from:
http://www.sciencedirect.com/science/article/pii/S0967064515002246.
REX, M.A., 1981. Community structure in the deep-sea benthos. Annual Review of Ecology
and Systematics, 12, pp. 331-353.
REX, M.A. & ETTER, R.J., 2010. Deep-sea biodiversity: pattern and scale. United States:
Harvard University Press.
149
RICHMOND, M.D., 2001. The marine biodiversity of the Western Indian Ocean and its
biogeography: How much do we know? In: M.D. RICHMOND & J. FRANCIS, eds,
Marine science development in Tanzania and eastern Africa. Proceedings of the 20th
anniversary
conference
on
advances
in
marine
science
in
Tanzania.
(IMS/WIOMSA), pp. 241-261.
RIDDERINKHOF, H., LUTJEHARMS, J.R.E. & DE RUIJTER, W.P.M., 2001. A research
cruise to investigate the Mozambique Current. South African Journal of Science,
97(11/12), pp. 461-464.
RIDLEY, S.O., 1884. Spongiida. Report on the zoological collections made in the IndoPacific Ocean during the voyage of H.M.S. ‘Alert’, 1881-2. London: British Museum
(Natural History), pp. 366-630.
RIDLEY, S.O. & DENDY, A., 1886. Preliminary report on the Monaxonida collected by
H.M.S. ‘Challenger’. Annals and Magazine of Natural History, 5(18), pp. 325-493.
ROBERTS, C.M., MCCLEAN, C.J., VERON, J.E.N., HAWKINS, J.P., ALLEN, G.R.,
MCALLISTER, D.E., MITTERMEIER, C.G., SCHUELER, F.W., SPALDING, M.,
WELLS, F., VYNNE, C. & WERNER, T.B., 2002. Marine biodiversity hotspots and
conservation priorities for tropical reefs. Science, 295(5558), pp. 1280-1284.
ROGERS, A.D., 1994. The biology of seamounts. Advances in Marine Biology, 30, pp. 305351.
ROGERS, A.D., 2012. An ecosystem approach to management of seamounts in the southern
Indian Ocean. Volume 1 – Overview of seamount ecosystems and biodiversity. Gland,
Switzerland: International Union for the Conservation of Nature.
ROGERS, A.D., ALVHEIM, O., BEMANAJA, E., BENIVARY, D., BOERSCH-SUPAN,
P.H., BORNMAN, T., CEDRAS, R., DU PLESSIS, N., GOTHEIL, S., HOINES, A.,
KEMP, K., KRISTIANSEN, J., LETESSIER, T., MANGAR, V., MAZUNGULA, N.,
150
MØRK, T., PINET, P., READ, J. & SONNEKUS, T., 2009. Cruise report "Dr.
Fridtjof Nansen" southern Indian Ocean seamounts (IUCN/ UNDP/ ASCLME/ NERC
/EAF Nansen Project 2009 Cruise 410) 12th November – 19th December, 2009
Gland, Switzerland: International Union for the Conservation of Nature.
ROMANOV, E.V., 2003. Summary and review of Soviet and Ukrainian scientific and
commercial fishing operations on the deepwater ridges of the southern Indian Ocean.
FAO Fisheries Circular No. 991. Rome, Italy: FAO.
ROWDEN, A.A., DOWER, J.F., SCHLACHER, T.A., CONSALVEY, M. & CLARK, M.R.,
2010. Paradigms in seamount ecology: fact, fiction and future. Marine Ecology,
31(s1), pp. 226-241.
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, pp. 1-37.
SAMAAI, T. & GIBBONS, M.J., 2005. Demospongiae taxonomy and biodiversity of the
Benguela region on the west coast of South Africa. African Natural History, 1, pp. 196.
SAMAAI, T., GIBBONS, M.J. & KELLY, M., 2006. Revision of the genus Latrunculia du
Bocage, 1869 (Porifera: Demospongiae: Latrunculiidae) with descriptions of new
species from New Caledonia and the Northeastern Pacific. Zootaxa, 1127, pp. 1-71.
SAMAAI, T., GIBBONS, M.J., KELLY, M. & DAVIES-COLEMAN, M., 2003. South
African Latrunculiidae (Porifera: Demospongiae: Poecilosclerida): descriptions of
new species of Latrunculia du Bocage, Strongylodesma Levi, and Tsitsikamma
Samaai & Kelly. Zootaxa, 371, pp. 1-26.
151
SAMAAI, T., GIBBONS, M.J., KERWATH, S., YEMANE, D. & SINK, K., 2010. Sponge
richness along a bathymetric gradient within the iSimangaliso Wetland Park, South
Africa. Marine Biodiversity, 40(3), pp. 205-217.
SAMAAI, T., JANSON, L. & KELLY, M., 2012. New species of Latrunculia from the
Agulhas shelf, South Africa, with designation of a type species for subgenus
Biannulata (Demospongiae, Poecilosclerida, Latrunculiidae). Zootaxa, 3395, pp. 3345.
SAMAAI, T. & KELLY, M., 2002. Family Latrunculiidae Topsent, 1922. In: J.N.A.
HOOPER & R.W.M. VAN SOEST, eds, Systema Porifera: a guide to the
classification of sponges. New York: Kluwer Academic/ Plenum Publishers,
pp.708-719.
SAMADI, S., SCHLACHER, T. & DE FORGES, B.R., 2007. Seamount benthos. In: T.J.
PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N. HAGGAN & R.S.
SANTOS, eds, Seamounts: ecology, fisheries & conservation. First edn. Oxford,
United Kingdom: Blackwell Publishing, pp. 119-140.
SANTOS, M.A., BOLTEN, A.B., MARTINS, H.R., RIEWALD, B. & BJORNDAL, K.A.,
2007. Air-breathing visitors to seamounts: sea turtles. In: T.J. PITCHER, T.
MORATO, P.J.B. HART, M.R. CLARK, N. HAGGAN & R.S. SANTOS, eds,
Seamounts: ecology, fisheries & conservation. First edn. Oxford, United Kingdom:
Blackwell Publishing, pp. 239-244.
SARÀ, M., 2002. Family Tethyidae Gray, 1848. In: J.N.A. HOOPER & R.W.M. VAN
SOEST, eds, Systema Porifera: a guide to the classification of sponges. First edn.
New York, Boston, Dordrecht, London, Moscow: Kluwer Academic/Plenum
Publishers, pp. 245-265.
152
SAUTYA, S., INGOLE, B., RAY, D., STÖHR, S., SAMUDRALA, K., RAJU, K.A.K. &
MUDHOLKAR, A., 2011. Megafaunal community structure of Andaman seamounts
including the back-arc basin - a quantitative exploration from the Indian Ocean.
PloS ONE, 6(1), pp. e16162.
SCHLACHER-HOENLINGER, M.A., PISERA, A. & HOOPER, J.N.A., 2005. Deep-sea
"lithistid" assemblages from the Norfolk Ridge (New Caledonia), with description of
seven new species and a new genus (Porifera, Demospongiae). Zoosystema, 27(4), pp.
649-698.
SCHLICH, R., SIMPSON, E.S.W., GIESKES, J., GIRDLEY, W.A., LECLAIRE, L.,
MARSHALL, B.V., MOORE, C., MULLER, C., SIGAL, J., VALUER, T.L.,
WHITE, S.M. & ZOBEL, B., 1974. Sites 246 and 247. In: T.L. VALUER & S.M.
WHITE, eds, Initial reports of the Deep Sea Drilling Project New York: U.S.
Government Printing Office, pp. 237-257.
SCHMIDT, O., 1862. Die Spongien des Adriatischen Meeres. Leipzig: Wilhelm Engelmann.
SCHMIDT, O., 1864. Supplement der Spongien des Adriatischen Meeres. Enthaltend die
Histologie und systematische Ergänzungen. Leipzig: Wilhelm Engelmann.
SCHMIDT, O., 1870. Grundzüge einer Spongien-Fauna des atlantischen Gebietes. Leipzig:
Wilhelm Engelmann.
SCHMIDT, R. & SCHMINCKE, H., 2000. Seamounts and island building. In: H.
SIGURDSSON, B. HOUGHTON, S.R. MCNUTT, H. RYMER & J. STIX, eds,
Encyclopedia of volcanoes. San Diego, California: Academic Press, pp. 383-402.
SCHUCHERT, P. & REISWIG, H.M., 2006. Brinckmannia hexactinellidophila, n. gen., n.
sp.: a hydroid living in tissues of glass sponges of the reefs, fjords, and seamounts of
Pacific Canada and Alaska. Canadian Journal of Zoology, 84(4), pp. 564-572.
153
SCHULZE, F.E., 1877. Untersuchungen über den Bau und die Entwicklung der Spongien. II.
Die Gattung Halisarca. Zeitschrift für wissenschaftliche Zoologie, 28, pp. 1-48.
SHANK, T.M., 2010. Seamounts: deep-ocean laboratories of faunal connectivity, evolution,
and endemism. Oceanography, 23(1), pp. 108-122.
SHOTTON, R., 2006. Management of demersal fisheries resources of the southern Indian
Ocean. FAO Fisheries Circular No. 1020. Rome, Italy: FAO.
SMITH, C.R., LEVIN, L.A., KOSLOW, A., TYLER, P.A. & GLOVER, A.G., 2008. The
near future of the deep seafloor ecosystems. In: N.V.C. POLUNIN, ed, Aquatic
ecosystems: trends and global prospects. United Kingdom, New York: Cambridge
University Press, pp. 334-349.
SMITH, D.K. & CANN, J.R., 1992. The role of seamount volcanism in crustal construction
at the Mid‐Atlantic Ridge (24°–30° N). Journal of Geophysical Research: Solid Earth
(1978–2012), 97(B2), pp. 1645-1658.
SOLLAS, W.J., 1885. A classification of the sponges. Annals and Magazine of Natural
History, 16(95), pp. 395.
SOLLAS, W.J., 1887. Sponges. In: A. BLACK & C. BLACK, eds, Encyclopaedia
britannica. Ninth edn. Edinburgh: pp. 412-429.
SOLLAS, W.J., 1888. Report on the Tetractinellida collected by H.M.S. 'Challenger', during
the years 1873-1876. Report on the scientific results of the voyage of H.M.S.
'Challenger', 1873-1876. Zoology, 25(63), pp. 1-458.
SPALDING, M.D., FOX, H.E., ALLEN, G.R., DAVIDSON, N., FERDAÑA, Z.A.,
FINLAYSON, M., HALPERN, B.S., JORGE, M.A., LOMBANA, A., LOURIE, S.A.,
MARTIN, K.D., MCMANUS, E., MOLNAR, J., RECCHIA, C.A. & ROBERTSON,
J., 2007. Marine ecoregions of the world: a bioregionalization of coastal and shelf
areas. BioScience, 57(7), pp. 573-583.
154
STAUDIGEL, H., KOPPERS, A.A., LAVELLE, J.W., PITCHER, T.J. & SHANK, T.M.,
2010. Defining the word “seamount”. Oceanography, 23(1), pp. 20-21.
STOCKS, K.I., CLARK, M.R., ROWDEN, A.A., CONSALVEY, M. & SCHLACHER,
T.A., 2012. CenSeam, an international program on seamounts within the Census of
Marine Life: achievements and lessons learned. PloS ONE, 7(2), pp. e32031.
STOCKS, K.I. & HART, P.J.B., 2007. Biogeography and biodiversity of seamounts. In: T.J.
PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N. HAGGAN & R.S.
SANTOS, eds, Seamounts: ecology, fisheries & conservation. First edn. Oxford,
United Kingdom: Blackwell Publishing, pp. 255-281.
SUNDAR, V.C., YABLON, A.D., GRAZUL, J.L., ILAN, M. & AIZENBERG, J., 2003.
Fibre-optical features of a glass sponge. Nature, 424(6951), pp. 899-900.
THIELE, J., 1900. Kieselschwämme von Ternate. I. Abhandlungen herausgegeben von der
Senckenbergischen Naturforschenden Gesellschaft, 25, pp. 19-80.
THIELE, J., 1903. Kieselschwämme von Ternate. II. Abhandlungen herausgegeben von der
Senckenbergischen Naturforschenden Gesellschaft, 25, pp. 933-968.
THIELE, J., 1905. Die Kiesel- und Hornschwämme der Sammlung Plate. Zoologische
Jahrbücher Supplement, 6, pp. 407-496.
THOMAS, P.A., 1973. Marine Demospongiae of Mahe Island in the Seychelles Bank (Indian
Ocean). Annales du Musée Royal De L'Afrique Centrale, 203, pp. 1-96.
THOMAS, P.A., 1979. Studies on sponges of the Mozambique Channel. I. Sponges of the
Inhaca Island. II. Sponges of Mambone and Paradise Islands. Annales du Musée
Royal De L'Afrique Centrale, 227, pp. 1-73.
THOMAS, P.A., 1984. Sponges collected aboard R.V. 'Skipjack' from the southeast coast of
India. Journal of the Marine Biological Association of India, 26(1&2), pp. 95-102.
155
THOMPSON, D.R., 2007. Air-breathing visitors to seamounts: importance of seamounts to
seabirds. In: T.J. PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK, N.
HAGGAN & R.S. SANTOS, eds, Seamounts: ecology, fisheries & conservation.
