Polar Biol (1992)12:559-585
© Springer-Verlag1992
Biogeographic traits and checklist of Antarctic demosponges
M. Sarfi, A. Balduzzi, M. Barbieri, G. Bavestrello and B. Burlando
Istituto di Zoologia dell'UniversitY, Via Balbi 5, 1-16126 Genova, Italy
Received 26 March 1991; accepted 30 March 1992
Summary. The biogeography of Antarctic demosponges
has been studied by dividing Antarctic and circumantarctic areas into geographic entities, and then assigning to
these entities all recorded species according to literature
reports. Correspondence analysis ordination based on the
presence or absence of species shows the existence of a
distinct Antarctic Faunistic Complex (AFC), including
continental Antarctica, most of the Antarctic and circumantarctic islands and the Magellan area. Such a result has
enabled us to drawup a checklist of 352 Antarctic demosponge species. Investigation of within-AFC patterns
indicates that the continent is a highly homogeneous area,
establishing closer relationships with the Scotia Arc and to
a lesser extent with the Magellan region. 'The AFC has low
specific affinities with the other circumantarctic regions
(South Africa, temperate Australia and New Zealand),
whereas at the generic level relationships appear more
pronounced. This biogeographic pattern may lead us to
suppose a common Gondwanian origin :for Antarctic and
circumantarctic sponge faunas, followed by differentiation
due to Gondwana fragmentation. Antarctica moved towards polar latitudes and became progressively isolated,
only maintaining active interchanges with South America.
Climatic changes possibly induced intense processes of
speciation in the Antarctic demosponge fauna, thus contributing to its differentiation.
In biogeographic studies, sponges are an important
element for characterizing faunal areas and tracing their
histories, since they have sessile adults and larval stages
with a short dispersal range. A first attempt to outline a
biogeographic pattern for the Antarctic sponge fauna was
made by Burton (1932), who found no marked differences
between Graham Land (Antarctic Peninsula) and Ross
Sea faunas, and a high degree of faunistic affinity between
Antarctica and some of the surrounding lands (Scotia Arc,
Falkland, Kerguelen and southern South America).
Koltun (1970) carried out a more comprehensive investigation of the Antarctica sponge fauna by considering
data on about 300 sponge species, which allowed him to
define some basic biogeographic traits: (a) a high level of
specific endemism coupled with a negligible generic endemism rate; (b) circumpolar distribution of many Antarctic
sponges, with relatively few species showing a restricted
distribution; (c) closer faunistic relationship with South
America and the Falkland Islands, than with Australia
and New Zealand.
After this work, several other monographs on the
sponge fauna of Antarctic and circumantarctic areas were
published, thus demanding a reexamination of faunistic
affinities among these areas. We therefore considered the
updated record of Antarctic and circumantarctic demosponge faunas in order to highlight a distinctive Antarctic
faunistic area and to reconstruct its affinities and possible
origins.
Introduction
Sponges are an important element of the Antarctic biota,
as suggested by their species diversity and by the finding of
sponge-dominated communities in several Antarctic areas
(Barthel et al. 1990; Dayton et al. 1970; Di Geronimo et al.,
in press). In Antarctic benthic environments, at about
100m depth, sponges can reach biomass values of
2,400 g/mq, which are comparable with the largest scores
of tropical areas (Beliaev and Ushakov 1957).
Correspondence to: A. Balduzzi
Material and methods
This study is based on the subdivision of a southernmost portion of
the Southern hemisphere into geographic entities representing
antarctic, circumantarctic and neighbouring areas (Fig. 1, Table 1).
These entities do not generally represent biogeographic areas as
recognized by other authors, but are simply a set of geographic
ranges to be compared, whilst keeping to a minimum a priori
assumptions about affinity patterns.
Sea areas pertaining to the Antarctic continent have been divided
into nine sector entities (Fig. 1) each of about 40° longitudinal range.
560
terms of endemism rates and shared species and genera in pairwise
comparisons. Multivariate statistical analysis has been carried out
on a desktop computer using the SAS package (SAS Institute Inc.,
Cary, NC).
f:A
SG
ss
\
GO
BO
VS
SA
E~
HE
KE
5/
Fig. 1. Map of the considered geographic entities. Abbreviations as
in Table 1
Sectors have been numbered eastwards starting from the one including the Greenwich meridian, and have been arranged in such a way
as to make the two sides of the Antarctic Peninsula fall in different
entities. This is the same pattern utilized by Rogick (1965) for
bryozoans. Most circumantarctic islands and archipelagos have
been treated as distinct entities, whereas some Antarctic coastal
islands (such as Balleny Islands, South Shetland, etc.) have been
included in coastal sectors.
Besides Antarctic and circumantarctic regions, the following
extra-Antarctic entities have been considered: South Africa (including the coasts of Namibia); Tasmania (including Australian coasts
pertaining to the Bass Strait); New Zealand; Vema Seamount (a
submerged plateau in the South-eastern Atlantic); Magellan region
(coasts of South America and related islands, excluding Falkland,
south of 39~ on the Pacific side and south of the Rio de la Plata on
the Atlantic side). Except for Vema Seamount, all of these latter
entities roughly correspond to biogeographic provinces.
The distribution of demosponges among the above geographic
entities has been reconstructed using records of species from the
following papers: Barthel et al. (1990); Bergquist (1968, 1970);
Bergquist and Warne (1980); Bergquist and Fromont (1988); BouryEsnault and Van Beveren (1982); Broendsted (1926); Burton (1929,
1930, 1932, 1934, 1938); Carter (1872, 1875, 1877, 1879); Dendy
(1924); Desqueyroux (1975); Desqueyroux-Faundez (1989); Desqueyroux and Moyano (1987); Hentschel (1914); Kirkpatrick
(1907a, b, 1908); Koltun (1964, 1976); Lendenfeld (1907a, b); L6vi
(1956, 1963, 1964, 1967, 1969); Ridley (1881); Ridley and Dendy
(1886, 1887); Sar/t (1978); Sarg et al. (1990); Sollas (1886, 1888);Tanita
(1959); Thiele (1905); Topsent (1901, 1902, 1907, 1908, 1913, 1915,
1916, 1917); Uriz (1984, 1987, 1988); Vacelet and Arnaud (1972);
Wiedenmayer (1989). The list of species obtained has been revised by
updating the nomenclature and eliminating synonymies. Koltun
(1964, 1976) and Desqueyroux and Moyano (1987) have been utilized
as a base for the Antarctic and Magellan fauna, respectively, but
original references have been used to solve taxonomic or distributional problems.
Data about presence-absence of species within entities have been
summarized by the ordination technique of correspondence analysis
(Benz6cri et al. 1982), also testing the significance of axes by Lebart's
(1975) tables. Affinities among entities have also been evaluated in
Results
The adopted geographic subdivision results in a total of 27
entities (Table 1), whilst the demosponge record from these
entities amounts to 817 species and 224 genera. Records
are unequally distributed among entities, not only due to
the variable extension of these geographic areas, but also
because sampling efforts have not been uniformly distributed throughout the Antarctic Ocean. For example, among
the extensive continental sectors, number 7 is completely
devoid of records and is thus excluded from any analysis,
whilst numbers 1 and 5 show much fewer species than
some circumantarctic islands.
The correspondence analysis made on all species and
entities exhibits a pattern consisting of a single cluster and
some outliers (Fig. 2A). The cluster includes the Antarctic
sectors, the Magellan area and all eircumantarctic islands
but Campbell. New Zealand and Tasmanian entities are
separated from the cluster along the first axis, showing a
certain degree of reciprocal affinity. Conversely, South
Africa and Vema Seamount are more distantly and independently separated along the second axis, thus appearing
faunistically distinct from the other regions. Campbell
Island separates from the cluster in the direction of New
Zealand, which is its closest neighbour. Among the eight
species recorded from Campbell, seven are in common
with New Zealand whereas only three are shared with
Antarctic entities.
The group of clustered entities enables us to delimit a
distinct Antarctic demosponge fauna, referred to henceforth as the Antarctic Faunistic Complex (AFC), and also
allows us to compile an updated checklist of Antarctic
species (Table 2). A provisional version of this checklist
has been published in the Reports of the Italian National
Program for Antarctic Research (Balduzzi et al. 1991).
Intra-AFC structuring has been explored through
correspondence analysis of AFC entities and related species only. Contrary to the former comprehensive correspondence analysis, overall differences are more shaded
and biases arise from considering all entities, due to
comparing highly unequal numbers of species. Hence, a
selection has been made excluding those entities showing
less than 60 species, based on the existence of a gap
between about 30 and 60 in the numbers of recorded
species per entity (Table 1).
After elimination of the less informative entities, correspondence analysis on Antarctic and neighbouring entities has been repeated, obtaining virtually the same pattern as in the comprehensive analysis (Fig. 2B), that is an
AFC cluster plus outliers. This indicates that entity elimination, whilst ridding the analysis of noisy data, leads to
no appreciable loss of information.
In the correspondence analysis ordination of AFC
entities (Fig. 2C), continental sectors are clustered, whereas South Georgia, Falkland and Magellan progressively
561
Table 1, Geographic entities considered
in this ~;tudy, with abbreviations and
numbers of recorded species and genera
of Demospongiae. Entities falling in the
Antarctic Faunistic Complex, as defined
later in the text, are marked with an asterisk ('~)
Entity
Abbrev.
