Svenzea, a new genus of Dictyonellidae (Porifera: Demospongiae) from tropical reef environments, with description of two new species

Abstract

The new genus Svenzea is created to group three sponge species from tropical reef environments of the Caribbean Sea and Indonesia: Pseudaxinella (?) zeai Alvarez, Van Soest & Rützler, Svenzea cristinae n. sp. and S. devoogdae n. sp. The genus shows affinities with members of both Halichondrida and Haplosclerida but it is assigned to the family Dictyonellidae based on shared microanatomical and developmental features. The higher taxonomic position of Svenzea is a subject for future investigations.

Full text

Contributions
to
Zoology,
71
(4)
171-176
(2002)
SPB
Academic
Publishing
bv,
The
Hague
Svenzea
,
a
new
genus
of
Dictyonellidae
(Porifera:
Demospongiae)
from
tropical
reef
environments,
with
description
of
two
new
species
Belinda
Alvarez
¹,
Rob+W.M.
van
Soest
²
&
Klaus
Rützler
³
Keywords:
Porifera,
Demospongiae,
Dictyonellidae,
new
genus,
new
species,
Svenzea
cristinae,
Svenzea
devoogdae
Abstract
The
new
genus
Svenzea
is
created
to
group
three
sponge
spe-
cies
from
tropical
reef
environments
of
the
Caribbean
Sea
and
Indonesia:
Pseudaxinella
(?)
zeai
Alvarez,
Van
Soest
&
Rützler,
Svenzea
cristinae
n.
sp. and
S.
devoogdae
n.
sp.
The
genus
shows
affinities
with
members
of
both
Halichondrida
and
Haplosclerida
but
it
is
assigned
to
the
family
Dictyonellidae
based
on
shared
microanatomical
and
developmental
features.
The
higher
taxo-
nomic
position
of
Svenzea
is
a
subject
for
future
investigations.
Contents
Introduction
171
Systematics
172
Discussion
175
Acknowledgements
175
References
176
Intruduction
Material
examined
in
this
work
is
deposited
at
the
National
Museum
of
Natural
History,
Washing-
ton
D.C.
(USNM),
Zoological
Museum
of
Amster-
dam
(ZMA),
National
Natural
History
Museum,
Leiden
(RMNH)
and
Museum
and
Art
Gallery
of
the
Northern
Territory,
Darwin
(NTM).
Thick
sec-
tions,
spicule
slides,
Transmission
and
Scanning
Electron
Microscopy
(TEM,
SEM)
preparations
were
made
using
the
methods
described
by
Riitzler
(1978),
Riitzler
et
al.
(in
press),
and
Hooper)
1996).
1
Museum
and
Art
Gallery
of
the
Northern
Territory,
PO
Box
4646,
Darwin
NT
0801,
Australia.
E-mail:
belinda.glasby@nt.gov.au;
2
Zoological
Museum,
Faculty
of
Science,
University
of
Amsterdam,
PO
Box
94766,
1090
GT,
Amsterdam,
The
Netherlands.
E-mail:
soest@science.uva.nl;
department
of
Inver-
tebrate
Zoology.
National
Museum
of
Natural
History,
Smithsonian
Institution.
Washington
DC
20560-0163,
U.S.A.
E-mail:
ruetzler.klaus@nmnh.si.edu
Van
Soest
et
ah,
1990).
sensu
The
discovery
of
two
additional
species
with
simi-
lar
skeletal
structure
and
spicule
composition
jus-
tifies
the
erection
of
a
new
genus
which,
based
on
histological
and
development
evidence
(Riitzler
et
ah,
in
press)
is
assigned
to
the
family
Dictyonellidae
of
the
order
Halichondrida
(
are
styles
with
a
small
percentage
of
oxeas,
such
as
in
many
axinellid
species,
and
also
in
a
similar
size
range,
and
unlike
the
spicule
combination
found
in
the
order
Haplo-
sclerida.
P.
zeai
(Haplosclerida:
Petrosiidae).
The
spicules
that
form
the
skeleton
of
Xesto-
spongia
(Halichondrida:
Dictyonellidae).
The
consistency
is
firm
but
crumbly
as
in
some
species
of
Scopalina
tion
common
in
Haplosclerida
and
obscured
by
masses
of
granular
cells,
also
present
in
is
a
nearly
unispicular
reticula-
P.
zeai
Alvarez
&
Hooper,
2002)
of
the
family
Axinellidae
was
doubtful
at
that time
as
the
species
shows
a
mixture
of
fea-
tures
of
haplosclerid
and
halichondrid
sponges.
The
skeleton
of
Dragmacidon
(now
Pseuda-
xinella
(?)
zeai.
The
assignment
of
the
species
to
Pseudaxinella
and
in
field
guides
(i.e.
Humann,
1992)
was
de-
scribed
by
Alvarez
et
al.
(1998)
as
1997a)
etal.,
1998;
Rodriguez
etal.,
(De
Laubenfels,
1934)
in
chemical
literature
(e.g.
Carballeira
Calyx
poda-
typa
A
very
common
sponge
species
found
in
the
Car-
ibbean
and
generally
mis-identified
as