First edn. Oxford, United Kingdom: Blackwell Publishing, pp. 245-251.
THRESHER, R., ALTHAUS, F., ADKINS, J., GOWLETT-HOLMES, K., ALDERSLADE,
P., DOWDNEY, J., CHO, W., GAGNON, A., STAPLES, D., MCENNULTY, F. &
WILLIAMS, A., 2014. Strong depth-related zonation of megabenthos on a rocky
continental margin (∼ 700–4000 m) off southern Tasmania, Australia. PloS ONE,
9(1), pp. e85872.
TOPSENT, E., 1892. Contribution à l’étude des Spongiaires de l’Atlantique Nord (Golfe de
Gascogne, Terre-Neuve, Açores). Résultats des campagnes scientifiques accomplies
par le Prince Albert I. Monaco, 2, pp. 1-165.
TOPSENT, E., 1894a. Etude monographique des spongiaires de France. I.Tetractinellida.
Archives de Zoologie expérimentelle et genérale, 3(2), pp. 259-400.
TOPSENT, E., 1894b. Une réforme dans la classification des Halichondrina. Mémoires de la
Société Zoologique de France, 7, pp. 5-26.
TOPSENT, E., 1918. Éponges de San Thomé. Essai sur les genres Spirastrella, Donatia et
Chondrilla. Archives de Zoologie expérimentelle et genérale, 57(6), pp. 535-618.
TOPSENT, E., 1922. Les mégasclères polytylotes des Monaxonides et la parenté des
Latrunculiines. Bulletin de l'Institut Océanographique, Monaco, 415, pp. 1-8.
TOPSENT, E., 1928. Spongiaires de l'Atlantique et de la Méditerrannée, provenant des
croisières du Prince Albert I er de Monaco. Résultats des campagnes scientifiques
accomplies par le Prince Albert I. Monaco., 74, pp. 1-376.
URIZ, M.J., 2002. Family Ancorinidae Schmidt, 1870. In: J.N.A. HOOPER & R.W.M. VAN
SOEST, eds, Systema Porifera: a guide to the classification of sponges. First edn.
156
New York, Boston, Dordrecht, London, Moscow: Kluwer Academic/Plenum
Publishers, pp. 108-126.
VACELET, J., VASSEUR, P. & LÉVI, C., 1976. Spongiaires de la pente externe des récifs
coralliens de Tuléar (Sud-Ouest de Madagascar). Mémoires du Muséum National
d'Histoire Naturelle, 49, pp. 1-116.
VAN SOEST, R.W.M., 1994. Demosponge distribution patterns, R.W.M. VAN SOEST,
T.M.G. VAN KEMPEN & J.C. BRAEKMAN, eds. In: Sponges in time and space:
biology, chemistry, paleontology, 19 - 23 April 1994, Balkema, pp. 213.
VAN SOEST, R.W.M., 2002a. Family Desmacididae Schmidt, 1870. In: J.N.A. HOOPER &
R.W.M. VAN SOEST, eds, Systema Porifera: a guide to the classification of
sponges. First edn. New York, Boston, Dordrecht, London, Moscow: Kluwer
Academic/Plenum Publishers, pp. 572-574.
VAN SOEST, R.W.M., 2002b. Family Suberitidae Schmidt, 1870. In: J.N.A. HOOPER &
R.W.M. VAN SOEST, eds, Systema Porifera: a guide to the classification of
sponges. First edn. New York, Boston, Dordrecht, London, Moscow: Kluwer
Academic/Plenum Publishers, pp. 227-244.
VAN SOEST, R.W.M., 2007. Sponge biodiversity. Journal of the Marine Biological
Association of the United Kingdom, 87(6), pp. 1345-1348.
VAN SOEST, R.W.M., BOURY-ESNAULT, N., HOOPER, J.N.A., RÜTZLER, K., DE
VOOGD, N.J., ALVAREZ DE GLASBY, B., HAJDU, E., PISERA, A.B.,
MANCONI, R., SCHOENBERG, C., JANUSSEN, D., TABACHNICK, K.R.,
KLAUTAU, M., PICTON, B., KELLY, M., VACELET, J., DOHRMANN, M.,
DÍAZ, M. & CÁRDENAS, P., 2015-last update, World Porifera Database. Available:
http://www.marinespecies.org/porifera2015].
157
VAN SOEST, R.W.M., BOURY-ESNAULT, N., VACELET, J., DOHRMANN, M.,
ERPENBECK, D., DE VOOGD, N.J., SANTODOMINGO, N., VANHOORNE, B.,
KELLY, M. & HOOPER, J.N.A., 2012. Global diversity of sponges (Porifera). PLoS
ONE, 7(4), pp. e35105.
VAN SOEST, R.W.M. & HOOPER, J.N.A., 2002. Family Calthropellidae Lendenfeld,
1907. In: J.N.A. HOOPER & R.W.M. VAN SOEST, eds, Systema Porifera: a guide
to the classification of sponges. First edn. New York, Boston, Dordrecht, London,
Moscow: Kluwer Academic/Plenum Publishers, pp. 127-133.
VERESHCHAKA, A.L., 1995. Macroplankton in the near-bottom layer of continental slopes
and seamounts. Deep Sea Research Part I: Oceanographic Research Papers, 42(9),
pp. 1639-1668.
VERRILL, A.E., 1907. The Bermuda Islands: Part V. An account of the coral reefs
(characteristic life of the Bermuda coral reefs). Porifera: sponges. Transactions of the
Connecticut Academy of Arts and Sciences, 12, pp. 330-344.
VIEIRA, W.F., COSME, B. & HAJDU, E., 2010. Three new Erylus (Demospongiae,
Astrophorida, Geodiidae) from the Almirante Saldanha Seamount (off SE Brazil),
with further data for a tabular review of worldwide species and comments on
Brazilian seamount sponges. Marine Biology Research, 6(5), pp. 437-460.
VOULTSIADOU, E., 2007. Sponges: an historical survey of their knowledge in Greek
antiquity. Journal of the Marine Biological Association of the United Kingdom, 87(6),
pp. 1757-1763.
WAFAR, M., VENKATARAMAN, K., INGOLE, B., KHAN, S.A. & LOKABHARATHI,
P., 2011. State of knowledge of coastal and marine biodiversity of Indian Ocean
countries. PLoS ONE, 6(1), pp. e14613.
158
WATSON, R., KITCHINGMAN, A. & CHEUNG, W.W., 2007. Catches from world
seamount fisheries. In: T.J. PITCHER, T. MORATO, P.J.B. HART, M.R. CLARK,
N. HAGGAN & R.S. SANTOS, eds, Seamounts: ecology, fisheries & conservation.
First edn. Oxford, United Kingdom: Blackwell Publishing, pp. 400-412.
WEAVER, J.C., PIETRASANTA, L.I., HEDIN, N., CHMELKA, B.F., HANSMA, P.K. &
MORSE, D.E., 2003. Nanostructural features of demosponge biosilica. Journal of
Structural Biology, 144(3), pp. 271-281.
WESSEL, P., 2001. Global distribution of seamounts inferred from gridded Geosat/ERS‐1
altimetry. Journal of Geophysical Research, 106(B9), pp. 19431-19441.
WESSEL, P., 2007. Seamount characteristics. In: T.J. PITCHER, T. MORATO, P.J.B.
HART, M.R. CLARK, N. HAGGAN & R.S. SANTOS, eds, Seamounts: ecology,
fisheries & conservation. First edn. Oxford, United Kingdom: Blackwell Publishing,
pp. 3-25.
WESSEL, P., SANDWELL, D.T. & KIM, S., 2010. The global seamount census.
Oceanography, 23(1), pp. 24-33.
WHITE, M., BASHMACHNIKOV, I., ARÍSTEGUI, J. & MARTINS, A., 2007. Physical
processes and seamount productivity. In: T.J. PITCHER, T. MORATO, P.J.B. HART,
M.R. CLARK, N. HAGGAN & R.S. SANTOS, eds, Seamounts: ecology, fisheries &
conservation. First edn. Oxford, United Kingdom: Blackwell Publishing, pp. 65-84.
WILLIAMS, A., ALTHAUS, F. & SCHLACHER, T.A., 2015. Towed camera imagery and
benthic sled catches provide different views of seamount benthic diversity. Limnology
and Oceanography: Methods, 13(2), pp. 62-73.
WILSON, R.R. & KAUFMANN, R.S., 1987. Seamount biota and biogeography. In: B.H.
KEATING, P. FRYER, R. BATIZA & W. BOEHLERT, eds, Seamounts, islands, and
atolls. Washington, D.C.: American Geophysical Union, pp. 355-377.
159
WÖRHEIDE, G., SOLÉ-CAVA, A.M. & HOOPER, J.N.A., 2005. Biodiversity, molecular
ecology and phylogeography of marine sponges: patterns, implications and outlooks.
Integrative and Comparative Biology, 45(2), pp. 377-385.
WORM, B., LOTZE, H.K. & MYERS, R.A., 2003. Predator diversity hotspots in the blue
ocean. Proceedings of the National Academy of Sciences, 100(17), pp. 9884-9888.
WRIGHT, E.P., 1881. On a new genus and species of a sponge (Alemo seychellensis) with
supposed heteromorphic zooids. Transactions of the Royal Irish Academy, 28, pp. 1320.
XAVIER, J.R., TORKILDSEN, M., TANGEN, S., CÁRDENAS, P., HESTETUN, J.,
EILERTSEN, M.H., ALVIZU, A., CARVALHO, F., THORKILDSEN, S., OLSEN,
B.R. & RAPP, H.T., 2015. Sponge assemblages of the Schultz Seamount - Arctic
Mid-Ocean Ridge, 14th Deep-Sea Biology Symposium: abstract book, 31 August - 4
September 2015, UA Editora, pp. 259.
XAVIER, J.R. & VAN SOEST, R.W.M., 2007. Demosponge fauna of Ormonde and
Gettysburg
Seamounts (Gorringe Bank, North-East Atlantic): diversity and
zoogeographical affinities. Journal of the Marine Biological Association of the
United Kingdom, 87(6), pp. 1643-1653.
YESSON, C., CLARK, M.R., TAYLOR, M.L. & ROGERS, A.D., 2011. The global
distribution of seamounts based on 30 arc seconds bathymetry data. Deep Sea
Research Part I: Oceanographic Research Papers, 58(4), pp. 442-453.
ZEZINA, O.N., 2010. Check-list of Holocene brachiopods annotated with geographical
ranges of species. Paleontological Journal, 44(9), pp. 1176-1199.
ZIBROWIUS, H., 1982. Deep-water scleractinian corals from the South-Western Indian
Ocean with crypts excavated by crabs, presumably Hapalocarcinidae. Crustaceana,
43(2), pp. 113-120.
160
Appendix
Table A: Taxonomic sponge species list per ecoregion included in the biogeographical
analyses, compiled from the World Porifera Database (van Soest et al. 2015). Categorisation
follows Spalding et al. (2007), with numbers in the brackets indicating the number of species
recorded per ecoregion. Ecoregions 101 (Bight of Sofala/Swamp Coast) and 217 (Bouvet
Island) were excluded as they had one and zero species recorded respectively. Vema
Seamount is also included for comparison as an associate of the West Wind Drift Islands
Province. Last updated May 2015.