Species
Continental Antarctica
* sector 1
* sector 2
* sector 3
* sector 4
* sector 5
* sector 6
* sector 7
* sector 8
* sector 9
CONT
C1
C2
C3
C4
C5
C6
C7
C8
C9
205
4
63
117
74
125
5
98
4
46
69
49
69
5
80
106
49
59
Circumantarctic islands
* Tristan da Cunha
* Gough
* Bouvet
* Prince Edward-Marion
* Crozet
* Kerguelen
* Heard
* Macquarie
Campbell
* Falkland
* South Orkney
* South Georgia
* South Sandwich
TC
GO
BO
EM
CR
KE
HE
MQ
CA
FA
SO
SG
SS
6
4
1
19
7
95
21
17
8
77
8
73
5
5
4
1
14
7
59
17
14
7
50
7
50
4
Neighbouring areas
Vema Seamount
South Africa (incl. Namibia)
Tasmania and South Australia
New Zealand
* Magellan (excl. Falkland)
VS
SA
TA
NZ
MG
26
194
120
271
150
24
88
67
126
65
* Antarctic Faunistic Complex
AFC
352
127
separale from the cluster along the first axis, and Kerguelen lies far away along the second axis. This pattern
recalls geographic topology, though the clustering of
continental entities makes ordination spacing not proportional to geographic distance, thus emphasizing the
homog:.,neity of the Antarctic continental sponge
fauna.
Relationships within the continent have been further
investigated by correspondence analysis on continental
sector entities and related fauna. Results indicate greater
affinitie~ among sector numbers 2, 3, 4 and 9, whereas
numbers 5 and 8 separate along either of the first two axes
(Fig. 2D). The two latter entities are those which maximally s h o ~ the influence of their neighbouring circumantarctic or e~tra-Antarctic areas (see Fig. 1). This clearly appears in the ordination of AFC and neighbourhoods
(Fig. 2B), where within the AFC cluster C5 is drawn
towardi New Zealand, and in the analysis devoted to AFC
entities (Fig. 2C), where C8 is slightly drawn towards
South Georgia.
Bas~z,d on the distribution of AFC species in the entities
and in other extra-AFC areas, faunistic patterns have been
constructed for each AFC entity, for AFC as a whole, and
for cont nental Antarctica (i.e. all continental sectors taken
as a wh:,le). Entities with less than 60 species have been set
apart. The patterns consist of the percentage rates of local
endemi~ms, of AFC endemisms where meaningful, of
Genera
continental endemisms where meaningful, and of the
percentage of species shared in pairwise comparisons
among AFC entities, between AFC and extra-AFC entities, and between AFC entities and major world oceanic
areas (Figs. 3, 4).
Faunistic patterns have been constructed in the same
way for genera, though in this case a particular use of the
term 'endemism' has been made. The recognition of true
endemic genera would have required deriving the world
distribution of all AFC genera, which is beyond the aims of
this paper. Hence, the term 'endemism' refers here to those
genera which have been found in a particular entity only
among the considered ones, thus disregarding their possible occurrence elsewhere. For the same reason, at the
generic level, no pairwise affinity between entities and
world oceanic areas has been estimated.
The homogeneity of the AFC is reflected in the fact
that in each of its entities local endemisms are fewer than
Antarctic endemisms (Fig. 3), whilst in continental sectors
continental endemisms are intermediate between the former two endemism rates (Fig. 4). Species affinities in
pairwise comparisons are always higher among AFC
entities than between AFC and extra-AFC entities (Figs. 3,
4), while the continental sectors in particular show highest
species affinities with each other, intermediate affinities
with other AFC entities and lowest affinities with extraAFC entities (Fig. 4).
562
Tlc5
oNZ
CA MG GO
NZ
KE
~
~
A
.vs
B
c8
~
+
FA
.,.....c8 ~ c 9
MO
.C9
.c3
.c5
.c2
C4
C
D
Fig. 2A-D. Correspondence analysis ordinations, based on the
presence or absence of species within entities. (A) All considered
entities; (B) all entities with more than 60 species; (C) Antarctic
Faunistic Complex entities with more than 60 species;(D) Antarctic
continental sectors with more than 60 species. In all graphs x = first
axis, y=second axis; in (A) positive coordinate values have been
taken to the left and below the origin to highlight ordination
similarity with (B). Variance explained by the axes: (A) I: 10.1%, II:
9.59%; (B) I: 19.6%, II: 18%; (C) I: 26.5%, II: 17.8%; (D) I: 25.8%, II:
25.2%. All plotted axes are significant (P<0.05) according to
Lebart's (1975) tables
Conversely, at the generic level no affinity drop occurs
in pairwise comparisons between AFC and extra-AFC
entities (Figs. 3, 4). This has also been verified by statistically comparing (Kruskal-Wallis test), for each AFC entity, the numbers of taxa (species or genera) shared with
each other AFC entity and those shared with each extraAFC entity, always finding significant differences for species (P < 0.05) and no difference for genera.
Among the world ocean areas, the AFC shows the
highest affinities with West Atlantic and East Pacific
faunas (Fig. 3). This is also the case for singular continental
sectors and circumantarctic entities (Figs. 3, 4).
Discussion
A major question has been first addressed in this paper,
namely the possibility of a posteriori delimiting an Antarctic sponge fauna. A result has been achieved through
correspondence analysis ordination, which separates a
group of Antarctic and circumantarctic areas from neighbouring regions, thus showing the existence of a distinct
Antarctic Faunistic Complex (AFC). The use of presenceabsence data in this analysis could introduce some distortion, since regional variations in species density are overlooked. However, estimating local densities of Antarctic
563
Continental Antarctica
South Georgia
I.e. ~ ,
a.e. ~
I.e.
a.e.
MG
FA
KE
CONT
MG
FA
KE
SA
TA
NZ
SA
TA
NZ
.-
EP
CONT
SG
FA
KE
SA
TA
NZ
wP
I 205 species
98 genera
EA "*
WAy"
EA
0
0
40
6'0
8'0
EP
WP
73 species
IN
L
9
50 genera
mm
I
I
AR
100
C--
I,e.
a,e.
EP m
WP
IN i
AR
Magellan
0
Falkland Islands
20
40
60
80
100
"
150 species
IN
EA
WA
AR L
0
65 genera
20
I.e.
a.e.
CONT
SG
MG
KE
CONT
8G
MG
FA
I
40
60
80
100
Antarctic
Faunistic Complex
Kerguelen Islands
I.e.
a.e.
(excl. Falkland)
$ATA ~
~
b
SA
TA
NZ
Eml
~'9
IN
EA
WA l i i l l
AR
50 genera
/I
: i n
:
0
20
40
60
80
95 species
WP
77 species
100
0
i
WP
II
352 specks
IN
EP I
EP
WP
IN
EA
WA
AR
EP
59 genera
EA
II
WA
i
127 genera
AR
20
20
40
60
80
100
[
40
60
mmspecies N N g e n e r a
80
100
I
Fig. 3. Percentage affinity patterns at specific and generic levels of
the demosponge faunas pertaining to continental Antarctica (all
sectors considered as a whole), to each non-continental AFC entity
with morv. than 60 species, and to the Antarctic Faunistic Complex.
The first 1we bar pairs (first one in the AFC graph) refer to endemism
rates (see: text for the meaning of 'endemic' genera): i.e.=local
endemisms, a.e. = AFC endemisms. The other bars refer to percentage of taxa shared with other entities (abbreviations as in Table 1),
and with major oceanic subdivisions: EP = East Pacific, WP = West
Pacific, IN=Indian, EA=East Atlantic and Mediterranean, WA
= West Atlantic, AR = Arctic
sponge populations is impossible since samplings are
scattered and in most cases no information about biomass
is given. However, the obtained ordination patterns show
a good ~verall agreement with previous biogeographic
analyses of the Antarctic ocean (Hedgpeth 1970; Koltun
1970; White 1984), and should therefore be considered as
only mb:.imally biased.
Demarcation of the AFC emphasizes the individuality
and honlogeneity of the demosponge fauna throughout a
wide w(}rld area including Antarctic and subantarctic
waters. '!?his is confirmed by the clearcut difference between the high endemism rate of the AFC as a whole,
compare:l to the low rates of within-AFC local endemisms.
The most important northward outspreading of the
AFC is the inclusion of the Magellan area, indicating a
much stronger affinity of Antarctica with southern South
America than with other continents. Outside this very
particular area, the incorporation of most circumantarctic
islands in the AFC makes the northern boundary roughly
coincide with the Antarctic Convergence, though Tristan
da Cunha and Gough are situated beyond this to the
north. Information about the sponge fauna along the
Convergence is very scarce since all entities but Kerguelen
include less than 25 species. However, since the Kerguelen
entity remains within the AFC cluster even after elimination of entities with a few species, it is reasonable to think
that outside southern South America the AFC at least
564
I.e.
c.e.
a.e.
C2
C4
C5
C8
C9
SG
I
KE
SA
TA
NZ
63 species
46 genera
1
0
20
40
60
EP
WP
IN
EA
WA
AN
]
j
80
I,e, l
c.e. ~
!