B.
Alvarez
et
al.
—
Svenzea,
new
sponge
genus
from
tropical
reefs
172
Systematica
Order
Halichondrida
Gray,
1867
Family
Dictyonellidae
Van
Soest
et
al.
1990
Svenzea,
new
genus
Type
species.
-
Pseudaxinella
zeai
Alvarez
et
al
1998:
20
(Fig.
1).
Diagnosis.
-
Massive
or
thickly
encrusting.
Sur-
face
smooth
but
microscopically
hispid.
Ectosomal
skeleton
specialization
absent.
Choanosomal
skel-
eton
formed
by
an
uni-
or
paucispicular
reticula-
tion
of
megascleres
with
spongin
cementing
the
spicules
at
the
nodes.
Internal
structure
on
thick
sections
viewed
in
transmitted
light
obscured
by
numerous
black-looking
cells
of
18-35
pm
diam-
eter,
with
3
pm
spherical
or
angular
granules.
All
known
species
are
bacteriosponges,
with
a
substan-
tial
biomass
contribution
from
unicellular
bacteria
and
cyanobacteria
in
the
mesohyl
or
inside
vesicles
of
special
cells
(bacteriocytes).
Megascleres
are
styles
with
a
few
oxeas
or
oxeote
modifications.
Microscleres
are
absent.
Reproduction
viviparous;
large
larvae,
up
to
6
mm,
elongate
ovoidal
and
uniformly
flagellated.
Distribution.
Central
West
Atlantic
and
Indone-
sia.
Etymology.
After
Sven
Zea,
first
collector
of
the
type
species,
in
recognition
of
his
contribution
to
the
systematics
of
Caribbean
sponges.
Svenzea
zeai
(Alvarez,
Van
Soest
&
Rutzler,
1998)
Hymeniacidon
amphilecta.
-
Pulitzer-Finali,
1986:
1
17
(not
de
Laubcnfels,
1936:
137)
Calyx
podatypa.
~
Humann,
1992:
33
Pseudaxinella
sp.
-
Alvarez
&
Crisp,
1994:
119
Pseudaxinella
zeai.
-
Erhardt
&
Moosleitner,
1995:
52
[prema-
turely
referring
to
Alvarez
et
al
1998]
Pseudaxinella
(?)
zeai
Alvarez,
Van
Soest
&
Rutzler,
1998:
20.
-
Diaz
&
Ward,
1997:
97-107
[production
rates
of
dissolved
inorganic
nitrogen].
Material
examined.
-
Holotype
USNM
39361
(Colombia,
Isla
Tcsoro,
18
m,
S.
Zea
coll.)
NTM
Z4152
(CBC97-BAG7),
Came
Bow
Cay,
Belize,
16°48'N,
88°05'W,
20
m,
M.C.
Diaz,
6
Aug
1997.
Description.
-
The
material
examined
from
Belize
fits
with
the
original
description
of
the
species.
New
observations
allow
us
to
shed
some
light
on
the
nature
of
the
granulous
cells
which
characteristi-
cally
obscure
the
skeleton
of
S.
zeai,
and
which
were
described
as
pigment
(?)
grains
by
Alvarez
et
al.
(1998).
Prominent
inclusion
cells
are
of
two
kinds,
bacteriocytes
and
granular
cells
(Rutzler
et
al.,
in
press)
(Fig.
2).
Both
styles
[length
205-270
(237.88
±
16.8);
width
7-12
pm
(9.48
±
1.5)]
and
oxeas
[length
210-
320
pm
(276.4
±21.3);
width
7-12
pm
(9.36
±
1.5)]
in
the
specimen
from
Belize
were
of
similar
dimen-
sions
to
those
present
in
the
material
from
St.
Croix,
Colombia
and
Tobago
(Alvarez
et
ah,
1998).
Reproduction.
-
The
species
is
viviparous,
with
un-
usually
large
and
conspicuous
embryos
present
in
most
specimens
year
around.
Embryos
are
yellow-
Photomicrograph
of
characteristic
granular
cells
found
in
Fig.
2.
circular
bodies
are
spicule
cross-sections.
(Scale:
10
pm.)
Svenzea
zeai;
Fig
I.
Svenzea
zeai
(Alvarez,
Van
Soest
&
Rutzler,
1998)
in
situ,
Santa
Marta, Colombia
(photo,
S.
Zea),
Scale
bar
20
mm.