Western Indo-Pacific Realm
20. Western Indian Ocean Province
94. Northern Monsoon Current Coast Ecoregion (44)
Aciculites tulearensis Vacelet & Vasseur, 1965; Amphimedon rubida Pulitzer-Finali, 1993;
Aulospongus flabellum Pulitzer-Finali, 1993; Aulospongus involutus (Kirkpatrick, 1903);
Axinella arborescens Ridley & Dendy, 1886; Axinella donnani (Bowerbank, 1873);
Axinyssa tenax Pulitzer-Finali, 1993; Callipelta thoosa Lévi, 1964; Callyspongia subtilis
Pulitzer-Finali, 1993; Calyx infundibulum Pulitzer-Finali, 1993; Chelotropella sphaerica
Lendenfeld, 1907; Chondrocladia (Chondrocladia) multichela Lévi, 1964; Coelosphaera
(Coelosphaera) crumena Pulitzer-Finali, 1993; Crambe erecta Pulitzer-Finali, 1993; Crella
(Grayella) cyathophora Carter, 1869; Echinodictyum jousseaumi Topsent, 1892; Ecionemia
acervus Bowerbank, 1864; Erylus globulifer Pulitzer-Finali, 1993; Hemiasterella
complicata Topsent, 1919; Hemiasterella intermedia Dendy, 1922; Hemiasterella magna
Pulitzer-Finali, 1993; Higginsia kenyensis Pulitzer-Finali, 1993; Higginsia lamella
Pulitzer-Finali, 1993; Higginsia pulcherrima Pulitzer-Finali, 1993; Jaspis manihinei
Pulitzer-Finali, 1993; Lithoplocamia indica Pulitzer-Finali, 1993; Lithoplocamia
tuberculata Pulitzer-Finali, 1993; Manihinea conferta Pulitzer-Finali, 1993; Oceanapia
exigua Pulitzer-Finali, 1993; Oceanapia fistulosa (Bowerbank, 1873); Oceanapia globosa
Pulitzer-Finali, 1993; Penares intermedia (Dendy, 1905); Petrosia (Petrosia) nigricans
Lindgren, 1897; Phorbas palmatus Pulitzer-Finali, 1993; Spheciospongia inconstans
(Dendy, 1887); Stelletta digitata (Pulitzer-Finali, 1993); Stelletta purpurea Ridley, 1884;
Tabulocalyx pedunculatus Pulitzer-Finali, 1993; Tethyopsis plurima (Pulitzer-Finali, 1993);
Thenea tyla Lendenfeld, 1907; Theonella swinhoei Gray, 1868; Xestospongia clavata
Pulitzer-Finali, 1993; Xestospongia informis Pulitzer-Finali, 1993; Xestospongia tuberosa
Pulitzer-Finali, 1993
95. East African Coral Coast Ecoregion (172)
Acanthostylotella cornuta (Topsent, 1897); Acarnus ternatus Ridley, 1884; Amorphinopsis
foetida (Dendy, 1889); Amphimedon navalis Pulitzer-Finali, 1993; Amphimedon rubiginosa
Pulitzer-Finali, 1993; Amphimedon spinosa Pulitzer-Finali, 1993; Aplysina primitiva
Burton, 1959; Astrosclera willeyana Lister, 1900; Aulospongus involutus (Kirkpatrick,
1903); Axinella aruensis (Hentschel, 1912); Axinella flabelloreticulata Burton, 1959;
Axinella massalis Burton, 1959; Axinella ventilabrum Burton, 1959; Axinella weltnerii
161
(Lendenfeld, 1897); Axinyssa aplysinoides (Dendy, 1922); Axinyssa topsenti Lendenfeld,
1897; Biemna bihamigera (Dendy, 1922); Biemna fistulosa (Topsent, 1897); Biemna fortis
(Topsent, 1897); Biemna humilis Thiele, 1903; Biemna microstrongyla (Hentschel, 1912);
Biemna trirhaphis (Topsent, 1897); Bubaris conulosa Vacelet & Vasseur, 1971;
Callyspongia (Cladochalina) diffusa (Ridley, 1884); Callyspongia (Toxochalina) robusta
(Ridley, 1884); Callyspongia abnormis Pulitzer-Finali, 1993; Callyspongia contorta
Pulitzer-Finali, 1993; Callyspongia hirta Pulitzer-Finali, 1993; Callyspongia perforata
Pulitzer-Finali, 1993; Callyspongia reticulata (Keller, 1889); Callyspongia violacea
Pulitzer-Finali, 1993; Calyx nyaliensis Pulitzer-Finali, 1993; Carteriospongia foliascens
(Pallas, 1766); Chondrilla mixta Schulze, 1877; Chondrilla sacciformis Carter, 1879;
Cinachyrella arabica (Carter, 1869); Cinachyrella lacerata (Bösraug, 1913); Ciocalypta
digitata (Dendy, 1905); Cladocroce tubulosa Pulitzer-Finali, 1993; Clathria (Microciona)
affinis (Carter, 1880); Clathria (Microciona) anonyma (Burton, 1959); Clathria
(Microciona) richmondi Hooper, Kelly & Kennedy, 2000; Coelosphaera (Coelosphaera)
navicelligera (Ridley, 1885); Crella shimonii Pulitzer-Finali, 1993; Diplastrella gardineri
Topsent, 1918; Discodermia discifera (Lendenfeld, 1907); Dragmacidon coccineum
(Keller, 1891); Dragmacidon durissimum (Dendy, 1905); Ecionemia acervus Bowerbank,
1864; Epipolasis suluensis (Wilson, 1925); Erylus lendenfeldi Sollas, 1888; Fangophilina
hirsuta Lendenfeld, 1907; Fascaplysinopsis reticulata (Hentschel, 1912); Fasciospongia
friabilis (Hyatt, 1877); Fasciospongia operculum (Lendenfeld, 1897); Geodia carcinophila
(Lendenfeld, 1897); Geodia crustosa Bösraug, 1913; Geodia pleiades (Sollas, 1888);
Geodia sollasi (Lendenfeld, 1888); Geodia spheranthastra Pulitzer-Finali, 1993;
Halichondria (Halichondria) cartilaginea (Esper, 1794); Halichondria (Halichondria)
lendenfeldi Lévi, 1961; Halichondria (Halichondria) tenuiramosa Dendy, 1922; Haliclona
(Gellius) amboinensis (Lévi, 1961); Haliclona (Gellius) cellaria (Rao, 1941); Haliclona
(Gellius) toxia (Topsent, 1897); Haliclona (Reniera) debilis Pulitzer-Finali, 1993;
Haliclona bawiana (Lendenfeld, 1897); Haliclona cavernosa (Pulitzer-Finali, 1993);
Haliclona cerebrum (Burton, 1928); Haliclona decidua (Topsent, 1906); Haliclona digitata
(Baer, 1906); Haliclona fistulosa (Pulitzer-Finali, 1993); Haliclona irregularis
(Kirkpatrick, 1900); Haliclona mollis (Baer, 1906); Haliclona pigmentifera (Dendy, 1905);
Halisarca ferreus Bergquist & Kelly, 2004; Hemiasterella bouilloni (Thomas, 1973);
Hyalonema (Cyliconema) molle Schulze, 1904; Hyalonema (Prionema) validum Schulze,
1904; Hyattella intestinalis (Lamarck, 1814); Hymedesmia (Hymedesmia) murrayi Burton,
1959; Iotrochota baculifera Ridley, 1884; Iotrochota nigra (Baer, 1906); Iotrochota
purpurea (Bowerbank, 1875); Jaspis sansibarensis (Baer, 1906); Lamellodysidea herbacea
(Keller, 1889); Lendenfeldia plicata (Esper, 1794); Leucandra brumalis Jenkin, 1908;
Leucandrilla wasinensis (Jenkin, 1908); Liosina paradoxa Thiele, 1899; Lissodendoryx
(Lissodendoryx) monticularis Baer, 1906; Lissodendoryx (Waldoschmittia) schmidti (Ridley,
1884); Lithoplocamia minor Pulitzer-Finali, 1993; Monorhaphis chuni Schulze, 1904;
Mycale (Aegogropila) crassissima (Dendy, 1905); Mycale (Aegogropila) sulevoidea (Sollas,
1902); Mycale (Mycale) grandis Gray, 1867; Mycale (Zygomycale) parishii (Bowerbank,
1875); Mycale imperfecta Baer, 1906; Mycale multisclera Pulitzer-Finali, 1993;
Myrmekioderma granulatum (Esper, 1794); Negombata kenyensis (Pulitzer-Finali, 1993);
Negombo kellyae Hooper, 2002; Neopetrosia contignata (Thiele, 1899); Neopetrosia exigua
(Kirkpatrick, 1900); Oceanapia cagayanensis (Wilson, 1925); Oceanapia media (Thiele,
1899); Oceanapia minuta (Vacelet, Vasseur & Lévi, 1976); Oceanapia polysiphonia
(Dendy, 1922); Oceanapia zoologica (Dendy, 1905); Oscarella nigraviolacea Bergquist &
Kelly, 2004; Paratetilla bacca (Selenka, 1867); Penares intermedia (Dendy, 1905); Petrosia
(Petrosia) expansa (Thiele, 1903); Petrosia (Petrosia) seychellensis Dendy, 1922; Petrosia
(Petrosia) shellyi Pulitzer-Finali, 1993; Petrosia (Strongylophora) mauritiana (Carter,
1885); Phakellia radiata (Dendy, 1916); Phakettia ridleyi (Dendy, 1887); Phorbas frutex
Pulitzer-Finali, 1993; Phyllospongia lamellosa (Esper, 1794); Placospongia carinata
(Bowerbank, 1858); Placospongia melobesioides Gray, 1867; Plakinastrella ceylonica
(Dendy, 1905); Plakortis copiosa Pulitzer-Finali, 1993; Plakortis kenyensis Pulitzer-Finali,
1993; Plakortis nigra Lévi, 1953; Platylistrum platessa Schulze, 1904; Polymastia
162
megasclera Burton, 1934; Raspailia colorans Pulitzer-Finali, 1993; Rhabdastrella
globostellata (Carter, 1883); Soleneiscus irregularis (Jenkin, 1908); Spheciospongia
excentrica (Burton, 1931); Spheciospongia florida (Lendenfeld, 1897); Spheciospongia
inconstans (Dendy, 1887); Spheciospongia vagabunda (Ridley, 1884); Spongia (Spongia)
cookii Hyatt, 1877; Spongia (Spongia) hospes (Lendenfeld, 1889); Spongia (Spongia)
mollicula Hyatt, 1877; Spongosorites topsenti Dendy, 1905; Stelletta brevioxea PulitzerFinali, 1993; Stelletta herdmani Dendy, 1905; Stelletta purpurea Ridley, 1884; Stelletta
tulearensis Vacelet, Vasseur & Lévi, 1976; Stellettinopsis laviniensis (Dendy, 1905);
Strongylacidon fasciculatum Pulitzer-Finali, 1993; Strongylacidon sansibarense
Lendenfeld, 1897; Sycettusa simplex (Jenkin, 1908); Sycon munitum Jenkin, 1908; Tedania
(Tedania) conica Baer, 1906; Tedania (Tedania) fragilis Baer, 1906; Tedania (Tedania)
sansibarensis Baer, 1906; Tedania (Tedania) vulcani Lendenfeld, 1897; Tethya
globostellata Lendenfeld, 1897; Tethya parvistella (Baer, 1906); Tethya seychellensis
(Wright, 1881); Tetilla globosa (Baer, 1906); Tetilla sansibarica (Lendenfeld, 1907);
Tetrapocillon minor Pulitzer-Finali, 1993; Thenea malindiae Lendenfeld, 1907; Thenea
pendula Lendenfeld, 1907; Thenea rotunda Lendenfeld, 1907; Theonella conica
(Kieschnick, 1896); Theonella swinhoei Gray, 1868; Timea spherastraea Burton, 1959;
Timea tethyoides Burton, 1959; Topsentia halichondrioides (Dendy, 1905); Topsentia
megalorrhapis (Carter, 1881); Topsentia salomonensis (Dendy, 1922); Xestospongia
testudinaria (Lamarck, 1815); Zyzzya fuliginosa (Carter, 1879)
96. Seychelles Ecoregion (147)
Acanthella cavernosa Dendy, 1922; Acanthostylotella cornuta (Topsent, 1897);
Acanthotetilla seychellensis (Thomas, 1973); Acarnus bicladotylotus Hoshino, 1981;
Acarnus ternatus Ridley, 1884; Acarnus topsenti Dendy, 1922; Agelas ceylonica Dendy,
1905; Astrosclera willeyana Lister, 1900; Aulospongus gardineri (Dendy, 1922); Axinella
donnani (Bowerbank, 1873); Axinella minor Thomas, 1981; Axinella proliferans Ridley,
1884; Axinyssa aplysinoides (Dendy, 1922); Biemna bihamigera (Dendy, 1922); Biemna
fortis (Topsent, 1897); Biemna seychellensis Thomas, 1973; Biemna trirhaphis (Topsent,
1897); Biemna tubulata (Dendy, 1905); Callyspongia (Callyspongia) differentiata (Dendy,
1922); Callyspongia (Callyspongia) reticutis (Dendy, 1905); Carteriospongia foliascens
(Pallas, 1766); Chalinula camerata (Ridley, 1884); Chalinula confusa (Dendy, 1922);
Chondrilla australiensis Carter, 1873; Chondrocladia (Chondrocladia) clavata Ridley &