"t
L
SG
LE
KE
SA
TA
NZ
i
L
i
EP
WP
IN
EA
WA
AR
117 species
69 genera
i
i
i
i
20
40
60
80
I.e.
c.e.
a.e,
c2
c3
c4
c8
c9
C2
C3
C4
C5
C9
SG
MG
FA
KE
SG
$A
TA
NZ
i
"9
1
20
40
49 genera
i
~
i
i
20
4O
6O
80
60
80
100
100
C9
I.e. , ~
c.e.
a.e.
L.
L
c2
C3
C4
C5
r
L,
SG
MG
FA
KE
,,,,
L
L
L
i
sA I,,
TA ~...................................
NZ
i
EP
WP
IN L,
EA ~=
WA
125 species
69 genera
i
9
0
100
KE
i
74 species
._
C8
I.e,
c.e.
a.e.
EP
WP
IN
EA
WA
AR
C2 l,,,,,,,,~,
C3 [
C5 L
C9
0
100
i
a.e.
o
C5
SA
TA
NZ
C4
C3
C2
0
wP
80 species
40
60
80
106 species 1
IN
EA
WA
49 genera
20
~
100
f
59 genera
II
J
0
20
40
60
80
100
Fig. 4. Percentage affinitypatterns at specificand generic levelsof the demospongefaunas pertaining to the Antarctic continental sectors with
more than 60 species. Symbols and bars as in Fig. 3, with the followingaddition: c.e.= continental endemisms
reaches the Antarctic Convergence. In this respect, the
AFC results are largely coincident with Hedgpeth's (1970)
Antarctic Region, the main differences being such entities
as Magellan and Kerguelen which belong to a Subantarctic Region in Hedgpeth's scheme.
The analysis of the relationships among AFC entities,
aimed at delineating biogeographic patterns on a finer
scale, has been necessarily restricted to a subset of more
informative entities, excluded areas being considered as
too scarcely explored for this purpose. This analysis clearly
shows that the continent is homogeneous with respect to
the rest of the AFC, and hence does not seem to confirm
the distinction of two biogeographic subunits, West and
East, recognized in the Antarctic continent by various
authors (Ekman 1935; Knox 1970; Andriashev 1965; Kussakin 1967). Also, the differences in species numbers
between the Western and Eastern continent, noticed for
sponges by Koltun (1970), now appear to be contradicted
by the large number of species recorded by Barthel et al.
(1990) from the Weddell sea.
Apart from the continent, no other grouping of AFC
entities is supported by our data, although future research
might better define the position of poorly known entities,
or even modify current opinions about the most investigated ones. However, a particular mention is due to the
geographically-consistent ordination pattern of Magellan,
Falklands and South Georgia, pointing to the continental
sector C8 which includes part of the Antarctic Peninsula.
A first-rank topic of Antarctic biogeographic studies seems
confirmed here, namely the existence of faunal exchanges
between Antarctica and South America via the Scotia Arc,
also proposed by Knox and Lowry (1977) as one of the
mechanisms of origin and dispersal of the Antarctic marine fauna. The influence of this process probably spreads
far northwards on both sides of South America, as witnessed by the fact that the East Pacific and West Atlantic are
565
the wcJ'ld oceanic portions showing maximum affinity
with the AFC.
The homogeneity of the continent is noteworthy, but
an expansion of the analysis also reveals some differentiation. For example, of the two entities including the
Antarctic Peninsula (C8 and C9) only C8 shows a moderately higher affinity with the Scotia Arc. Such a difference
is not ikely to be a biogeographic issue, but rather a
conseq~;ence of the fact that in C8 many samplings have
been made along the coasts of the Antarctic Peninsula,
wherea_~ most of the C9 records are from the Weddell Sea.
Another sector showing peculiar affinities is C5, whose
fauna is slightly more similar than the rest of the continent
to that ~f the relatively proximal entity of New Zealand.
NotwitiLstanding such a connection, New Zealand falls
definitely outside the AFC, indicating that the Antarctic
demosl:,~nge fauna does not support the hypothesis of
Udvarc y (1987), who considers New Zealand as a province
of his Antarctic Realm.
At ~:he generic level, the affinities of the AFC with
South ~!~friea,Tasmania and New Zealand are decisively
more ir~Lportant than at the species level. Moreover, New
Zealand and South Africa, and to a lesser extent Tasmania
and Sol.LthAustralia, show a high percentage of endemic
species, zomparable to that of the AFC. Such a coexistence
of both high specific endemism rates and generic homogeneity suggests a common Gondwanian origin of these
sponge ~aunas.
After Gondwana fragmentation, by the end of the
Mesozo:c, land masses pertaining to the Antarctic Continent m:~ved towards polar latitudes, thus experiencing
progres!ive climatic changes. This probably caused the
rise of ~ew selective pressures which could have triggered
intense !',peciation processes, also favoured by geographic
isolatio~,~t. Yet, a permanent connection with South America was guaranteed by the Antarctic Peninsula, which is
actually a portion of the Andean Orogen. This tectonic
connection is probably mirrored in the Magellan fauna
being ir~:zorporated within the AFC. In conclusion, what
we obse~rve today would be the relic of the Gondwanian
sponge paleofauna at the generic level and the predominant e~:'.ct of vicariance at the specific level.
The homogeneity of the AFC is primarily underlined
by the longitudinal distribution range of many of its
species, of which nearly a half covers, although discontinuously, 200 ~ or more. Within the domain of the
Antarct~: continental shelf, these wide distributions raise
no problem, since, as stressed by Koltun (1970), the shelf
shows a fair degree of environmental uniformity. Hence, it
is quite ,~:onceivable that species can diffuse from place to
place, tt:~us colonizing huge areas within time intervals
shorter than any reasonable estimate of species duration.
The ~ame cannot be said for a consistent number of
species which occur both in the continent and in cireumantarctic islands or in different islands only. Most of these
species have been sampled at depths roughly corresponding to the continental shelf and upper slope, no information existing about their presence at deeper levels. Starting
from this premise, a suite of hypotheses can be advanced to
account for these wide-range, apparently disjunct distributions.
Firstly, if one admits that these areas are really disconnected by intervening abyssal sea bottoms, then some
long-distance dispersal mechanism must be hypothesized.
The West Wind Drift is generally considered to be a main
homogenizing factor of the Antarctic fauna (White 1984),
but the existence in sponges of short-living lecithotrophic
larvae argues against the fitting of this model to Porifera.
However, a dispersal role could be played by floating
propagules such as buds or small sponge body fragments,
a mechanism already proposed by Burton (1932) to account for some correspondence between sponge distributions and oceanic current systems.
Secondly, as many shelf sponge species are supposed to
be eurybathic elements (Koltun 1970), one could think
that most Antarctic sponge species actually colonize abyssal domains, possibly with low population densities.
Hence their ranges, appearing disjunct as judged from
coastal sampling data, would be on the contrary continuous. A main consequence of such a hypothesis is that the
northern boundary of the Antarctic sponge fauna should
be imagined as a fuzzy contour penetrating far beyond the
Antarctic Convergence, possibly reaching with its extreme
fringes to the Arctic Ocean. The existence of bipolarlydistributed sponge species (Koltun 1970) is fairly in line
with this hypothesis.
Finally, a possibility that cannot be discarded is that
the supposed wide-range species could actually be complexes of cryptic species, highly conservative from a morphological point of view but genetically well separated.
This assumption is supported by the recent discovery of a
number of species complexes in allozymic studies of
sponge genetic variability (Sol6-Cava and Thorpe 1986,
1990; Sol6-Cava et al. 1991; Bavestrello and Sarfi 1992).
Based on the present knowledge, little can be said as to
which of the above hypotheses should be considered valid,
though the most reasonable explanation is probably a
combined one. Therefore, even though this study has
shown that the current level of knowledge of the Antarctic
demosponge fauna enables us to draw a reliable biogeographic picture, much remains to be learnt about the
dispersal mechanisms and the genetic structure of sponge
populations before a real understanding of the origin and
evolution of sponge faunas can be achieved.
Acknowledgements. We thank G. Pulitzer-Finalifor his help in the
analysis of synonymies,and M. Pansini and R. Pronzato for their
suggestionsand help in bibliographicresearch. We are also grateful
to the anonymous referees, whose criticism resulted in a definite
improvementofthe text.This workhas been supportedby the Italian
National Program for Antarctic Research (PNRA).
Table 2. Checklist and distribution of the demosponges of the Antarctic Faunistic Complex. Synonyms used in the reference literature are indicated in parentheses. Abbreviations for geographic
entities as in Table 1, and for the references as follows: Ba: Barthel et al. (1990) [Weddel Sea]; Be: Bergquist (1968, 1970), Bergquist and Warne (1980), Bergquist and F r o m o n t (1988) [New
Zealand]; Bo: Boury-Esnault and Van Beveren (1982) [Kerguelen-Heard]; DI: Desqueyroux (1975), Desqueyroux-Faflndez (1989) [Antarctic Peninsula]; D2: Desqueyroux and Moyano (1987)
[review on Magellan area]; K: Kottun (1964, 1976) [review on Antarctic continent and islands]; LI: L6vi (1956, 1964) [Kerguelen]; L2: L6vi (1963, 1967) [South Africa]; L3: L6vi (1969) [Vema
Seamount]; S: Sarfi et al. (1990) [Terra Nova Bay]; U: Uriz (1984, 1987, 1988) [Namibia]; V: Vacelet and Arnaud (1972) [Adblie Land]; W: Wiedenmayer (1989) [Tasmanian province]
Antarctic faunistic area
Continental sectors
C1
C2
C3
-
I
l
I
I
I
C4
Neighbouring
extra-Antarctic
entities
Non-continental entities
C5
C6
C8
C9
SG SS
SO M G F A
CR K E HE EM BO G O TC M Q C A
References
N Z TA SA VS
Homosclerophorida
1. Oscarella lobularis
(Schmidt, 1862) . . . . . . .