Contributions
to
Zoology,
71(4)
-
2002
173
ish
to
cream,
spherical,
up
to
4
mm
in
diameter.
Larvae
are
ovoid
with
a
thicker
anterior
pole,
en-
tirely
and
evenly
flagellated,
up
to
6
mm
long.
Ecology.
-
Cyanobacterial
and
bacterial
endosym-
bionts
in
this
species
have
been
linked
to
the
fixa-
tion
of
dissolved
inorganic
nitrogen
(Diaz
&
Ward,
1997).
Distribution.
-
The
species
is
widely
distributed
through
the
Caribbean
area.
It
is
been
recorded
from
Florida
to
Puerto
Rico,
the
Virgin
Islands,
Tobago,
the
Atlantic
coast
of
Colombia,
Curasao
and
Belize,
between
20
and
40
m
depth.
Remarks.
-
Svenzea
zeai
was
wrongly
identified
as
Calyx
podatypa
by
Humann
(1992)
in
a
field
guide
of
reef
organisms.
After
this
publication
a
variety
of
novel
fatty
acids,
cyclopropene
sterols,
calyxamines
(a
type
of
alkaloid)
and
calyxolanes
(new
1,3-diphenylbutanoid
metabolites)
have
been
isolated
from
Caribbean
sponges
also
identified
as
Calyx
podatypa
(Carballeira
et
ah,
1998;
Rodriguez
etal.,
1997a;
Rodriguez
etak,
1997b).
We
assume,
that
the
target
species
of
these
chemical
investiga-
tions
was
Svenzea
zeai
given
that
the
species
is
wide-
spread
and
common
throughout
the
Caribbean
and
because
the
description
of
the
external
morphol-
ogy
given
in
these
papers
matches
perfectly
with
the
species.
Calyx
podatypa
(De
Laubenfels,
1934:
23)
has
no
resemblance
to
Svenzea
zeai.
It
is
mas-
sive
or
ramose,
brittle
in
consistency,
pale
drab
in
colour
with
characteristic
oscules
of
various
shapes:
heart-shaped,
Y-shaped,
and
some
resembling
hu-
man
footprints.
The
ectosomal
skeleton
is
thick
and
dense,
formed
by
a
tangential
layer
of
spicules.
The
choanosomal
skeleton
is
a
subrectangular
reticula-
tion
of
spongin
fibres
cored
by
multispicular
tracts
and
interstitial
spicules
in
a
vague
isodictyal
re-
ticulation.
The
spicules
are
exclusively
oxeas.
Svenzea
cristinae
sp.
nov.
(Figs.
3-4,
Table
1).
Material
examined.
-
Holotype:
USNM
1006559
(CBC97-52),
Cat
Cay
lagoon.
Pelican
Cays,
Belize,
I6”39.8'N,
88“
11.5'W,
Sven
Zea
coll.
3
Aug
1997.
Paratypes:
USNM
1006560
(KS96-
92),
Fisherman’s
Cay,
Pelican
Cays,
Belize,
I6"40.25'N,
88“
11.40'W,
K.
Smith
coll.
24
Nov
1996;
ZMA
17098
(CBC97-
57),
Cat
Cay
lagoon,
Pelican
Cays,
Belize,
16"39.8'N,
88°
11,5'W,
B,Alvarez
coll.
3
Aug
1997;
NTM
Z4155
(CBC97-58),
Cat
Cay
lagoon,
Pelican
Cays,
16°39,8'N,
88“
11,5'W,
Belize,
B.
Alvarez
coll.
3
Aug
1997.
Other
material:
NTM
Z4153
(CBC97-BAG3),
Blue
Ground
Range,
Belize,
I6°48.
6'N,
88°08.9'W,
on
man-
grove
roots,
B.Alvarez
coll.,
31
Jul
1997;NTMZ4154(CBC97-
BAG4),
Blue
Ground
Range,
Belize,
16°48.6'N,
88°08.9'W,
on
mangrove
roots,
B.Alvarez
coll.,
31
Jul
1997.
ZMA
16613,
Discovery
Bay,
Jamaica,
H.
Lehnert
coll.
Description.
-
Thickly
encrusting.
Surface
brown-
yellowish
with
purple
or
brown-pinkish
areas;
smooth;
pierced
with
minute
pores
less
than
1mm
diameter,
aggregated
in
shallow
depressions;
with
Fig.
3.
Svenzea
cristinae
n.
sp. A,
holotype
USNM
1006559
(CBC97-52).
B,
ZMA
16613
in
situ,
Discovery
Bay,
Jamaica
(photo,
H.
Lehnert).
Scales
A-B
10
mm.
cristinae,
NTM
Z4154
(CBC97-BAG4).
A,
scan-
ning
electron
micrograph
(SEM)
of
the
choanosomal
skeleton.
B,
spicules,
(scales
B,
100
pm,
C,
25)
Fig.
4.
Svenzea