Dendy, 1886; Cinachyrella australiensis (Carter, 1886); Clathria (Clathria) decumbens
Ridley, 1884; Clathria (Clathria) maeandrina Ridley, 1884; Clathria (Clathria) spongodes
Dendy, 1922; Clathria (Thalysias) amirantiensis Hooper, 1996; Clathria (Thalysias)
procera (Ridley, 1884); Clathria (Thalysias) robusta (Dendy, 1922); Clathria (Thalysias)
vulpina (Lamarck, 1814); Coelosphaera (Coelosphaera) ramosa (Dendy, 1922); Cornulella
amirantensis van Soest, Zea & Kielman, 1994; Cornulella lundbecki Dendy, 1922;
Cornulella tyro van Soest, Zea & Kielman, 1994; Crambe acuata (Lévi, 1958); Crella
(Grayella) cyathophora Carter, 1869; Cyamon vickersii (Bowerbank, 1864); Damiria
toxifera van Soest, Zea & Kielman, 1994; Dictyodendrilla pallasi (Ridley, 1884); Didiscus
aceratus (Ridley & Dendy, 1886); Discodermia laevidiscus Carter, 1880; Dragmacidon
durissimum (Dendy, 1905); Dragmacidon durissimum var. massale (Dendy, 1922); Dysidea
gumminea Ridley, 1884; Ecionemia acervus Bowerbank, 1864; Erylus cylindriger Ridley,
1884; Erylus lendenfeldi Sollas, 1888; Euplectella cucumer Owen, 1857; Eurypon encrusta
(Thomas, 1981); Fasciospongia seychellensis (Thomas, 1973); Forcepia (Forcepia)
stephensi Dendy, 1922; Geodia auroristella Dendy, 1916; Geodia lindgreni (Lendenfeld,
1903); Geodia micraster (Lendenfeld, 1907); Halichondria (Halichondria) aldabrensis Lévi,
1961; Halichondria (Halichondria) lendenfeldi Lévi, 1961; Haliclona (Haliclona)
cribriformis (Ridley, 1884); Haliclona (Reniera) cribricutis (Dendy, 1922); Haliclona
(Reniera) tufoides (Dendy, 1922); Hemiasterella bouilloni (Thomas, 1973); Hemiasterella
intermedia Dendy, 1922; Higginsia fragilis Lévi, 1961; Higginsia higgini Dendy, 1922;
163
Higginsia petrosioides Dendy, 1922; Hyalonema (Cyliconema) madagascarense (Lévi,
1964); Hyattella sinuosa (Pallas, 1766); Hymedesmia (Hymedesmia) prostrata Thiele, 1903;
Hymeniacidon proteus (Ridley, 1884); Hymeniacidon variospiculata Dendy, 1922; Hyrtios
erectus (Keller, 1889); Igernella mirabilis Lévi, 1961; Iotrochota baculifera Ridley, 1884;
Iotrochota purpurea (Bowerbank, 1875); Jaspis penetrans (Carter, 1880); Leucaltis
nodusgordii (Poléjaeff, 1883); Leucandra anguinea (Ridley, 1884); Leucandra seychellensis
Hozawa, 1940; Leucetta chagosensis Dendy, 1913; Levinella prolifera (Dendy, 1913);
Liosina paradoxa Thiele, 1899; Lithoplocamia lithistoides Dendy, 1922; Microscleroderma
herdmani (Dendy, 1905); Monanchora unguiculata (Dendy, 1922); Mycale (Aegogropila)
crassissima (Dendy, 1905); Mycale (Grapelia) vansoesti Hajdu, 1995; Mycale (Mycale)
gelatinosa (Ridley, 1884); Myrmekioderma granulatum (Esper, 1794); Myxilla (Myxilla)
seychellensis Thomas, 1981; Neopetrosia retiderma (Dendy, 1922); Oceanapia fistulosa
(Bowerbank, 1873); Oceanapia pellucida (Ridley, 1884); Oceanapia seychellensis (Dendy,
1922); Oceanapia toxophila Dendy, 1922; Paraleucilla proteus (Dendy, 1913); Pericharax
orientalis Van Soest & De Voogd, 2015; Petrosia (Petrosia) nigricans Lindgren, 1897;
Petrosia (Strongylophora) durissima (Dendy, 1905); Phakellia radiata (Dendy, 1916);
Phlyctaenopora (Barbozia) primitiva (Dendy, 1922); Phorbas clathrodes (Dendy, 1922);
Phorbas papillatus (Dendy, 1922); Phyllospongia alcicornis (Esper, 1794); Phyllospongia
supraoculata Ridley, 1884; Plakinastrella minor (Dendy, 1916); Rhabdastrella cribriporosa
(Dendy, 1916); Rhabdastrella globostellata (Carter, 1883); Rhabdastrella oxytoxa
(Thomas, 1973); Rhabdastrella providentiae (Dendy, 1916); Rhabdastrella rowi (Dendy,
1916); Rhabderemia bistylifera Lévi, 1961; Siphonodictyon minutum (Thomas, 1973);
Spheciospongia globularis (Dendy, 1922); Spheciospongia inconstans (Dendy, 1887);
Spheciospongia inconstans var. digitata (Dendy, 1887); Spheciospongia transitoria (Ridley,
1884); Spirastrella decumbens Ridley, 1884; Spirastrella pachyspira Lévi, 1958;
Spongionella retiara (Dendy, 1916); Spongosorites niger (Dendy, 1922); Stelletta cylindrica
Thomas, 1973; Stelletta jonesi (Thomas, 1973); Stelletta purpurea Ridley, 1884;
Stellettinopsis cherbonnieri Lévi, 1961; Stellettinopsis laviniensis (Dendy, 1905);
Strongylamma wilsoni (Dendy, 1922); Stylissa carteri (Dendy, 1889); Stylissa conulosa
(Dendy, 1922); Stylissa massa (Carter, 1887); Terpios cruciata (Dendy, 1905); Tethya
japonica Sollas, 1888; Tethya peracuta (Topsent, 1918); Tethya robusta (Bowerbank,
1873); Tethya seychellensis (Wright, 1881); Tethya stellagrandis (Dendy, 1916); Theonella
complicata (Carter, 1880); Theonella conica (Kieschnick, 1896); Theonella swinhoei Gray,
1868; Thoosa radiata Topsent, 1887; Thrombus ornatus Sollas, 1888; Timea anthastra Lévi,
1961; Timea curvistellifera (Dendy, 1905); Topsentia stellettoides (Lévi, 1961);
Xestospongia testudinaria (Lamarck, 1815); Zyzzya fuliginosa (Carter, 1879)
97. Cargados Carajos/Tromelin Island Ecoregion (27)
Acanthella calyx (Dendy, 1922); Acarnus topsenti Dendy, 1922; Auletta lyrata var.
brevispiculata Dendy, 1905; Aulocalyx serialis Dendy, 1916; Axinyssa aplysinoides (Dendy,
1922); Clathria (Clathria) whiteleggii Dendy, 1922; Clathria (Thalysias) lendenfeldi Ridley
& Dendy, 1886; Clathria (Thalysias) procera (Ridley, 1884); Dictyonella conglomerata
(Dendy, 1922); Didiscus placospongioides Dendy, 1922; Discodermia tuberosa Dendy,
1922; Dragmacidon durissimum var. erectum (Dendy, 1922); Dragmacidon durissimum var.
tethyoides (Dendy, 1922); Erylus proximus Dendy, 1916; Grantia indica Dendy, 1913;
Hemigellius calyx var. indica (Dendy, 1922); Hymedesmia (Stylopus) dendyi Burton, 1930;
Leucetta pyriformis Dendy, 1913; Monanchora lipochela (Dendy, 1922); Neopetrosia
tuberosa (Dendy, 1922); Oceanapia porosa (Dendy, 1922); Paracornulum strepsichela
(Dendy, 1922); Petrosia (Petrosia) mammiformis Dendy, 1922; Phorbas clathrodes (Dendy,
1922); Plakinastrella minor (Dendy, 1916); Stelletta cavernosa (Dendy, 1916); Stylissa
conulosa (Dendy, 1922)
98. Mascarene Islands Ecoregion (35)
164
Agelas mauritiana (Carter, 1883); Chondrilla sacciformis Carter, 1879; Clathrina
compacta (Schuffner, 1877); Cliona jullieni Topsent, 1891; Dysidea enormis (Hyatt, 1877);
Dysidea spinosa (Hyatt, 1877); Echinodictyum pykii (Carter, 1879); Eurypon cactoides
(Burton & Rao, 1932); Fasciospongia pikei (Hyatt, 1877); Heterotella corbicula
(Bowerbank, 1862); Hippospongia mauritiana (Hyatt, 1877); Hyattella intestinalis
(Lamarck, 1814); Ircinia intertexta (Hyatt, 1877); Laocoetis perion Lévi, 1986; Leucaltis
mauritiana Schuffner, 1877; Leucandra claviformis Schuffner, 1877; Leucandra echinata
Schuffner, 1877; Leucandra falcigera Schuffner, 1877; Lithoplocamia lithistoides Dendy,
1922; Monanchora laevissima (Dendy, 1922); Mycale (Zygomycale) parishii (Bowerbank,
1875); Petrosia (Strongylophora) mauritiana (Carter, 1885); Phlyctaenopora (Barbozia)
primitiva (Dendy, 1922); Phyllospongia lamellosa (Esper, 1794); Polymastia tubulifera
Dendy, 1922; Raspailia laciniata (Carter, 1879); Rhaphidhistia spectabilis Carter, 1879;
Sigmosceptrella quadrilobata Dendy, 1922; Spongia (Spongia) hispida Lamarck, 1814;
Spongia (Spongia) irregularis (Lendenfeld, 1889); Stelletta mauritiana (Dendy, 1916);
Stelletta purpurea Ridley, 1884; Stylissa massa (Carter, 1887); Sycettusa sycilloides
(Schuffner, 1877); Sycon tabulatum (Schuffner, 1877)
99. Southeast Madagascar Ecoregion (4)
Geodia crustosa Bösraug, 1913; Geodia piriformis Bösraug, 1913; Geodia poculata
Bösraug, 1913; Spongia (Spongia) hispida Lamarck, 1814
100. Western and Northern Madagascar Ecoregion (150)
Acanthancora stylifera Burton, 1959; Acanthostylotella cornuta (Topsent, 1897);
Acanthotriaena crypta Vacelet, Vasseur & Lévi, 1976; Acarnus bergquistae van Soest,
Hooper & Hiemstra, 1991; Acarnus wolffgangi Keller, 1889; Aciculites spinosa Vacelet &
Vasseur, 1971; Aciculites tulearensis Vacelet & Vasseur, 1965; Agelas bispiculata Vacelet,
Vasseur & Lévi, 1976; Agelas marmarica Lévi, 1958; Agelas mauritiana (Carter, 1883);
Alectona primitiva Topsent, 1932; Amorphinopsis fistulosa (Vacelet, Vasseur & Lévi,
1976); Ancorina nanosclera Lévi, 1967; Astrosclera willeyana Lister, 1900; Aulospongus
gardineri (Dendy, 1922); Axinyssa aplysinoides (Dendy, 1922); Batzella aurantiaca (Lévi,
1958); Biemna anisotoxa Lévi, 1963; Biemna bihamigera (Dendy, 1922); Biemna laboutei
Hooper, 1996; Bubaris conulosa Vacelet & Vasseur, 1971; Callipelta cavernicola (Vacelet
& Vasseur, 1965); Callipelta mixta Vacelet, Vasseur & Lévi, 1976; Callipelta ornata
Sollas, 1888; Callyspongia (Toxochalina) robusta (Ridley, 1884); Carteriospongia foliascens
(Pallas, 1766); Carteriospongia pennatula Ridley, 1884; Chondrilla australiensis Carter,
1873; Chondrilla mixta Schulze, 1877; Chondrilla sacciformis Carter, 1879; Chondropsis
lamella (Lendenfeld, 1888); Chondrosia debilis Thiele, 1900; Cinachyrella australiensis
(Carter, 1886); Cinachyrella schulzei (Keller, 1891); Ciocalypta microstrongylata Vacelet,
Vasseur & Lévi, 1976; Cladorhiza nematophora Lévi, 1964; Clathria (Clathria) foliascens
Vacelet & Vasseur, 1971; Clathria (Clathria) spongodes Dendy, 1922; Clathria
(Microciona) microxea (Vacelet & Vasseur, 1971); Clathria (Microciona) vacelettia
Hooper, 1996; Clathria (Thalysias) abietina (Lamarck, 1814); Clathria (Thalysias) vulpina
(Lamarck, 1814); Clathria (Wilsonella) cercidochela (Vacelet & Vasseur, 1971); Clathria
dichela sensu Vacelet, Vasseur & Lévi, 1976; Cliona mucronata Sollas, 1878;
Coelodischela diatomorpha Vacelet, Vasseur & Lévi, 1976; Cornulella minima (Vacelet,
Vasseur & Lévi, 1976); Crambe acuata (Lévi, 1958); Diacarnus globosus (Vacelet,
Vasseur & Lévi, 1976); Didiscus aceratus (Ridley & Dendy, 1886); Didiscus anisodiscus
Vacelet & Vasseur, 1971; Didiscus placospongioides Dendy, 1922; Discodermia dubia
Vacelet & Vasseur, 1971; Discodermia japonica Döderlein, 1884; Discodermia panoplia