I
-
K
2. Plakina trilopha
Schulze, 1880 . . . . . . . .
I
I
I
I
I
-
I
I
-
!
Ba, Be, Bo, D2, K
3. Plakina monolopha
Schulze, 1880 . . . . . . . .
-
I
-
I
-
I
-
I
-
Ba, Be,
-
K
K,
Astrophorida
4. Cladothenea andriashevi
Koltun, 1964 . . . . . . . . .
5. Geodia magellani
(Sollas, 1886) . . . . . . . . .
I
D2
I
6. Geodinella vestigifera
Dendy, 1924 . . . . . . . . .
-
-
I
I
I
-
Be, K
-
Ba,
7. Monosyringa longispina
(Lendenfeld, 1907) . . . . .
I
I
I
K
8. Pachastrella monilifera
Schmidt, 1868 . . . . . . . .
I
I
9. Penares tylotaster
Dendy, 1924 . . . . . . . . .
10. Poecillastra compressa
(Bowerbank, 1866) . . . .
I
(Pachastrella, compressa
antarctica)
11. Poecillastra schulzii
.
.
.
.
I
I
(Sollas, 1886) . . . . . . . . .
12. Stelletta crater Dendy,
1924 . . . . . . . . . . . . . . .
13. Stelletta purpurea
Ridley, 1884 . . . . . . . . .
.
|
-
-
I
Bo,
I
K
K
Be, K
I
I
I
I
I
Be, K
I
I
(M yriastr a)
14. Stelletta maori Dendy,
1924 . . . . . . . . . . . . . . .
15. Stelletta phrissens
Sollas, 1886 . . . . . . . . .
16. Stelletta clarella De
Laubenfels, 1930 . . . . . .
17. Thenea delicata (Sollas,
1886) . . . . . . . . . . . . . . .
.
K, L2, U
I
I
I
-
Be, K, L2
I
Be, K
-
D2
-
D2
Bo,
K
S
Epipolasida
18. Jaspis novaezealandiae
D e n d y , 1924 . . . . . . . . .
-
I
Re !(_
|
19. Cinachyra coactifera
L e n d e n f e l d , 1907
(Tetilla) . . . . . . . . . . . . .
20. Cinachyra barbata
Sollas, 1886 . . . . . . . . .
21. Cinachyra antarctica
( C a r t e r , 1872) . . . . . . . .
22. Tetilla metaclada
( L e n d e n f e l d , 1907) . . . . .
23. Tetilla leptoderma
Sollas, 1886 . . . . . . . . . .
24. Tetilla coronida S o l l a s ,
1888 . . . . . . . . . . . . . . .
Bo, L1
I
-
I
I
I
I
-
!
!
-
I
I
I
I
I
I
I
I
!
!
I
I
I
Ba, Bo, D 1 , K , S
-
Ba, D 1 , K , V
!
-
!
-
I
I
I
I
-
Ba, K
I
I
Bo, D 1 , D 2 , K , S
I
I
Bo, K
Hadromerida
25. Aaptos unispiculus
( C a r t e r , 1880) . . . . . . . .
26. Ateryia acanthoxa
K o l t u n , 1964 . . . . . . . . .
27. Cliona euryphylla
T o p s e n t , 1888 . . . . . . . .
28. Cliona chilensis T h i e l e ,
1905 . . . . . . . . . . . . . . .
29. Cliona platei (Thiele,
1905) . . . . . . . . . . . . . . .
-
D2
-
!
K
I
-
I
.
i
.
.
.
.
.
.
.
.
I
.
Be
--
D2
.
(C liono psis)
D2
30. Cliona diversityla S a r a ,
1978 . . . . . . . . . . . . . . .
31. Cliona azzaroliae S a r a ,
1978 . . . . . . . . . . . . . . .
32. Cliothosa hancocki
( T o p s e n t , 1887) . . . . . . .
33. Latrunculia biformis
K i r k p a t r i c k , 1908 . . . . .
34. Latrunculia carlinae
Boury-Esnault & Van
B e v e r e n , 1982 . . . . . . . .
35. Latrunculia apicalis
Ridley & Dendy, 1886.
36. Latrunculia brevis
R i d l e y & D e n d y , 1886 .
-
I
!
!
!
-
(lendenfeldi, antarctica,
bocagei )
37. Polymastia isidis T h i e l e ,
-
I
I
-
I
1905 . . . . . . . . . . . . . . .
38., Polymastia invaginata
K i r k p a t r i c k , 1907 . . . . .
39. Proteleia burtoni
K o l t u n , 1964 . . . . . . . . .
-
D2
-
D2
D2
-
!
K
|
-
-
I
I
!
I
!
I
!
I
I
I
l
L
Bo
_
Ba, D1, K
I
I
I
-
I
I
_
!
I
I
-
-
I
I
I
I
I
-
I
Ba, Be, Bo, D 2 , K , U
I
Ba, Bo, D1, D 2 , K , U
Ba, Bo, D 1 , D 2 , K , V
K
Table
2 (continued)
oo
Antarctic faunistic area
Continental sectors
C1
Hadromerida (continued)
40. Pseudosuberites hyalinus
(Ridley & Dendy, 1887)
41. Pseudosuberites
digitatus Thiele, 1905..
42. Pseudosuberites melanos
de Laubenfels, 1934 . ..
43. Pseudosuberites
antarcticus (Carter,
1876) . . . . . . . . . . . . . . .
44. Pseudosuberites sulcatus
Thiele, 1905 . . . . . . . . .
45. Pseudosuberites mollis
Ridley & Dendy, 1887 .
C3
C4
C5
C6
C8
C9
I
I
I
I
-
I
I
I
(Suberites)
46. Pseudosuberites nudus
Koltun, 1964 . . . . . . . . .
47. Rhizaxinella
australiensis Hentschel,
1909 . . . . . . . . . . . . . . .
48. Sphaerotylus schoenus
Kirkpatrick, 1908 . . . . .
(schoenus vanhoffeni,
capitatus)
49. Sphaerotylus capitatus
Non-continental entities
C2
I
I
I
I
I
I
I
-
I
I
-
I
I
I
I
I
I
SO M G F A
CR KE HE EM BO G O TC M Q C A
N Z TA SA VS
I
I
-
I
-
I
I
I
I
Ba, D1,
I
D2
I
D2
I
I
I
I
I
I
I
I
I
I
-
K, U
Ba, Bo, K, U, V
Ba, Be, Bo, D1, D2, K
-
I
Ba, Bo, K
Ba, K
I
I
I
SG SS
References
Neighbouring
extra-Antarctic
entities
I
I
I
K
I
Ba, K
(Vosmaer, 1885) . . . . . .
(capitatus vanhoffeni)
50. Sphaerotylus borealis
antarcticus Kirkpatrick,
I
I
I
-
Ba, Bo, K, U
1908 . . . . . . . . . . . . . . .
(antarcticus)
51. Stylocordyla borealis
I
I
I
I
-
I
I
I
I
I
I
I
-
I
I
I
Ba, D1, K, V
(Loven, 1868) . . . . . . . .
(stipitata)
52. Suberites microstomus
Ridley & Dendy, 1887 .
53. Suberites montiniger
Carter, 1880 . . . . . . . . .
54. Suberites strongylatus
Sar~i, 1978. . . . . . . . . . .
55. Suberites punturatus
Thiele, 1905 . . . . . . . . .
56. Suberites caminatus
Ridley & Dendy, 1887.
I
-
I
I
-
I
I
-
I
Ba, Bo, D1, K, V
I
Ba, Bo
I
I
I
K
I
D2
I
I
I
I
I
I
-
D2
-
Ba,
Bo, K
57. Suberites ruber Thiele,
1905 . . . . . . . . . . . . . . .
58. Tentorium semisuberites
(Schmidt, 1870) . . . . . . .
Suberites caminatus pp.)
59. Tentorium papillatum
(Kirkpatrick, 1908) . . . .
-
-
I
I
I
I
I
-
I
D2
I
I
I
I
--
I
I
I
-
I
-
Ba, Bo, K
-
Ba,
Bo
Axinellida
60. Axinectya mariana
(Ridley & D e n d y , 1887)
(Axinella)
61. Axinella crinita Thiele,
1905 . . . . . . . . . . . . . . .
62. Axinella egregia (Ridley,
1881) . . . . . . . . . . . . . . .
I
Ba, Bo,
I
K
D2
-
D2
(Pseudaxinella)
63. Bubaris antarctica
K o l t u n , 1954 . . . . . . . . .
(Axinella)
I
I
64. Bubaris vermiculata
(Bowerbank, 1866) . . . .
65. Eurypon miniaceum
Thiele, 1905 . . . . . . . . .
66. Hemiastrella digitata
Burton, 1929 . . . . . . . . .