Sollas, 1888; Echinodictyum jousseaumi Topsent, 1892; Ecionemia cinerea Thiele, 1900;
165
Erylus lendenfeldi Sollas, 1888; Farrea occa Bowerbank, 1862; Farrea occa occa
Bowerbank, 1862; Gelliodes flagellifera Vacelet, Vasseur & Lévi, 1976; Gelliodes
nossibea Lévi, 1956; Gelliodes petrosioides Dendy, 1905; Geodia carcinophila (Lendenfeld,
1897); Geodia composita Bösraug, 1913; Geodia peruncinata Dendy, 1905; Geodia sollasi
(Lendenfeld, 1888); Geodia sphaerulifer (Vacelet & Vasseur, 1965); Haliclona (Gellius)
cymaeformis (Esper, 1794); Haliclona (Gellius) friabilis (Lévi, 1956); Haliclona (Gellius)
ridleyi (Hentschel, 1912); Haliclona (Halichoclona) cioniformis (Lévi, 1956); Haliclona
fragilis (Vacelet, Vasseur & Lévi, 1976); Haliclona madagascarensis Vacelet, Vasseur &
Lévi, 1976; Haliclona polypoides (Vacelet, Vasseur & Lévi, 1976); Haliclona striata
Vacelet, Vasseur & Lévi, 1976; Haliclona tulearensis Vacelet, Vasseur & Lévi, 1976;
Halisarca ectofibrosa Vacelet, Vasseur & Lévi, 1976; Hemiasterella complicata Topsent,
1919; Hemiasterella strongylophora Lévi, 1956; Higginsia petrosioides Dendy, 1922;
Hippospongia laxa Lendenfeld, 1889; Homophymia lamellosa Vacelet & Vasseur, 1971;
Hyrtios cavernosus (Vacelet, Vasseur & Lévi, 1976); Igernella mirabilis Lévi, 1961;
Iotrochota purpurea (Bowerbank, 1875); Ircinia conulosa (Ridley, 1884); Ircinia
cylindracea Vacelet, Vasseur & Lévi, 1976; Jaspis diastra (Vacelet & Vasseur, 1965);
Kaliapsis incrustans (Vacelet & Vasseur, 1971); Lelapiella incrustans Vacelet, 1977;
Lepidoleucon inflatum Vacelet, 1967; Liosina arenosa (Vacelet & Vasseur, 1971);
Monanchora unguiculata (Dendy, 1922); Mycale (Carmia) microxea Vacelet, Vasseur &
Lévi, 1976; Mycale (Grapelia) vaceleti Hajdu, 1995; Mycale (Mycale) gravelyi Burton,
1937; Mycale (Naviculina) cleistochela Vacelet & Vasseur, 1971; Mycale (Naviculina)
flagellifera Vacelet & Vasseur, 1971; Mycale (Zygomycale) parishii (Bowerbank, 1875);
Mycale imperfecta Baer, 1906; Myrmekioderma granulatum (Esper, 1794); Oceanapia
cribrirhina (Vacelet & Vasseur, 1971); Oceanapia dura (Vacelet & Vasseur, 1971);
Oceanapia incrustata (Dendy, 1922); Oceanapia minuta (Vacelet, Vasseur & Lévi, 1976);
Oceanapia mucronata (Vacelet, Vasseur & Lévi, 1976); Oscarella ochreacea Muricy &
Pearse, 2004; Paracornulum strepsichela (Dendy, 1922); Paramurrayona corticata Vacelet,
1967; Petrosia (Petrosia) microxea (Vacelet, Vasseur & Lévi, 1976); Phakellia labellum
(Lamarck, 1814); Phorbas scabida (sensu Vacelet, Vasseur & Lévi, 1976); Phyllospongia
papyracea (Esper, 1794); Pione margaritiferae (Dendy, 1905); Plakina corticioides Vacelet,
Vasseur & Lévi, 1976; Plakinastrella ceylonica (Dendy, 1905); Plectroninia minima
Vacelet, 1967; Plectroninia pulchella Vacelet, 1967; Plectroninia radiata Vacelet, 1967;
Plectroninia tecta Vacelet, 1967; Plectroninia vasseuri Vacelet, 1967; Rhabdocalyptus
monstraster Tabachnick, 1994; Scopalina rubra (Vacelet & Vasseur, 1971);
Sigmosceptrella quadrilobata Dendy, 1922; Spheciospongia florida (Lendenfeld, 1897);
Spheciospongia inconstans (Dendy, 1887); Spheciospongia poterionides (Vacelet &
Vasseur, 1971); Spirastrella decumbens Ridley, 1884; Spirastrella pachyspira Lévi, 1958;
Spirorhabdia alata Vacelet, Vasseur & Lévi, 1976; Spongosorites hentscheli Lévi, 1956;
Stelletta discolor Bösraug, 1913; Stelletta osculifera (Lévi, 1964); Stelletta tulearensis
Vacelet, Vasseur & Lévi, 1976; Stelletta variohamata Thiele, 1900; Strongylamma arenosa
(Vacelet & Vasseur, 1971); Stylissa carteri (Dendy, 1889); Terpios granulosa Bergquist,
1967; Tetilla ridleyi Sollas, 1888; Tetrapocillon minor Pulitzer-Finali, 1993; Theonella
conica (Kieschnick, 1896); Theonella swinhoei Gray, 1868; Thorecta madagascarensis
Lendenfeld, 1889; Timea curvistellifera (Dendy, 1905); Tulearinia stylifera Vacelet, 1977;
Vaceletia crypta (Vacelet, 1977); Xestospongia testudinaria (Lamarck, 1815); Xestospongia
viridenigra (Vacelet, Vasseur & Lévi, 1976); Zyzzya fuliginosa (Carter, 1879)
101. Bight of Sofala/Swamp Coast Ecoregion (1) (excluded)
Axinella tenuidigitata var. oxeata Thomas, 1979
102. Delagoa Ecoregion (34)
Acanthotetilla enigmatica (Lévi, 1964); Amorphinopsis foetida (Dendy, 1889); Ancorina
166
corticata Lévi, 1964; Astrosclera willeyana Lister, 1900; Auletta elongata Dendy, 1905;
Axinella donnani (Bowerbank, 1873); Axinella tenuidigitata Dendy, 1905; Callipelta thoosa
Lévi, 1964; Callyspongia (Cladochalina) diffusa (Ridley, 1884); Chondrilla australiensis
Carter, 1873; Clathria (Clathria) indica Dendy, 1889; Clathria (Clathria) inhacensis
Thomas, 1979; Clathria (Thalysias) vulpina (Lamarck, 1814); Cliona mucronata Sollas,
1878; Coelosphaera (Coelosphaera) navicelligera (Ridley, 1885); Coelosphaera
(Coelosphaera) solenoidea (Lévi, 1964); Dragmacidon agariciforme (Dendy, 1905);
Dysidea gumminea Ridley, 1884; Echinoclathria rimosa (Ridley, 1884); Fasciospongia
benoiti (Thomas, 1979); Hyrtios erectus (Keller, 1889); Iotrochota baculifera Ridley, 1884;
Liosina paradoxa Thiele, 1899; Phakettia ridleyi (Dendy, 1887); Pione margaritiferae
(Dendy, 1905); Rhabdastrella actinosa (Lévi, 1964); Rhabdastrella rowi (Dendy, 1916);
Spheciospongia inconstans (Dendy, 1887); Spirastrella punctulata Ridley, 1884; Stelletta
freitasi Lévi, 1964; Stelletta osculifera (Lévi, 1964); Stelletta purpurea Ridley, 1884;
Terpios cruciata (Dendy, 1905); Tethya robusta (Bowerbank, 1873)
Temperate South America Realm
49. Tristan Gough Province
189. Tristan Gough Ecoregion (21)
Amphilectus rugosus (Thiele, 1905); Amphoriscus gastrorhabdifer (Burton, 1932); Antho
(Acarnia) simplicissima (Burton, 1932); Axinyssa paradoxa (Ridley & Dendy, 1886);
Bubaris murrayi Topsent, 1913; Caulocalyx tener Schulze, 1886; Caulophacus
(Caulodiscus) polyspicula Tabachnick, 1990; Ceratopsion incrustans (Burton, 1932);
Clathria (Clathria) discreta (Thiele, 1905); Clathria (Microciona) antarctica (Topsent,
1917); Desmacella suberitoides (Burton, 1932); Gelliodes licheniformis (Lamarck, 1814);
Haliclona petrosioides Burton, 1932; Hexactinella divergens Tabachnick, 1990;
Hyalonema (Leptonema) campanula longispicula Tabachnick, 1990; Hyrtios altus
(Poléjaeff, 1884); Leucascus leptoraphis (Jenkin, 1908); Leucetta homoraphis Poléjaeff,
1883; Pericharax carteri Poléjaeff, 1883; Poecillastra incrustans Sollas, 1888;
Pseudosuberites exalbicans Topsent, 1913
Temperate Southern Africa Realm
50. Benguela Province
190. Namib Ecoregion (excluded)
191. Namaqua Ecoregion (138)
Aaptos alphiensis Samaai & Gibbons, 2005; Amphilectus informis (Stephens, 1915);
Antho (Acarnia) kellyae Samaai & Gibbons, 2005; Aplysina minuta Lendenfeld, 1889;
Artemisina vulcani Lévi, 1963; Biemna anisotoxa Lévi, 1963; Biemna megalosigma var.
sigmodragma Lévi, 1963; Biemna polyphylla Lévi, 1963; Biemna rhabdostyla Uriz,
1988; Callyspongia (Callyspongia) tubulosa sensu (Esper, 1797); Callyspongia hospitalis
(Stephens, 1915); Clathria (Axosuberites) benguelaensis Samaai & Gibbons, 2005;
Clathria (Clathria) axociona Lévi, 1963; Clathria (Clathria) conica Lévi, 1963; Clathria
(Clathria) dayi Lévi, 1963; Clathria (Clathria) hexagonopora Lévi, 1963; Clathria
(Clathria) omegiensis Samaai & Gibbons, 2005; Clathria (Clathria) pachystyla Lévi,
1963; Clathria (Clathria) parva Lévi, 1963; Clathria (Clathria) rhaphidotoxa Stephens,
1915; Clathria (Isociella) oudekraalensis Samaai & Gibbons, 2005; Clathria
(Microciona) ixauda (Lévi, 1969); Clathria (Microciona) namibiensis (Uriz, 1984);
Clathria (Microciona) stephensae Hooper, 1996; Clathria (Microciona) tenuis (Stephens,
1915); Clathria (Microciona) urizae Hooper, 1996; Clathria (Thalysias) hooperi Samaai
& Gibbons, 2005; Clathria (Thalysias) lissoclada (Burton, 1934); Crambe acuata (Lévi,
1958); Craniella australis Samaai & Gibbons, 2005; Craniella cranium (Müller, 1776);
Desmacidon clavatum Lévi, 1969; Echinochalina (Echinochalina) isochelifera (Uriz,
167
1988); Echinoclathria dichotoma (Lévi, 1963); Echinodictyum macroxiphera Lévi, 1969;
Ectyonopsis flabellata (Lévi, 1963); Ectyonopsis pluridentata (Lévi, 1963); Erylus
amorphus Burton, 1926; Erylus gilchristi Burton, 1926; Eurypon fulvum Lévi, 1969;
Eurypon miniaceum Thiele, 1905; Fibulia ramosa (Ridley & Dendy, 1886); Forcepia
(Leptolabis) australis (Lévi, 1963); Gelliodes coscinopora Lévi, 1969; Geodia libera
Stephens, 1915; Geodia littoralis Stephens, 1915; Guitarra indica Dendy, 1916;
Halichondria (Halichondria) capensis Samaai & Gibbons, 2005; Halichondria
(Halichondria) gilvus Samaai & Gibbons, 2005; Haliclona (Gellius) glacialis (Ridley &
Dendy, 1886); Haliclona (Gellius) jorii (Uriz, 1984); Haliclona (Haliclona) anonyma
(Stephens, 1915); Haliclona (Haliclona) stilensis Burton, 1933; Haliclona (Reniera)
ciocalyptoides Burton, 1933; Haliclona saldanhae (Stephens, 1915); Haliclona stephensi
Burton, 1932; Haliclona submonilifera Uriz, 1988; Haliclonissa sacciformis Burton,
1932; Halisarca pachyderma Lévi, 1969; Hamacantha (Vomerula) esperioides Ridley &
Dendy, 1886; Hexadella kirkpatricki Burton, 1926; Hymedesmia (Hymedesmia)
aurantiaca Lévi, 1963; Hymedesmia (Hymedesmia) parva Stephens, 1915; Hymenancora
tenuissima (Thiele, 1905); Hymeniacidon stylifera (Stephens, 1915); Hymeniacidon
sublittoralis Samaai & Gibbons, 2005; Inflatella belli (Kirkpatrick, 1907); Iophon
cheliferum Ridley & Dendy, 1886; Isodictya alata (Stephens, 1915); Isodictya
chichatouzae Uriz, 1984; Isodictya compressa (Esper, 1794); Isodictya conulosa (Ridley
& Dendy, 1886); Isodictya ectofibrosa (Lévi, 1963); Isodictya elastica (Vosmaer, 1880);
Isodictya frondosa (Lévi, 1963); Isodictya multiformis (Stephens, 1915); Latrunculia
(Biannulata) lunaviridis Samaai, Gibbons, Kelly & Davies-Coleman, 2003; Latrunculia
(Latrunculia) brevis Ridley & Dendy, 1886; Lissodendoryx (Anomodoryx)
coralgardeniensis Samaai & Gibbons, 2005; Lissodendoryx (Lissodendoryx) digitata
(Ridley & Dendy, 1886); Lissodendoryx (Lissodendoryx) simplex (Baer, 1906); Mycale
(Aegogropila) tapetum Samaai & Gibbons, 2005; Mycale (Carmia) levii Samaai &
Gibbons, 2005; Mycale (Carmia) pulvinus Samaai & Gibbons, 2005; Mycale (Mycale)
anisochela Lévi, 1963; Mycale (Mycale) massa (Schmidt, 1862); Mycale (Mycale) trichela
Lévi, 1963; Mycale (Oxymycale) stephensae Samaai & Gibbons, 2005; Mycale
(Paresperella) atlantica (Stephens, 1917); Mycale (Paresperella) curvisigma Lévi, 1969;
Mycale (Paresperella) levii (Uriz, 1987); Mycale (Paresperella) toxifera (Lévi, 1963);
Mycale diastrophochela Lévi, 1969; Myxilla (Burtonanchora) sigmatifera (Lévi, 1963);
Myxilla (Ectyomyxilla) chilensis Thiele, 1905; Myxilla (Ectyomyxilla) kerguelensis
(Hentschel, 1914); Oceanapia atlantica Lévi, 1969; Paracornulum coherens Lévi, 1963;
Penares sphaera (Lendenfeld, 1907); Petrosia (Strongylophora) vulcaniensis Samaai &
Gibbons, 2005; Phorbas bardajii (Uriz, 1988); Phorbas benguelensis (Uriz, 1984);
Phorbas dayi (Lévi, 1963); Phorbas lamellatus (Lévi, 1963); Phorbas pustulosus (Carter,
1882); Plocamiancora walvisensis (Uriz, 1988); Plocamionida ambigua (Bowerbank,
1866); Poecillastra compressa (Bowerbank, 1866); Polymastia atlantica Samaai &
Gibbons, 2005; Polymastia bouryesnaultae Samaai & Gibbons, 2005; Polymastia
infrapilosa Topsent, 1927; Polymastia isidis Thiele, 1905; Polymastia littoralis Stephens,
1915; Protosuberites hendricksi Samaai & Gibbons, 2005; Pseudosuberites hyalinus
(Ridley & Dendy, 1886); Raspailia (Hymeraphiopsis) irregularis Hentschel, 1914;
Raspailia (Raspailia) urizae Hooper, 2012; Rossella antarctica Carter, 1872;
Smenospongia nuda (Lévi, 1969); Spongia (Spongia) brunnea Lévi, 1969; Spongia
(Spongia) violacea Lévi, 1969; Stelletta agulhana Lendenfeld, 1907; Stelletta farcimen
Lendenfeld, 1907; Stelletta rugosa Burton, 1926; Stelletta sphaerica Burton, 1926;
Stelletta trisclera Lévi, 1967; Strongylodesma areolata Lévi, 1969; Suberea pedunculata
(Lévi, 1969); Tedania (Tedania) brondstedi Burton, 1936; Tedania (Tedania) scotiae
Stephens, 1915; Tedania (Tedania) stylonychaeta Lévi, 1963; Tedania (Tedania)
tubulifera Lévi, 1963; Tethya rubra Samaai & Gibbons, 2005; Tetilla capillosa Lévi,
1967; Tetilla casula (Carter, 1871); Trachycladus spinispirulifer (Carter, 1879);
Tsitsikamma scurra Samaai, Gibbons, Kelly & Davies-Coleman, 2003; Xestospongia
hispida (Ridley & Dendy, 1886)
168
51. Agulhas Province
192. Agulhas Bank Ecoregion (131)
Acanthascus (Rhabdocalyptus) baculifer (Schulze, 1904); Acarnus claudei van Soest,
Hooper & Hiemstra, 1991; Alectona wallichii (Carter, 1874); Amphilectus informis
(Stephens, 1915); Amphiute lepadiformis Borojevic, 1967; Amphoriscus kryptoraphis
Urban, 1908; Ancorina corticata Lévi, 1964; Aplysina capensis Carter, 1875; Arthuria
africana (Klautau & Valentine, 2003); Arthuria hirsuta (Klautau & Valentine, 2003);
Biemna anisotoxa Lévi, 1963; Biemna pedonculata Lévi, 1963; Callyspongia
(Cladochalina) foliacea (Esper, 1797); Caulophacus (Caulophacus) basispinosus Lévi,
1964; Caulophacus (Caulophacus) galatheae Lévi, 1964; Ceratopsion microxephora
(Kirkpatrick, 1903); Chelotropella sphaerica Lendenfeld, 1907; Cinachyrella hamata
(Lendenfeld, 1907); Ciocalypta tyleri Bowerbank, 1873; Cladorhiza ephyrula Lévi, 1964;
Clathria (Clathria) elastica Lévi, 1963; Clathria (Clathria) lobata Vosmaer, 1880;
Clathria (Clathria) zoanthifera Lévi, 1963; Clathria (Thalysias) delaubenfelsi (Lévi,
1963); Clathria (Thalysias) flabellata (Burton, 1936); Clathria (Thalysias) nervosa (Lévi,
1963); Clathria (Thalysias) oxitoxa Lévi, 1963; Clathrina cordata (Haeckel, 1872);
Clathrina rotunda Klautau & Valentine, 2003; Crambe acuata (Lévi, 1958); Craniella
metaclada (Lendenfeld, 1907); Crella (Grayella) erecta Lévi, 1963; Crella caespes
(Ehlers, 1870); Cyclacanthia bellae (Samaai, Gibbons, Kelly & Davies-Coleman, 2003);
Echinoclathria dichotoma (Lévi, 1963); Ectyonopsis flabellata (Lévi, 1963); Erylus
polyaster Lendenfeld, 1907; Esperiopsis papillata (Vosmaer, 1880); Fibulia ramosa
(Ridley & Dendy, 1886); Forcepia (Forcepia) agglutinans Burton, 1933; Geodia gallica
(Lendenfeld, 1907); Geodia globosa (Baer, 1906); Geodia perarmata Bowerbank, 1873;
Geodia robusta Lendenfeld, 1907; Geodia stellata Lendenfeld, 1907; Grantessa ramosa
(Haeckel, 1872); Grantessa rarispinosa Borojevic, 1967; Grantia socialis Borojevic,
1967; Guitarra indica Dendy, 1916; Haliclona (Gellius) glacialis (Ridley & Dendy,
1886); Haliclona (Haliclona) stilensis Burton, 1933; Haliclona (Reniera) ciocalyptoides
Burton, 1933; Haliclona simplicissima (Burton, 1933); Hamacantha (Vomerula)
esperioides Ridley & Dendy, 1886; Heteropia glomerosa (Bowerbank, 1873); Higginsia
bidentifera (Ridley & Dendy, 1886); Homaxinella flagelliformis (Ridley & Dendy, 1886);
Hymedesmia (Hymedesmia) aurantiaca Lévi, 1963; Hymeniacidon kerguelensis var.
capensis Hentschel, 1914; Iophon cheliferum Ridley & Dendy, 1886; Isodictya conulosa
(Ridley & Dendy, 1886); Isodictya ectofibrosa (Lévi, 1963); Isodictya elastica (Vosmaer,
1880); Isodictya foliata (Carter, 1885); Isodictya grandis (Ridley & Dendy, 1886);
Isodictya lenta (Vosmaer, 1880); Isodictya multiformis (Stephens, 1915); Latrunculia
(Biannulata) algoaensis Samaai, Janson & Kelly, 2012; Latrunculia (Biannulata) gotzi
Samaai, Janson & Kelly, 2012; Latrunculia (Biannulata) kerwathi Samaai, Janson &
Kelly, 2012; Latrunculia (Biannulata) microacanthoxea Samaai, Gibbons, Kelly &
Davies-Coleman, 2003; Leiosella caliculata Lendenfeld, 1889; Leucandra algoaensis
(Bowerbank, 1864); Leucandra armata (Urban, 1908); Leucandra bathybia (Haeckel,
1869); Leucandra bleeki (Haeckel, 1872); Leucandra hentschelii Brøndsted, 1931;
Leucandra minor (Urban, 1908); Leucandra spissa (Urban, 1909); Leucetta trigona
Haeckel, 1872; Leucilla australiensis (Carter, 1886); Leucilla capsula (Haeckel, 1870);
Leucosolenia eustephana Haeckel, 1872; Lissodendoryx (Ectyodoryx) arenaria Burton,
1936; Lissodendoryx (Lissodendoryx) areolata Lévi, 1963; Lissodendoryx (Lissodendoryx)
digitata (Ridley & Dendy, 1886); Lissodendoryx (Lissodendoryx) simplex (Baer, 1906);
Lissodendoryx (Lissodendoryx) stephensoni Burton, 1936; Lissodendoryx (Lissodendoryx)
ternatensis (Thiele, 1903); Lithochela conica Burton, 1929; Macandrewia auris
Lendenfeld, 1907; Mycale (Aegogropila) meridionalis Lévi, 1963; Mycale (Aegogropila)
simonis (Ridley & Dendy, 1886); Mycale (Mycale) anisochela Lévi, 1963; Mycale
(Mycale) sulcata Hentschel, 1911; Neopetrosia similis (Ridley & Dendy, 1886);
Pachymatisma areolata Bowerbank, 1872; Penares alata (Lendenfeld, 1907); Penares
obtusus (Lendenfeld, 1907); Penares sphaera (Lendenfeld, 1907); Phorbas clathratus
(Lévi, 1963); Phorbas fibrosus (Lévi, 1963); Phyllospongia schulzei Lendenfeld, 1889;
169
Poecillastra tenuirhabda (Lendenfeld, 1907); Polymastia atlantica Samaai & Gibbons,
2005; Proteleia sollasi Dendy & Ridley, 1886; Raspailia rigida Ridley & Dendy, 1886;
Rhabdocalyptus baculifer Schulze, 1904; Spheciospongia capensis (Carter, 1882);
Stelletta agulhana Lendenfeld, 1907; Stelletta capensis Lévi, 1967; Stelletta grubioides
Burton, 1926; Strongylodesma algoaensis Samaai, Gibbons, Kelly & Davies-Coleman,
2003; Strongylodesma tsitsikammaensis Samaai, Gibbons, Kelly & Davies-Coleman,
2003; Stryphnus progressus (Lendenfeld, 1907); Stryphnus unguiculus Sollas, 1886;
Suberites stilensis Burton, 1933; Sycodorus hystrix Haeckel, 1870; Sycon defendens
Borojevic, 1967; Sycon dunstervillia (Haeckel, 1872); Sycon lunulatum (Haeckel, 1872);
Tedania (Tedania) scotiae Stephens, 1915; Tedania (Tedania) stylonychaeta Lévi, 1963;
Tedania (Tedania) tubulifera Lévi, 1963; Tetilla bonaventura Kirkpatrick, 1902; Tetilla
casula (Carter, 1871); Tetilla pedonculata Lévi, 1967; Tetrapocillon novaezealandiae
Brøndsted, 1924; Trachycladus spinispirulifer (Carter, 1879); Tsitsikamma favus Samaai
& Kelly, 2002; Tsitsikamma pedunculata Samaai, Gibbons, Kelly & Davies-Coleman,
2003
193. Natal Ecoregion (101)
Aaptos nuda (Kirkpatrick, 1903); Ancorina corticata Lévi, 1964; Ancorina nanosclera
Lévi, 1967; Aulospongus involutus (Kirkpatrick, 1903); Axinella natalensis (Kirkpatrick,
1903); Axinella weltnerii (Lendenfeld, 1897); Axinyssa tethyoides Kirkpatrick, 1903;
Callyspongia (Toxochalina) dendyi (Burton, 1931); Callyspongia (Toxochalina) ridleyi
(Dendy, 1905); Callyspongia (Toxochalina) robusta (Ridley, 1884); Callyspongia
mammillata (Burton, 1933); Clathria (Clathria) indica Dendy, 1889; Clathria (Clathria)
irregularis (Burton, 1931); Clathria (Clathria) juncea Burton, 1931; Clathria (Clathria)
oculata Burton, 1933; Clathria (Clathria) whiteleggii Dendy, 1922; Clathria (Thalysias)
anomala (Burton, 1933); Clathria (Thalysias) cullingworthi Burton, 1931; Clathria
(Thalysias) delaubenfelsi (Lévi, 1963); Clathria (Thalysias) procera (Ridley, 1884);
Coelosphaera (Coelosphaera) navicelligera (Ridley, 1885); Coscinoderma nardorus
(Lendenfeld, 1886); Crateromorpha (Crateromorpha) lankesteri Kirkpatrick, 1902;
Crella (Grayella) erecta Lévi, 1963; Cyclacanthia cloverlyae Samaai, Govender & Kelly,
2004; Cyclacanthia mzimayiensis Samaai, Govender & Kelly, 2004; Cymbastela
sodwaniensis Samaai, Pillay & Kelly, 2009; Darwinella warreni Topsent, 1905; Dercitus
natalensis (Burton, 1926); Dictyodendrilla caespitosa (Carter, 1886); Discodermia
natalensis Kirkpatrick, 1903; Dragmacidon sanguineum (Burton, 1933); Dysidea
chalinoides (Burton, 1931); Echinodictyum jousseaumi Topsent, 1892; Echinodictyum
marleyi
Burton, 1931; Ecionemia baculifera (Kirkpatrick, 1903); Endectyon
gorgonioides (Kirkpatrick, 1903); Erylus amorphus Burton, 1926; Fangophilina
gilchristi (Kirkpatrick, 1902); Gastrophanella mammilliformis Burton, 1929; Geodia
basilea Lévi, 1964; Geodia dendyi Burton, 1926; Geodia labyrinthica (Kirkpatrick,