6% Higginsia papillosa
Thiele, 1905 . . . . . . . . .
68. Homaxinella
flagelliformis (Ridley &
D e n d y , 1886) . . . . . . . .
I
-
I
I
I
Be, Bo, D2, K
I
-
I
I
-
Bo,
U
D2
(Raspailia)
I
I
I
I
I
I
l
I
I
I
I
I
!
I
-
I
K
-
I
Bo, D1, K, L1, V
D2
I
l
-
I
I
I
I
-
I
-
I
-
I
-
Ba,
-
Ba, Bo,
K
K,
Bo
Be, K
-
I
I
I
-
-
I
-
D2, K , V
Halichondrida
76. Eumastia attenuata
Topsent, 1915 . . . . . . . .
D2,
K
I
69. Homaxinella
balfourensis (Ridley &
D e n d y , 1886) . . . . . . . .
70. Plicatellopsis flabellata
Burton, 1932 . . . . . . . . .
71. Plicatellopsis fragilis
K o l t u n , 1964 . . . . . . . . .
72. Raspailia irregularis
Hentschel, 1914 . . . . . .
(Eurypon miniaceum pp.)
73. Raspaxilla phakellina
Topsent, 1913 . . . . . . . . .
7 4 . Rhabderemia coralloides
D e n d y , 1924 . . . . . . . . .
75. Thieleia rubiginosa
(Thiele, 1905) . . . . . . . .
(H ymeniacidon)
Ba, D2, K
-
-
I
K
S, U
Table
2 (continued)
Neighbouring
extra-Antarctic
entities
A n t a r c t i c faunistic area
C o n t i n e n t a l sectors
C1
C2
C3
H a l i c h o n d r i d a (continued)
77. Halichondria cristata
Sarfi, 1978 . . . . . . . . . . .
78. Halichondria hentscheli
K o l t u n , 1964 . . . . . . . . .
79. Halichondria panicea
Pallas, 1766 . . . . . . . . .
80. Haliehondria variabilis
(Ridley, 1884) . . . . . . . .
81. H ymeniacidon torquata
Topsent, 1916 . . . . . . . .
82. Hymeniacidon fernandezi Thiele, 1905 . . . . . .
83. Hymeniacidon
N o n - c o n t i n e n t a l entities
C4
C5
C6
C8
C9
SG
SS
SO
MGFA
CR
KE
HE
EM BO GO TC
MQCA
NZ TA
References
SA
VS
D2
i
Ba, K
.
.
.
I
.
I
I
I
.
I
.
-
I
I
I
I
I
I
.
.
-
I
-
i
.
I
I
I
I
.
I
-
I
.
.
.
I
-
Be, Bo, K, L1
I
-
K
Ba, K
.
Bo, D2, K
I
longistylus
D e s q u e y r o u x , 1972 . . . .
84. Hymeniacidon centrotyla
Hentschel, 1914 . . . . . .
85. Hymeniacidon insuta
K o l t u n , 1964 . . . . . . . . .
86. Hymeniacidon
kerguelensis Hentschel,
1914 . . . . . . . . . . . . . . .
87. Leucophlaeus oxiparvus
Sarfi, 1978 . . . . . . . . . . .
88. Leucophaleus
membranaceus Sar~i,
1978 . . . . . . . . . . . . . . .
89. Leucophlaeus flexuosus
Sar/t, 1978 . . . . . . . . . . . .
90. Spongosorites compacta
Sar~i, 1978 . . . . . . . . . . .
91. Sponoosorites ineisa
Sar~i, 1978 . . . . . . . . . . .
i
D2
K
m
-
K
II
!
!
!
Bo, K
-
i
D2
D2
D2
D2
D2
Poecilosclerida
92. Acanthorhabdus fragilis
Burton, 1929 . . . . . . . . .
93. Acheliderma topsenti
Burton, 1932 . . . . . . . . .
94. Anomomycale titubans
(Schmidt, 1870) . . . . . . .
95. Artemisina jovis D e n d y ,
1924 . . . . . . . . . . . . . . .
96. Artemisina apollinis
(Ridley & D e n d y , 1886)
I
-
I
-
I
N
i
Ba, K
K
-
Bo
I
I
I
i
I
!
i
Be, K
Bo, D1, K
97.
Artemisina plumosa
( H e n t s c h e l , 1914) . . . . .
98.
I
Ko!tun, ! 964 ....
99.
!
I
!
|
-
-
Ba,
D1,
K
-~..z, K
....
-
K
!
!
!
!
!
Ba, K
I
-
K
!
K
!
Axociella rameus
Ba, K
-
!
!
-
!
!
!
!
!
!
I
I
I
!
I
I
I
Axociella fiabellata
( T o p s e n t , 1916) . . . . . . .
105.
-
I
Asbestopluma obae
Koltun, 1964 .........
104.
I
Asbestopluma calyx
Koltun, 1964 ........
103.
-
I
Hentschel, 1914 ......
102.
!
Asbestopluma belgicae
( T o p s e n t , 1902) . . . . . . .
101.
I
Asbestopluma callithrix
Hentschel, 1914 ......
100.
i
Artemisina tubulosa
Ba, K
Axociella nidificata
( K i r k p a t r i c k , 1907) . . . .
Biemna polyphylla L6vi,
1963 ...............
107. Biemna chilensis T h i e l e ,
1905 ...............
108. Bipocillopsis nexus
Koltun, 1964 .........
109. Cercidochela lankesteri
Kirkpatrick, 1907 .....
1 I0. Chondrocladia clavata
Ridley & Dendy, 1886 .
Ba, D1, K, S
!
106.
!
-
-
I
-
|
!
!
I
!
I
I
~
-
L2
Bo, D1, D2, K
!
-
-
!
Ba, K
I
I
Ba, D1, K
!
-
!
!
l
Be, B o , K
-
111. Chondrocladia
antarctica H e n t s c h e l ,
1914 ...............
112.
K
I
Chondrocladiafatimae
Boury-Esnault & Van
Beveren 1982 ........
113.
Boury-Esnault & Van
Beveren, 1982 ........
114.
-
!
K
!
-
!
-9-
K
!
-
K
!
-
K
Clathria pauper
!
I
!
-
|
I
Ba, D2, K
--
Clathria microxa
Desqueyroux,
120.
-
Cladorhiza tridentata
Broendsted, 1926 .....
119.
!
Cladorhiza mani
Ridley & Dendy, 1887 .
118.
Bo
Bo
g
Koltun, 1964 .........
117.
-
Cladorhiza moruliformis
Ridley & Dendy, 1886 .
116.
!
Chondropsis
chaliniformis ( C a r t e r ,
1886) . . . . . . . . . . . . . . .
115.
....
Chondrocladia nani
1972 ....
-
!
I
!
D2
Clathria terraenovae
Dendy, 1924 .........
(Dictyociona)
-
-
!
!
-
Be,
Bo,
K
Table
2 (continued)
Antarctic faunistic area
Continental sectors
C1
C2
C3
C4
Neighbouring
extra-Antarctic
entities
N o n - c o n t i n e n t a l entities
C5
C6
C8
C9
SG
SS
SO
MGFA
CR
KE
H E E M BO G O T C
MQCA
N Z TA
References
SA
VS
Poecilosclerida (continued)
121. Clathria " elastica" Sarfi
1978 . . . . . . . . . . . . . . .
(non C. elastica L~vi,
1963)
122. Clathria lipochela
Burton, 1932 . . . . . . . . .
123. Clathria spinifera Sara,
1978 . . . . . . . . . . . . . . .
124. Coelosphaera
D2
I
-
I
I
I
D2, K
I
D2
I
appendiculata antarctica
Koltun, 1976 . . . . . . . .
125. Coelosphaera #lobosa
Bergquist, 1961 . . . . . . .
126. Crella stylifera
Hentschel, 1914 . . . . . .
127. Crellina tubifex
Hentschel, 1914 . . . . . .
128. Desmacella vestibularis
(Wilson, 1904) . . . . . . . .
K
I
-
-
I
-
-
I
-
I
I
(Tylodesma)
Be, K
I
K
-
-
Bo,
I
I
I
I
129. DesmaceUa alba
(Wilson, 1904) . . . . . . . .
(Tylodesma)
130. Desmacidon nebulosum
Boury-Esnault & Van
Beveren, 1982 . . . . . . . .
131. Desmacidon fruticosa
(Montagu, 1818) . . . . . .
132. Desmacidon ramosa Ridley & Dendy, 1886 . . . .
I
no
!
-
-
I
-
I
-
I
-
I
I
I
K, L2
-
I
Ba, K, L2,
D2
K
I
D2
I
-
D2, K, U
Bo
-
-
(Plumocolumella,
ramosus)
133. Dictyociona discreta
(Thiele, 1905) . . . . . . . .
134. Dolichocantha macrodon
Hentschel, 1914 . . . . . .
135. Echinoclathria contexta
Sarh, 1978 . . . . . . . . . . .
136. Echinoclathria atlantica
Sarfi, 1978 . . . . . . . . . . .
137. Ectyodoryx antarctica
(Hentschel, 1914) . . . . .
138. Ectyodoryx ramilobosa
(Topsent, 1916) . . . . . . .
K
I
-
I
I
I
I
-
D2
K
-
Ba, K, V
U
139.