1903); Geodia littoralis Stephens, 1915; Geodia megaster Burton, 1926; Geodia
ovifractus Burton, 1926; Geodia ovifractus var. cyathioides Burton, 1926; Grantessa
ramosa (Haeckel, 1872); Guitarra indica Dendy, 1916; Hemiasterella vasiformis
(Kirkpatrick, 1903); Hemiasterella vasiformis var. minor (Kirkpatrick, 1903); Higginsia
natalensis Carter, 1885; Histodermella natalensis (Kirkpatrick, 1903); Hyalonema
(Corynonema) natalense (Lévi, 1964); Hyalonema (Cyliconema) abyssale (Lévi, 1964);
Hyalonema (Cyliconema) curvisclera (Lévi, 1964); Lissodendoryx (Lissodendoryx)
pygmaea (Burton, 1931); Lissodendoryx (Lissodendoryx) ternatensis (Thiele, 1903);
Lithobactrum forte Kirkpatrick, 1903; Lophophysema gilchristi Tabachnick & Lévi,
1999; Microscleroderma hirsutum Kirkpatrick, 1903; Mycale (Carmia) phyllophila
Hentschel, 1911; Mycale (Grapelia) burtoni Hajdu, 1995; Mycale (Mycale) sulcata
Hentschel, 1911; Mycale (Zygomycale) parishii (Bowerbank, 1875); Oceanapia eumitum
(Kirkpatrick, 1903); Pachastrella isorrhopa Kirkpatrick, 1902; Penares orthotriaena
Burton, 1931; Penares sphaera (Lendenfeld, 1907); Petromica (Petromica) digitata
170
(Burton, 1929); Petromica (Petromica) plumosa Kirkpatrick, 1903; Petromica
(Petromica) tubulata (Kirkpatrick, 1903); Phorbas clathratus (Lévi, 1963); Phorbas
clathrodes (Dendy, 1922); Phorbas mollis (Kirkpatrick, 1903); Podospongia natalensis
(Kirkpatrick, 1903); Poecillastra tuberosa (Lévi, 1964); Polymastia disclera Lévi, 1964;
Protosuberites reptans (Kirkpatrick, 1903); Psammoclema inordinatum (Kirkpatrick,
1903); Rhabdastrella actinosa (Lévi, 1964); Rhabdastrella primitiva (Burton, 1926);
Rhabdastrella spinosa (Lévi, 1967); Rhabderemia spirophora (Burton, 1931);
Rhabdocalyptus plumodigitatus Kirkpatrick, 1901; Sigmaxinella arborea Kirkpatrick,
1903; Sigmaxinella incrustans Kirkpatrick, 1903; Spheciospongia excentrica (Burton,
1931); Stelletta agulhana Lendenfeld, 1907; Stelletta agulhana var. paucistella Burton,
1926; Stelletta cyathioides Burton, 1926; Stelletta horrens var. subcylindrica Burton,
1926; Stelletta purpurea Ridley, 1884; Stelletta retroclada (Lévi, 1967); Stelletta rugosa
Burton, 1926; Strongylodesma aliwaliensis Samaai, Keyzers & Davies-Coleman, 2004;
Sycon natalense Borojevic, 1967; Tethya magna Kirkpatrick, 1903; Tetilla casula
(Carter, 1871); Triptolemma incertum (Kirkpatrick, 1903); Waltherarndtia caliculatum
(Kirkpatrick, 1903)
52. Amsterdam-St Paul Province
194. Amsterdam-St Paul Ecoregion (8)
Ancorella paulini Lendenfeld, 1907; Erylus megaster Lendenfeld, 1907; Farrea seiri
Duplessis & Reiswig, 2004; Thenea centrotyla Lendenfeld, 1907; Thenea megaspina
Lendenfeld, 1907; Thenea mesotriaena Lendenfeld, 1907; Thenea microspina
Lendenfeld, 1907; Thenea multiformis Lendenfeld, 1907
Southern Ocean Realm
59. Subantarctic Islands Province
212. Macquarie Island Ecoregion (excluded)
213. Heard and Macdonald Islands Ecoregion (7)
Calyx kerguelensis (Hentschel, 1914); Lissodendoryx (Lissodendoryx) fusca (Ridley &
Dendy, 1886); Poecillastra schulzei (Sollas, 1886); Polymastia invaginata Kirkpatrick,
1907; Polymastia isidis Thiele, 1905; Tetilla coronida Sollas, 1888; Tetilla leptoderma
Sollas, 1886
214. Kerguelen Islands Ecoregion (63)
Artemisina apollinis (Ridley & Dendy, 1886); Biemna chilensis Thiele, 1905; Calyx
kerguelensis (Hentschel, 1914); Chondrocladia (Chondrocladia) fatimae Boury-Esnault
& van Beveren, 1982; Chondrocladia (Chondrocladia) nani Boury-Esnault & van
Beveren, 1982; Cinachyra barbata Sollas, 1886; Craniella coactifera (Lendenfeld, 1907);
Craniella crassispicula (Lendenfeld, 1907); Crella (Pytheas) crassa (Hentschel, 1914);
Dendya clathrata (Carter, 1883); Desmacidon nebulosum Boury-Esnault & van
Beveren, 1982; Ectyonopsis panis (Boury-Esnault & van Beveren, 1982); Grantia
aculeata Urban, 1908; Grantia hirsuta (Topsent, 1907); Grantia tenuis Urban, 1908;
Haliclona (Gellius) constans (Boury-Esnault & van Beveren, 1982); Haliclona (Gellius)
latisigmae (Boury-Esnault & van Beveren, 1982); Haliclona (Reniera) topsenti (Thiele,
1905); Haliclona divulgata Koltun, 1964; Haliclona pedunculata (Ridley & Dendy,
1886); Homaxinella balfourensis (Ridley & Dendy, 1886); Homaxinella flagelliformis
(Ridley & Dendy, 1886); Hymedesmia (Hymedesmia) antarctica Boury-Esnault & van
Beveren, 1982; Hymedesmia (Hymedesmia) mariondufresni Boury-Esnault & van
Beveren, 1982; Hymeniacidon kerguelensis Hentschel, 1914; Iophon proximum var.
reticulare Hentschel, 1914; Isodictya dufresni Boury-Esnault & van Beveren, 1982;
171
Isodictya kerguelenensis (Ridley & Dendy, 1886); Latrunculia (Latrunculia) apicalis
Ridley & Dendy, 1886; Latrunculia (Latrunculia) bocagei Ridley & Dendy, 1886;
Leucandra anfracta (Urban, 1908); Leucandra astricta Tanita, 1942; Leucandra cirrhosa
(Urban, 1908); Leucandra gaussii (Brøndsted, 1931); Leucandra kerguelensis (Urban,
1908); Leucandra minor (Urban, 1908); Leucandra ovata (Poléjaeff, 1883); Leucettusa
vera Poléjaeff, 1883; Leucosolenia australis Brøndsted, 1931; Leucosolenia discoveryi
Jenkin, 1908; Leucosolenia incerta Urban, 1908; Leucosolenia minchini Jenkin, 1908;
Megaciella pilosa (Ridley & Dendy, 1886); Mycale (Oxymycale) acerata Kirkpatrick,
1907; Mycale fibrosa Boury-Esnault & van Beveren, 1982; Myxilla (Ectyomyxilla)
chilensis Thiele, 1905; Myxilla (Ectyomyxilla) kerguelensis (Hentschel, 1914); Phorbas
domini (Boury-Esnault & van Beveren, 1982); Plicatellopsis antarctica (Carter, 1876);
Polymastia invaginata Kirkpatrick, 1907; Polymastia isidis Thiele, 1905;
Pseudosuberites sulcatus (Thiele, 1905); Sigmosceptrella carlinae (Boury-Esnault & van
Beveren, 1982); Spanioplon werthi (Hentschel, 1914); Stelodoryx multidentata (BouryEsnault & van Beveren, 1982); Stylocordyla borealis var. globosa Ridley & Dendy,
1886; Suberites microstomus Ridley & Dendy, 1887; Sycon kerguelense Urban, 1908;
Tedania (Trachytedania) spinata (Ridley, 1881); Tentorium papillatum (Kirkpatrick,
1908); Tetilla leptoderma Sollas, 1886; Xestospongia hispida (Ridley & Dendy, 1886);
Xestospongia variabilis (Ridley, 1884)
215. Crozet Islands Ecoregion (8)
Bathydorus spinosus Schulze, 1886; Esperiopsis profunda Ridley & Dendy, 1886;
Haliclona (Gellius) carduus (Ridley & Dendy, 1886); Hyalonema (Ijimaonema)
clavigerum Schulze, 1886; Iophon cheliferum Ridley & Dendy, 1886; Lissodendoryx
(Ectyodoryx) nobilis (Ridley & Dendy, 1886); Suberites mollis Ridley & Dendy, 1886;
Thenea delicata Sollas, 1886
216. Prince Edward Islands Ecoregion (18)
Amphoriscus elongatus Poléjaeff, 1883; Asbestopluma (Asbestopluma) symmetrica (Ridley
& Dendy, 1886); Aulocalyx irregularis Schulze, 1886; Chondrocladia (Meliiderma)
stipitata (Ridley & Dendy, 1886); Cladorhiza tridentata Ridley & Dendy, 1886;
Esperiopsis profunda Ridley & Dendy, 1886; Fibulia ramosa (Ridley & Dendy, 1886);
Haliclona (Gellius) flagellifera (Ridley & Dendy, 1886); Haliclona (Gellius) glacialis
(Ridley & Dendy, 1886); Iophon abnormale Ridley & Dendy, 1886; Iophon cheliferum
Ridley & Dendy, 1886; Iophon laminale Ridley & Dendy, 1886; Leucandra levis
(Poléjaeff, 1883); Megaciella pilosa (Ridley & Dendy, 1886); Mycale mammiformis
(Ridley & Dendy, 1886); Myxilla (Ectyomyxilla) mariana Ridley & Dendy, 1886;
Raspailia (Raspaxilla) mariana (Ridley & Dendy, 1886); Suberites caminatus Ridley &
Dendy, 1886
217. Bouvet Island Ecoregion (0) (excluded)
No species recorded
218. Peter the First Island Ecoregion (excluded)
61. Continental High Antarctic Province
224. East Antarctic Wilkes Land Ecoregion (174)
Acanthopolymastia acanthoxa (Koltun, 1964); Achramorpha glacialis Jenkin, 1908;
Achramorpha grandinis Jenkin, 1908; Achramorpha nivalis Jenkin, 1908; Amphilectus
rugosus (Thiele, 1905); Anoxycalyx (Anoxycalyx) ijimai Kirkpatrick, 1907; Anoxycalyx
172
(Scolymastra) joubini (Topsent, 1916); Aplysina minima Hentschel, 1914; Artemisina
jovis Dendy, 1924; Artemisina plumosa Hentschel, 1914; Artemisina strongyla Hentschel,
1914; Artemisina tubulosa Koltun, 1964; Asbestopluma (Asbestopluma) belgicae
(Topsent, 1901); Asbestopluma (Asbestopluma) callithrix Hentschel, 1914; Asbestopluma
(Asbestopluma) calyx Hentschel, 1914; Asbestopluma (Asbestopluma) obae Koltun, 1964;
Biemna macrorhaphis Hentschel, 1914; Breitfussia chartacea (Jenkin, 1908); Breitfussia
vitiosa (Brøndsted, 1931); Calyx arcuarius (Topsent, 1913); Caulophacus (Caulophacus)
antarcticus Schulze & Kirkpatrick, 1910; Caulophacus (Caulophacus) oviformis
(Schulze, 1886); Chondrocladia (Chondrocladia) antarctica Hentschel, 1914;
Chonelasma choanoides Schulze & Kirkpatrick, 1910; Cinachyra antarctica (Carter,
1872); Cinachyra barbata Sollas, 1886; Cladocroce gaussiana (Hentschel, 1914);
Cladorhiza moruliformis Ridley & Dendy, 1886; Cladothenea andriashevi Koltun, 1964;
Clathria (Axosuberites) flabellata (Topsent, 1916); Clathria (Axosuberites) nidificata
(Kirkpatrick, 1907); Clathria (Axosuberites) ramea (Koltun, 1964); Clathria (Clathria)
lipochela Burton, 1932; Clathria (Clathria) paucispicula (Burton, 1932); Clathria
(Clathria) pauper Brøndsted, 1927; Clathria (Clathria) toxipraedita Topsent, 1913;
Clathria (Thalysias) koltuni Hooper in Hooper & Wiedenmayer, 1994; Coelosphaera
(Coelosphaera) antarctica Koltun, 1976; Craniella sagitta (Lendenfeld, 1907); Crella
(Crella) tubifex (Hentschel, 1914); Crella (Pytheas) crassa (Hentschel, 1914); Crella
(Pytheas) stylifera Hentschel, 1914; Dermatreton hodgsoni Jenkin, 1908; Dolichacantha
macrodon Hentschel, 1914; Eurypon miniaceum Thiele, 1905; Fibulia maeandrina
(Kirkpatrick, 1907); Grantia hirsuta (Topsent, 1907); Grantia transgrediens Brøndsted,
1931; Guancha apicalis Brøndsted, 1931; Guitarra antarctica Hentschel, 1914; Guitarra
dendyi (Kirkpatrick, 1907); Halichondria (Halichondria) prostrata Thiele, 1905;
Haliclona (Gellius) glacialis (Ridley & Dendy, 1886); Haliclona (Gellius) rudis (Topsent,