Ecryodoryx nobilis
(Ridley & Dendy, 1886)
140.
141.
(paupertas nobile)
Ectyodoryx anacantha
-
rlentscnel,
-
I
I
-
I
I
1~14 ......
I
I
Ectyodoryx plumosa
Hentschel, 1914 . . . . . .
142.
I
I
I
I
I
I
I
Ectyomyxilla
kerguelensis Hentschel,
-
Bo, K
-
Ba, K
-
K
1914 . . . . . . . . . . . . . . .
(Crellomyxilla, Myxilla)
143. Ectyomyxilla mariana
-
I
-
l
Be, Bo, L1, L2
I
(Ridley & Dendy, 1887)
144.
(Crellomyxilla)
Ectyomyxilla tenuissima
Bo, K
(Thiele, 1905) . . . . . . . .
145.
Ectyomyxilla chilensis
I
-
D2, L2
!
-
Bo, D1, D2, K, L1, U
(Thiele, 1905) . . . . . . . .
146.
(Crellomyxilla, M yxilla)
Ectyonancora panis
I
I
I
l
-
I
I
I
--
Boury-Esnault & Van
Beveren, 1982 . . . . . . . .
147.
Esperiopsis villosa
148.
(Carter, 1874) . . . . . . . .
Esperiopsis varia Sara,
1978 . . . . . . . . . . . . . . .
149.
Esperiopsis heard!
-
I
I
!
|
I
-
-
Bo
-
K
-
D2
I
Boury-Esnault & Van
Beveren, 1982 . . . . . . . .
150.
|
-
Esperiopsis symmetrica
Bo
I
Ridley & Dendy, 1886.
I
151. Esperiopsis scoriae
K
-
Topsent, 1915 . . . . . . . .
152.
K
Esperiopsis rugosa
Thiele, 1905 . . . . . . . . .
153.
(Amphilectus)
Esperiopsis profunda
I
I
I
I
-
l
Ridley & Dendy, 1886 .
154.
I
!
l
--
|
-
I
I
-
i
I
i
!
-
Ba, Bo, K, U
K
_
Esperiopsis fucorum
(Esper, 1974) . . . . . . . . .
155.
156.
157.
(Amphilectus)
Esperiopsis edwardii
(Bowerbank, 1866) . . . .
Guitarra antarctica var.
novaezealandiae Dendy,
1924 . . . . . . . . . . . . . . .
([imbriata)
Guitarra fimbriata
Carter, 1874 . . . . . . . . .
158.
I
Homoeodictya
microchela Topsent,
-
I
-
I
I
I
-
I
a
i
l
-
I
-
K
I
Be, D 2
!
Be, K
!
I
I
!
Ba, Be, L2
1915 . . . . . . . . . . . . . . . .
(lsodictya)
-
I
I
-
I
-
I
-
K
Table 2 (continued)
Antarctic faunistic area
C o n t i n e n t a l sectors
C1
Neighbouring
extra-Antarctic
entities
N o n - c o n t i n e n t a l entities
C2
C3
C4
C5
1
I
1
I
C6
C8
C9
SG
SS
SO
MGFA
CR
KE
HE
EM BO GO TC
MQCA
NZ TA
References
SA
VS
Poecilosclerida (continued)
159.
Hoplakithara dendyi
Kirkpatrick, 1908 . . . . .
160.
I
Ba, K
Hymedesmia
mariondufresni B o u r y E s n a u l t & V a n Beveren,
1982 . . . . . . . . . . . . . . .
161.
-
H ymedesmia irritans
(Thiele, 1905) . . . . . . . .
162.
-
I
D2
I
I
D2
H ymedesmia 9aussiana
Hentschel, 1914 . . . . . .
164.
Bo
Hymedesmia laevis
Thiele, 1905 . . . . . . . . .
163.
I
-
K
Hymedesmia antarctica
(Hentschel, 1914) . . . . .
165.
(simillima antarctica)
H ymedesmia unguifera
I
1
I
Burton, 1929 . . . . . . . . .
166.
|
.
.
.
.
.
.
-
K
K
K
m
(.
I
1
-
K
Hymenancora exioua
1
-
K
InflateUa beUi
(Kirkpatrick, 1907) . . . .
I
1
1
1
I
1
1
1
I
I
I
1
I
l
170. Inflatella latrunculioides
(Ridley & D e n d y , 1887)
171.
Inflatella
coelosphaeroides K o l t u n ,
172.
lophon chelifer Ridley
1964 . . . . . . . . . . . . . . .
1
-
l
-
l
I
Ba, Bo, K, S, U, V
Ba, K
1
Ba, K
& D e n d y , 1886 . . . . . . .
173.
Bo,
-
Hymenancora tufa
(Kirkpatriek, 1907) . . . .
169.
I
I
(Kirkpatriek, 1907) . . . .
168.
I
Hymenancora
rhaphidophora
Hentschel, 1914 . . . . . .
167.
I
1
I
Bo,
L2,
U
lophon unicornis
Topsent, 1907 . . . . . . . .
(spatulatus)
174. Iophon abnormalis
I
I
1
I
-
1
I
1
1
I
l
I
l
Ridley & D e n d y , 1886 .
175.
Iophon aceratus
176.
Hentschel, 1914 . . . . . .
Iophon laminalis Ridley
& Dendy, 1886 . . . . . . .
Bo, D1, K, S, V
l
K
1
!
I
I
I
-
I
Ba,
K
K
177. Iophon proximum
(Ridley, 1881) . . . . . . . .
178. Iophon radiatus
Topsent, 1901 . . . . . . . .
!79. I oph~n !ae::i::ty!::~
Dendy, 1924 . . . . . . . . .
180. Iotrochota somovi
(Koltun, 1964) . . . . . . . .
I
I
(M yxichela)
I
I
I
-
I
I
I
I
I
I
I
I
I
-
Be, Bo, D2, K, L2, U
Ba, Bo, K
I
Be, K
(Iotaota)
181. Isodictya cavicornuta
Dendy, 1924 . . . . . . . . .
182. Isodictya dufresni
Boury-Esnault & Van
Beveren, 1982 . . . . . . . .
183. Isodictya erinacea
(Topsent, 1916) . . . . . . .
184. Isodictya cactoides
(Kirkpatrick, 1908) . . . .
185. Isodictya verrucosa
(Topsent, 1913) . . . . . . .
186. Isodietya delicata
megachela (Thiele, 1905)
187. lsodictya toxophila
Burton, 1932 . . . . . . . . .
188. Isodictya obliquidens
(Hentschel, 1914) . . . . .
189. Isodictya antarctica
(Kirkpatrick, 1908) . . . .
190. lsodictya setiJera
(Topsent, 1901) . . . . . . .
191. lsodictya kerguelensis
(Ridley & Dendy, 1886)
192. Kirkpatrickia coulmani
(Kirkpatrick, 1907) . . . .
193. Kirkpatrickia variolosa
(Kirkpatrick, 1907) . . . .
194. Lissodendoryx
paucispinata (Ridley &
Dendy, 1887) . . . . . . . .
195. Lissodendoryx
complicata (Hansen,
1885) . . . . . . . . . . . . . . .
196. Lissodendoryx styloderma Hentschel, 1914.
197. Lissodendoryx flabellata
Burton, 1929 . . . . . . . . .
198. Lissodendoryx
innominata Burton, 1929
199. Lissodendoryx fusca
(Ridley & Dendy, 1886)
200. Megaciella pilosa
(Ridley & Dendy, 1886)
I
-
l
.
.
.
-
l
I
l
.
Ba, K
-
I
I
Bo
I
I
I
I
I
I
I
I
I
Ba, D1, K, S, V
I
K
I
I
I
-
I
I
-
I
I
I
I
I
I
I
I
I
I
I
I
I
Ba, Be, K
I
I
D1, K
I
I
Ba, K
I
Ba, K
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
-
Bo
-
I
I
I
I
I
Ba, K, V
I
Ba, K
I
I
-
Ba, Bo, D1, K, L1, V
I
K,V
I
Ba, K
Bo
Bo
I
|
I
--
I
.
Ba, K
I
I
K,V
I
Ba, K
Bo, K
I
Ba, Bo, K
Table
2
(continued)
N o n - c o n t i n e n t a l e nt i t i e s
C o n t i n e n t a l sectors
C1
C2
C3
C4
C5
C6
C8
C9
SG
SS, S O
MGFA
References
Neighbouring
extra-Antarctic
en tities
A n t a r c t i c faunistic a r e a
CR
KE
HE
EM BO GO TC
MQCA
NZ
TA
SA
VS
Poecilosc/erida (continued)
201.
Meliiderma stipitata
(Ridley & D e n d y , 1886)
202.
-
-
Burton, 1934 . . . . . . . . .
203.
I
I
D2, K
-
K
-
I
I
I
I
-
-
I
I
-
-
I
I
I
|
--
Ba, D2, K
_
M ycale mammiformis
(Ridley & Dendy, 1886)
206.
Mycale tridens
207.
Hentschel, 1914 . . . . . .
Mycale tenuis Sar~t,
1978 . . . . . . . . . . . . . . .
208.
Mycale doellojuradoi
Burton, 1940 . . . . . . . . .
209.
I
Mycate magellanica
(Ridley, 188l) . . . . . . . .
205.
K
-
Mycale tylotornota
K o l t u n , 1964 . . . . . . . .
204.
I
I
Microciona basispinosa
-
K
I
-
I
I
-
I
I
-
-
-
I
Ba, D2, K
I
D2
I
D2
I
Ba, D2, K
M ycale gaussiana
Hentschel, 1914 . . . . . .
-
!
!
I
I
--
|
I
210. Mycale profunda
Koltu n, 1964 . . . . . . . . .
21 I.
-
K
MycaIefibrosa
Boury-Esnault & Van
Beveren, 1982 . . . . . . . .
212.
Mycale macrochela
213.
Burton, 1932 . . . . . . . . .
M ycale diminuta Sara,
1978 . . . . . . . . . . . . . . .
214.
M yxilla novaezealandiae
Dendy, 1924 . . . . . . . . .
215.
Myxilla pistiUaris
216.
Topsent, 1917 . . . . . . . .
Myxilla mollis Ridley &
Dendy, 1886 . . . . . . . . .
-
I
-
-
|
!
--
-
Bo, D1
--
K
D2
I
-
I
-
-
!
I
I
-
l
-
l
I
I
I
l
l
I
-
-
Be, K
-
K
-
Ba, Bo, D1,
217. Myxilla decepta
K i r k p a t r i c k , 1907 . . . . .
218.
Topsent, 1916 . . . . . . . .
219.
-
!
!
I
l
I
l
-
I
I
l
-
Ba, K
I
Ba, K
I
Ba, K
l
M yxilla basimucronata
Burton, 1932 . . . . . . . . .
222.
-
Myxilla insolens
Koltun, 1964 . . . . . . . . .
221.
l
MyxiUa asigmata
Topsent, 1901 . . . . . . . .
220.
K
Myxilla elongata
-
I
Bo,
K
M yxilla lissostyla
Burton, 1938 . . . . . . . . .
!
I
Ba, D1, K
K
223. MyxiUa australis
(Topsent, 1901) . . . . . . .
(incrustans australis)
-
I
224. Myxilla calieifnrmi~
Sara, 1978 . . . . . . . . . . .
225. Myxilla verrucosa
B u r t o n , 1932 . . . . . . . . .
226. Myxodoryx hanitschi
(Kirkpatrick, 1908) . . . .
227. Ophlitaspongia mutans
Sar~i, 1978 . . . . . . . . . . .
228. Ophlitaspongia
membranacea Thiele,
1905 . . . . . . . . . . . . . . .
I
I
I
I
I
Ba, D1, K, S
I
-
i
t
-
l
I
D2
I
D2
Ba, D1, K
i
D2
(thielei)
D1, D2, K
I
-
I
-
I
I
I
I
I
I
I
i
-
229. Oxymycale acerata
(Kirkpatrick, 1907) . . . .
(M ycale)
I
I
I
I
230. Oxytedania bifaria Sara,
1978 . . . . . . . . . . . . . . .
231. Phelloderma radiatum
Ridley & D e n d y , 1887 .
232. Phorbas areolata
(Thiele, 1905) . . . . . . . .
(Anchinoe, Hymedesmia)
Ba, Bo, D 1 , D 2 , K , L1, S , V
I
D2
K
-
I
I
l
-
l
I
D1, D2, K, V
I
233. Phorbas acanthochela
(Koltun, 1964) . . . . . . . .
(Anchinoe)
i
K,S
I
234. Phorbas leptochela
(Hentschel, 1914) . . . . .
(Anchinoe)
-
I
Bo,
i
K
235. Phorbas longurioides
(Burton, 1932) . . . . . . . .
(Hymedesmia, Anchinoe)
Bo, K
I
236. Phorbas glaberrima
(Topsent, 1917) . . . . . . .
(Anchinoe)
237. Plocamia gaussiana
Hentschel, 1914 . . . . . .
238. Plumocolumella
cribroporosa Burton,
1929 . . . . . . . . . . . . . . .
239. Plumocolumella
meandrina (Kirkpatrick,
1907) . . . . . . . . . . . . . . .
240. Pronax domini
Boury-Esnault & Van
Beveren, 1982 . . . . . . . .
241. Pronax pustulosa
(Carter) . . . . . . . . . . . . .
242. Pseudanchinoe papillosa
(Thiele, 1905) . . . . . . . .
(Clathria)
I
I
I
I
I
I
I
I
I
I
--
!
Ba, K
--
K
I
Ba, K
I
K,V
!
i
i
-
.
.
.
Bo
.
!
L2
D2
Table 2 (continued)
Oo
N o n - c o n t i n e n t a l entities
Continental sectors
C1
C2
C3
C4
C5
I
-
I
C6
C8
C9
SG
I
I
I
SS
SO
MGFA
References
Neighbouring
extra-Antarctic
entities
Antarctic faunistic area
CR
KE
HE
E M BO G O T C
MQCA
N Z TA
SA
VS
Poecilosclerida (continued)
243. Pseudanehinoe toxifera
(Topsent, 1913) . . . . . . .
244. Pseudomyxilla
multidentata BouryEsnault & Van Beveren,
1982 . . . . . . . . . . . . . . .
I
I
-
-
Ba, D1, K
I
Bo
I
245. Pytheas crassa
Hentschel, 1914 . . . . . .
(Crella)
I
I
I
I
-
I
I
I
I
-
I
Ba, Bo, K
246. Rhaphidophlus
toxipraeditus (Topsent,
1913) . . . . . . . . . . . . . . .
( C lat hria)
I
247. Rhaphidophlus
paucispiculus Burton,
1932 . . . . . . . . . . . . . . .
248. Spanioplon werthi
(Hentschel, 1914) . . . . .
(Acanthoxa)
-
-
I
.
.
.
.
I
I
I
Ba, K
I
I
D1, D2, K
I
Bo,
I
249. Stylopus longurius
antarcticus (Hentschel,
1914) . . . . . . . . . . . . . . .
(H ymedesmia)
!
!
I
K,V
I
250. Stylopus fristedti
Topsent, 1917 . . . . . . . .
(H ymede smia )
1
K
-
251. Stylotella aominata
(Ridley, 1884) . . . . . . . .
(H ymeniacido n)
252. StyloteUopsis antarcticus
Koltun, 1964 . . . . . . . .
253. Stylotellopsis amabilis
Thiele, 1905 . . . . . . . . .
254. Tedania laminariae
Sara, 1978 . . . . . . . . . . .
255. Tedania inflata Sarfi,
1978 . . . . . . . . . . . . . . .
256. Tedania charcoti
Topsent, 1908 . . . . . . . .
257. Tedania fueyensis
Thiele, 1905 . . . . . . . . .
258. Tedania trirhaphis
Koltun, 1964 . . . . . . . . .
259. Tedania pectinicola
Thiele, 1905 . . . . . . . . .
K
I
I
-
I
Be, Bo, K
K
I
D2, K
I
D2
D2
l
-
I
I
-
I
I
I
I
I
I
I
I
I
-
Ba, Bo, D1, D2, K, L1
D2
I
-
l
Ba,
I
I
-
D2
K
260. Tedania armata Sarfi,
1978 . . . . . . . . . . . . . . .
261. Tedania tenuicapita
Ridley, 1881 . . . . . . . . .
262. Tedania vanhoeffeni
Hentschel, 1914 . . . . . .
263. Tedania spinata (Ridley,
1881) . . . . . . . . . . . . . . .
264. Tedania corticata Sar~i,
1978 . . . . . . . . . . . . . . .
265. Tedania lanceta Koltun,
1964 . . . . . . . . . . . . . . .
266. Tedania gracilis
(Hentschel, 1914) . . . . .
267. Tedania tantula
(Kirkpatrick, 1908) . . . .
268. Tedania oxeata
Topsent, 1916 . . . . . . . .
269. Tedania mucosa Thiele,
1905 . . . . . . . . . . . . . . .
270. Tedania patagonica
(Ridley & Dendy, 1887)
271. Tedania excavata
Thiele, 1905 . . . . . . . . .
D2
D2
I
I
Ba, Bo, K
I
I
I
D2, K
-
D2
K
!
I
K,V
I
I
I
I
I
I
-
I
I
I
I
I
-
I
I
Ba, K, V
I
Ba, D1, K
D2
Bo,
!
K
D2
Petrosida
272. Petrosia hispida Ridley
& Dendy, 1886 . . . . . . .
(Haliclona)
I
I
Bo, K, U
273. Xestospongia fistulata
(Kirkpatrick, 1907) . . . .
(Pachypellina}
I
K
I
274. Xestospongia aulopora
(Schmidt) . . . . . . . . . . .
D2
Haplosclerida
275. Adocia conica (Thiele,
1905) . . . . . . . . . . . . . . .
276. Adocia tenella (Topsent,
1916) . . . . . . . . . . . . . . .
!
I
-
I
I
K
D1, K
277. Amphimedon decurtata
(Sar~i, 1978) . . . . . . . . . .
(Pachychalina)
D2
278. Amphimedon validissima
(Thiele, 1905) . . . . . . . .
(Pachychalina,
Dasychalina)
279. Amphimedon magellanica (Thiele, 1905) . . . . .
(Pachychalina)
280. Amphimedon paradisus
Desqueyroux-Fa6ndez,
1989 . . . . . . . . . . . . . . .