1901); Haliclona (Gellius) tylotoxa (Hentschel, 1914); Haliclona (Reniera) topsenti
(Thiele, 1905); Haliclona (Rhizoniera) dancoi (Topsent, 1913); Haliclona virens
(Topsent, 1908); Hemigellius bidens (Topsent, 1901); Hemigellius calyx (Ridley &
Dendy, 1886); Hemigellius pachyderma Burton, 1932; Holascus tenuis Schulze, 1904;
Homaxinella balfourensis (Ridley & Dendy, 1886); Homoieurete macquariense Reiswig
& Kelly, 2011; Hyalonema (Cyliconema) drygalskii Schulze & Kirkpatrick, 1910;
Hymedesmia (Hymedesmia) antarctica Boury-Esnault & van Beveren, 1982;
Hymedesmia (Hymedesmia) gaussiana Hentschel, 1914; Hymedesmia (Hymedesmia)
leptochela Hentschel, 1914; Hymedesmia (Stylopus) antarctica Hentschel, 1914;
Hymedesmia (Stylopus) dermata var. antarctica Hentschel, 1914; Hymenancora
rhaphidophora Hentschel, 1914; Hymeniacidon centrotyla Hentschel, 1914; Inflatella
belli (Kirkpatrick, 1907); Inflatella coelosphaeroides Koltun, 1964; Iophon aceratum
Hentschel, 1914; Iophon gaussi Hentschel, 1914; Iophon unicorne Topsent, 1907;
Iotroata somovi (Koltun, 1964); Isodictya cavicornuta Dendy, 1924; Isodictya delicata
(Thiele, 1905); Isodictya delicata var. megachela Burton, 1934; Isodictya doryphora
(Brøndsted, 1927); Isodictya erinacea (Topsent, 1916); Isodictya kerguelenensis (Ridley
& Dendy, 1886); Isodictya lankesteri (Kirkpatrick, 1907); Isodictya obliquidens
(Hentschel, 1914); Isodictya setifera (Topsent, 1901); Jenkina articulata Brøndsted,
1931; Jenkina glabra Brøndsted, 1931; Jenkina hiberna (Jenkin, 1908); Kirkpatrickia
variolosa (Kirkpatrick, 1907); Latrunculia (Latrunculia) apicalis Ridley & Dendy, 1886;
Latrunculia (Latrunculia) basalis Kirkpatrick, 1908; Leucandra comata Brøndsted,
1931; Leucandra gausapata Brøndsted, 1931; Leucandra mawsoni Dendy, 1918;
Leucascus leptoraphis (Jenkin, 1908); Leucetta antarctica Dendy, 1918; Leucosolenia
aboralis Brøndsted, 1931; Leucosolenia australis Brøndsted, 1931; Leucosolenia solida
Brøndsted, 1931; Lissodendoryx (Ectyodoryx) anacantha (Hentschel, 1914);
Lissodendoryx (Ectyodoryx) antarctica (Hentschel, 1914); Lissodendoryx (Ectyodoryx)
plumosa (Hentschel, 1914); Lissodendoryx (Ectyodoryx) ramilobosa (Topsent, 1916);
Lissodendoryx (Lissodendoryx) flabellata Burton, 1929; Lissodendoryx (Lissodendoryx)
styloderma Hentschel, 1914; Megapogon crispatus Jenkin, 1908; Megapogon raripilus
Jenkin, 1908; Microxina benedeni (Topsent, 1901); Microxina phakellioides
173
(Kirkpatrick, 1907); Mycale (Aegogropila) magellanica (Ridley, 1881); Mycale (Carmia)
gaussiana Hentschel, 1914; Mycale (Mycale) tridens Hentschel, 1914; Mycale
(Oxymycale) acerata Kirkpatrick, 1907; Mycale profunda Koltun, 1964; Myxilla
(Burtonanchora) asigmata (Topsent, 1901); Myxilla (Burtonanchora) lissostyla Burton,
1938; Myxilla (Myxilla) insolens Koltun, 1964; Myxilla (Myxilla) mollis Ridley & Dendy,
1886; Myxodoryx hanitschi (Kirkpatrick, 1907); Phelloderma radiatum Ridley & Dendy,
1886; Phorbas acanthochela (Koltun, 1964); Phorbas glaberrimus (Topsent, 1917);
Phorbas nexus (Koltun, 1964); Plakina monolopha var. antarctica Lendenfeld, 1907;
Plakina trilopha var. antarctica Lendenfeld, 1907; Plicatellopsis antarctica (Carter,
1876); Plicatellopsis fragilis Koltun, 1964; Plocamionida gaussiana (Hentschel, 1914);
Poecillastra compressa antarctica Koltun, 1964; Poecillastra schulzei (Sollas, 1886);
Polymastia invaginata Kirkpatrick, 1907; Polymastia invaginata var. gaussi Hentschel,
1914; Polymastia isidis Thiele, 1905; Proteleia burtoni Koltun, 1964; Pseudosuberites
hyalinus (Ridley & Dendy, 1886); Pseudosuberites nudus Koltun, 1964; Pyloderma
latrunculioides (Ridley & Dendy, 1886); Raspailia (Hymeraphiopsis) irregularis
Hentschel, 1914; Rhizaxinella australiensis Hentschel, 1909; Rossella antarctica Carter,
1872; Rossella fibulata Schulze & Kirkpatrick, 1910; Rossella gaussi Schulze &
Kirkpatrick, 1910; Rossella lychnophora Schulze & Kirkpatrick, 1910; Rossella mixta
Schulze & Kirkpatrick, 1910; Rossella racovitzae Topsent, 1901; Rossella vanhoeffeni
(Schulze & Kirkpatrick, 1910); Rossella vanhoeffeni var. armata (Schulze &
Kirkpatrick, 1910); Rossella vanhoeffeni var. vanhoeffeni (Schulze & Kirkpatrick,
1910); Rossella villosa Burton, 1929; Rossella vitiosa Wiedenmayer in Hooper &
Wiedenmayer, 1994; Soleneiscus apicalis (Brøndsted, 1931); Soleneiscus hispidus
(Brøndsted, 1931); Sphaerotylus antarcticus Kirkpatrick, 1907; Sphaerotylus antarcticus
var. drygalskii Hentschel, 1914; Sphaerotylus vanhoeffeni Hentschel, 1914; Stelletta
crater Dendy, 1924; Stylocordyla borealis var. irregularis Hentschel, 1914; Stylocordyla
chupachups Uriz, Gili, Orejas & Perez-Porro, 2011; Suberites caminatus Ridley &
Dendy, 1886; Suberites microstomus Ridley & Dendy, 1887; Suberites topsenti (Burton,
1929); Sycantha longstaffi (Jenkin, 1908); Sycetta antarctica (Brøndsted, 1931); Sycon
australe (Jenkin, 1908); Tedania (Tedania) trirhaphis Koltun, 1964; Tedania
(Tedaniopsis) charcoti Topsent, 1907; Tedania (Tedaniopsis) gracilis Hentschel, 1914;
Tedania (Tedaniopsis) massa Ridley & Dendy, 1886; Tedania (Tedaniopsis) oxeata
Topsent, 1916; Tedania (Tedaniopsis) vanhoeffeni Hentschel, 1914; Tethyopsis
longispinus (Lendenfeld, 1907); Tetilla leptoderma Sollas, 1886
225. East Antarctic Enderby Land Ecoregion (8)
Anoxycalyx (Anoxycalyx) ijimai Kirkpatrick, 1907; Anoxycalyx (Scolymastra) joubini
(Topsent, 1916); Caulophacus (Caulodiscus) valdiviae Schulze, 1904; Coelosphaera
(Coelosphaera) antarctica Koltun, 1976; Isodictya erinacea (Topsent, 1916); Rossella
antarctica Carter, 1872; Rossella racovitzae Topsent, 1901; Rossella villosa Burton,
1929
226. East Antarctic Dronning Maud Land Ecoregion (7)
Anoxycalyx (Scolymastra) joubini (Topsent, 1916); Cladorhiza mani Koltun, 1964;
Clathria (Thalysias) ongulensis (Hoshino, 1977); Isodictya echinata Thomas & Matthew,
1986; Rossella antarctica Carter, 1872; Rossella racovitzae Topsent, 1901; Rossella
villosa Burton, 1929
227. Weddell Sea Ecoregion (71)
Acanthopolymastia acanthoxa (Koltun, 1964); Anoxycalyx (Scolymastra) joubini
(Topsent, 1916); Aplysina minima Hentschel, 1914; Ascaltis abyssus Rapp, Janussen &
174
Tendal, 2011; Astrotylus astrotylus Plotkin & Janussen, 2007; Axinella antarctica
(Koltun, 1964); Calyx arcuarius (Topsent, 1913); Caulophacus (Caulodiscus) brandtae
Janussen, Tabachnick & Tendal, 2004; Caulophacus (Caulophacus) discohexactinus
Janussen, Tabachnick & Tendal, 2004; Caulophacus (Caulophacus) instabilis Topsent,
1910; Caulophacus (Caulophacus) scotiae Topsent, 1910; Caulophacus (Oxydiscus)
weddelli Janussen, Tabachnick & Tendal, 2004; Chondrocladia (Chondrocladia)
antarctica Hentschel, 1914; Cinachyra antarctica (Carter, 1872); Cladocroce gaussiana
(Hentschel, 1914); Cladorhiza penniformis Göcke & Janussen, 2013; Clathria
(Axosuberites) nidificata (Kirkpatrick, 1907); Clathria (Clathria) pauper Brøndsted,
1927; Clathrina broendstedi Rapp, Janussen & Tendal, 2011; Cornulum antarcticum
Göcke & Janussen, 2013; Crella (Pytheas) crassa (Hentschel, 1914); Desmacella koltuni
Göcke & Janussen, 2013; Esperiopsis scotiae Topsent, 1915; Halichondria
(Halichondria) prostrata Thiele, 1905; Haliclona (Gellius) flagellifera (Ridley & Dendy,
1886); Haliclona (Gellius) rudis (Topsent, 1901); Haliclona (Gellius) tylotoxa (Hentschel,
1914); Haliclona (Rhizoniera) dancoi (Topsent, 1913); Hemigellius bidens (Topsent,
1901); Hemigellius fimbriatus (Kirkpatrick, 1907); Holascus obesus Schulze, 1904;
Holascus pseudostellatus Janussen, Tabachnick & Tendal, 2004; Inflatella belli
(Kirkpatrick, 1907); Iophon unicorne Topsent, 1907; Isodictya doryphora (Brøndsted,
1927); Isodictya kerguelenensis (Ridley & Dendy, 1886); Isodictya setifera (Topsent,
1901); Isodictya toxophila Burton, 1932; Leucetta weddelliana Rapp, Janussen &
Tendal, 2011; Lissodendoryx (Ectyodoryx) ramilobosa (Topsent, 1916); Lissodendoryx
(Lissodendoryx) styloderma Hentschel, 1914; Lophocalyx profunda Janussen & Reiswig,
2009; Lophocalyx topsenti Janussen & Reiswig, 2009; Microxina benedeni (Topsent,
1901); Mycale (Oxymycale) acerata Kirkpatrick, 1907; Myxilla (Burtonanchora)
asigmata (Topsent, 1901); Myxilla (Myxilla) insolens Koltun, 1964; Myxodoryx hanitschi
(Kirkpatrick, 1907); Periphragella antarctica Janussen, Tabachnick & Tendal, 2004;
Phelloderma oxychaetoides Göcke, Hajdu & Janussen, 2014; Phorbas megasigma Rios
& Cristobo, 2007; Polymastia invaginata Kirkpatrick, 1907; Polymastia zitteli
(Lendenfeld, 1888); Pseudosuberites hyalinus (Ridley & Dendy, 1886); Pseudosuberites
nudus Koltun, 1964; Pyloderma latrunculioides (Ridley & Dendy, 1886); Raspailia
(Hymeraphiopsis) irregularis Hentschel, 1914; Rossella antarctica Carter, 1872; Rossella
fibulata Schulze & Kirkpatrick, 1910; Rossella levis (Kirkpatrick, 1907); Rossella nuda
Topsent, 1901; Rossella racovitzae Topsent, 1901; Rossella vanhoeffeni (Schulze &
Kirkpatrick, 1910); Rossella villosa Burton, 1929; Stylocordyla chupachups Uriz, Gili,
Orejas & Perez-Porro, 2011; Suberites topsenti (Burton, 1929); Tedania (Tedania)
trirhaphis Koltun, 1964; Tedania (Tedaniopsis) charcoti Topsent, 1907; Tedania
(Tedaniopsis) oxeata Topsent, 1916; Tedania (Tedaniopsis) tantula (Kirkpatrick, 1907);
Tetilla leptoderma Sollas, 1886
228. Amundsen/Bellingshausen Sea Ecoregion (excluded)
229. Ross Sea Ecoregion (excluded)
Other
Vema Seamount (13)
Desmacidon clavatum Lévi, 1969; Echinodictyum macroxiphera Lévi, 1969; Eurypon
fulvum Lévi, 1969; Gelliodes coscinopora Lévi, 1969; Haliclona (Reniera) alusiana (Lévi,
1969); Mycale (Paresperella) curvisigma Lévi, 1969; Mycale diastrophochela Lévi, 1969;
Oceanapia atlantica Lévi, 1969; Smenospongia nuda (Lévi, 1969); Spongia (Spongia)
brunnea Lévi, 1969; Spongia (Spongia) violacea Lévi, 1969; Strongylodesma areolata
Lévi, 1969; Suberea pedunculata (Lévi, 1969)
175