D2, K
D2
DI
Table
2 (continued)
Non-continental entities
Continental sectors
C1
C2
C3
C4
C5
C6
C8
C9
SG SS
SO M G F A
References
Neighbouring
extra-Antarctic
entities
Antarctic faunistic area
CR KE HE EM BO G O TC
MQCA
NZ TA SA VS
Haplosclerida (continued)
281. Amphimedon anomala
(Sara, 1978) . . . . . . . . . .
D2
I
(P achychalina)
282. Amphimedon maresi
(Sarfi, 1978) . . . . . . . . . .
284. Callyspongia communis
(Carter, 1881) . . . . . . . .
285. Callyspongia fusifera
(Thiele, 1905) . . . . . . . .
286. Callyspongia fortis
Ridley, 1881 . . . . . . . . .
287. Callyspongia gaussiana
(Hentschel, 1914) . . . . .
-
I
-
(Broendsted, 1924) . . . .
(Haliclona)
288. Callysponoia ramosa
D2
I
(P achychalina)
283. Callyspongia conica
Be
-
D2
I
-
I
I
I
I
I
I
I
-
D2
D2
-
K
-
(Gray, 1843) . . . . . . . . .
(Cladochalina)
289. GeUius kerguelensis
I
Be, K, W
I
-
(Hentschel, 1914) . . . . .
(Gelliodes, Calyx)
290. Gellius bidens Topsent,
I
I
I
I
I
I
I
Ba, Bo, K
I
1902 . . . . . . . . . . . . . . .
(H aliclona,
Plumocolumella)
291. Gellius latisigmae
l
I
I
I
Ba, K, V
Boury-Esnault & Van
Beveren, 1982 . . . . . . . .
Bo
I
292. Gellius constans
Boury-Esnault & Van
Beveren, 1982 . . . . . . . .
293. Gellius flagellifer Ridley
& Dendy, 1886 . . . . . . .
(Adocia, Si#madocia)
294. Gellius calyx Ridley &
I
-
I
I
-
I
I
-
I
-
Bo
I
Dendy, 1887 . . . . . . . . .
(Haliclona)
295. Gellius carduus Ridley
I
I
I
-
I
-
1
l
I
I
-
I
I
Be, Bo, K, L3, U
Ba, K
& Dendy, 1886 . . . . . . .
(Adocia)
I
l
l
l
Bo,
K
296. Gellius cucurbitiformis
Kirkpatrick, 1907 . . . . .
(Adocia)
I
l
l
I
D1, K, V
-
297. Gellius flabelliformis
P.id!ey & Dendy: ~ . ~
(Orina)
Bo, U
298. Haliclona pedunculata
(Ridley & Dendy, 1887)
(Reniera)
-
!
Bo, K, V
i
299. Haliclona similis (Ridley
& Dendy, 1887) . . . . . .
300. Haiiclona sordida
(Thiele, 1905) . . . . . . . .
301. Haliclona spongiosissima
(Topsent, 1908) . . . . . . .
302. Haliclona chilensis
(Thiele, 1905) . . . . . . . .
303. Haliclona bilamellata
Burton, 1932 . . . . . . . . .
-
-
I
I
Bo, K, L1
i
D2
I
D1, K
I
(Reniera)
304. Haliclona altera
(Topsent, 1901) . . . . . . .
305. Haliclona divulgata
Koltun, 1964 . . . . . . . . .
306. Haliclona texta Sani,
1978 . . . . . . . . . . . . . . .
307. Haliclona eterospiculata
(Sarh, 1978) . . . . . . . . . .
I
-
I
I
I
-
l
I
I
D2
I
I
Ba, D1, D2, K
D1, K
Bo, K
D2
i
(Reniera)
D2
308. Haliclona topsenti
(Thiele, 1905) . . . . . . . .
(Reniera, foraminosa)
309. Haliclona gemina Sarh,
1978 . . . . . . . . . . . . . . .
310. Haliclona tylotoxus
(Hentschel, 1914) . . . . .
I
-
!
-
!
!
-
|
Bo, D1, D2, K, L1, U
D2
!
-
-
!
K
311. H aliclona ionobilis
(Thiele, 1905) . . . . . . . .
312. Haliclona variabilis
(Thiele, 1905) . . . . . . . .
(Reniera fortior)
!
!
-
D2
i
i
-
D2
313. Haliclona delicata (Sars
1978) . . . . . . . . . . . . . . .
(Reniera)
-
i
i
D2, U
314. Haliclona macropora
(Thiele, 1905) . . . . . . . .
(Reniera)
315. Haliclona virens
Topsent, 1908 . . . . . . . .
I
-
!
-
I
--
D2
D2, K
i
316. Haliclona penieillata
(Topsent, 1908) . . . . . . .
317. Haliclona bifacialis
Sani, 1978 . . . . . . . . . . .
K
i
-
D2
Table 2 (continued)
Continental sectors
C1
C2
C3
C4
Non-continental entities
C5
C6
C8
C9
SG SS
SO
MGFA
References
Neighbouring
extra-Antarctic
entities
Antarctic faunistic area
CR KE HE EM BO G O TC M Q C A
NZ TA SA VS
Haplosclerida (continued)
318. Haliclona algicola
(Thiele, 1905) . . . . . . . .
(Reniera)
319. Haliclona proletaria
(Topsent, 1908) . . . . . . .
320. Haliclona dancoi (Topsent, 1902). . . . . . . . . . .
321. Haliclona nodosa
(Thiele, 1905) . . . . . . . .
322. Haliclonissa sacciformis
(Burton, 1932) . . . . . . . .
I
I
l
l
I
K
--
i
-
l
I
-
Ba, K
I
K
I
I
(H aliclona)
I
323. Haliclonissa verrucosa
Burton, 1932 . . . . . . . . .
324. Hemigellius pilosus
(Kirkpatrick, 1907) . . . .
l
(Haliclona)
|
325. HemigeUius pachyderma
Burton, 1932 . . . . . . . . .
326. Hemigellius fimbriatus
(Kirkpatrick, 1907) . . . .
(Gellius rudis, Haliclona
rudis)
327. Microxina simplex
(Topsent, 1916) . . . . . . .
328. Microxina benedeni
(Topsent, 1901) . . . . . . .
329. Orina phakelloides
(Kirkpatrick, 1907) . . . .
D2
i
l
I
Bo, U
I
Ba, K
I
K
--
D1, K
I
I
-
I
I
-
l
I
I
I
I
Ba, Bo, D1, K, V
I
I
Ba, K
i
I
(Haliclona, Gellius)
i
i
i
I
I
I
I
l
!
I
I
-
-
Ba, D1,
Ba, K
I
330. Orina spongiosa
(Topsent, 1916) . . . . . . .
(Haliclona)
331. Sigmadocia glacialis
I
I
I
l
I
l
K
K
(Ridley & Dendy, 1886)
(Haliclona, Adocia,
Gellius)
332. Sigmadocia
tubuloramosus (Dendy,
I
I
l
l
I
1924) . . . . . . . . . . . . . . .
333. Fagocia arcuarius (Topsent, 1913) . . . . . . . . . . .
(Calyx)
l
l
|
I
I
I
I
l
I
I
I
l
I
I
-
Ba, Be, Bo, D1, K, U
Be
Ba, D1, K, S
Dictyoceratida
334.
Dysidea oculata
(Burton, 1929) . . . . . . . .
-
I
K
I
D2
(Thiele, 1905) . . . . . . . .
336.
Dysidea fragilis
(Montagu, 18t4) . . . . . .
337.
I
I
I
I
-
Dysidea tenuifibra
Euryspongia repens
339.
(Thiele, 1905) . . . . . . . .
Hyrtios arenosa (Thiele,
1905) . . . . . . . . . . . . . . .
340.
U
D2
(Burton, 1932) . . . . . . . .
338.
Bo,
D2
(Oligoceras)
Hyrtios vinciguerrae
K
I
(Sarfi, 1978) . . . . . . . . . .
D2
(Oligoceras)
341. Ircinia variabilis
D2, K
I
Schmidt, 1862 . . . . . . . .
342. Ircinia clavata Thiele,
1905 . . . . . . . . . . . . . . .
343. Sponoia cerebralis
Thiele, 1905 . . . . . . . . .
344. Spongia hispida
Lamarck, 1814 . . . . . . .
345. Spongia maoellanica
Thiele, 1905 . . . . . . . . .
346. Spongionella pulchella
(Soverby, 1804) . . . . . . .
D2
D2
I
.
.
.
.
D2, W
I
D2, K
I
Bo
I
Dendroceratida
347.
348.
Aplysilla rosea (Barrois,
1876) . . . . . . . . . . . . . . .
(sulphurea)
Dendrilla antarctica
I
Topsent, 1905 . . . . . . . .
I
Dictyodendrilla membranosa (Pallas) . . . . . . .
(Dendrilla)
350. Halisarca magellanica
-
-
I
I
I
Bo, K, U, W
I
D1
I
I
349.
I
I
I
Topsent, 1901 . . . . . . . .
351.
I
-
I
I
I
I
I
I
--
-
Ba, Bo, D1,
D2
Halisarca dujardini
Johnston, 1842 . . . . . . .
I
I
I
I
-
I
-
I
I
Bo, K, U
Verongida
352.
Aplysina minima
Hentschel, 1914 . . . . . .
I
I
K
K
584
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