B.
Alvarez
et
al.
-
Svenzea,
new
sponge genus
from
tropical
reefs
174
meandering
and
transparent
channels.
Oscula
cir-
cular
and
flush
with
surface.
Species
Styles
Oxeas
Svenzea
zeai
(Alvarez,
Van
Soest,
Riitzler)
Colombia
(Hoiotype)
length
220-290
(270.4±17.7)
270-330
(301.6±15.5)
width
7.5-12.5
(11.1±1.5)
7.5-12.5
(10.4±l.2)
Belize
(CRC97-BAG7)
length
205-270(237.9±l6.8)
210-320(276.4±21.3)
width
7-12(9.5±1.5)
7-12(9.4±l.)
Svenzea
cristinae
n.
sp
Belize
(NTM
Z4155)
length
320-450(405.6±27.9)
60-200(131.6±42.3)
width
6.25-17,5(11.05±27)
3.75-10(6.17±2.1)
Jamaica
(ZMA
16613)
length
310-460(375.8±45.3)
120-340(212.5±102.4)[4]
width
4.25-11.25(7.97±1.5)
6.25-8.25(7.15±0.9)[5]
Svenzea
devoogdae
n,
sp
Sulawesi
(ZMA
17097)
length
160-270(218.4±25.1)
200-290(243.2±23.7)[14]
width
6.25-11.25(9.15±I.4)
7.5-12.5(9.7±1.4)[14]
Bali
(RMNH
Por.1363)
length
130-270(221
±38.8)
130-240(182.5±47.9)[4]
width
8.25-13.75(11.2±1.4)
7,5-12.5(10±2,0)[4]
Choanosome
white.
Consistency
soft,
crumbly.
Ectosomal
skeleton
specialization
absent.
Choano-
somal
skeleton
(Fig.
4A)
is
a
disorganised
uni-
spicular
reticulation
with
loose
spicules
forming
triangular
or
polygonal
like-meshes
with
very
little
spongin
at
the
nodes;
granular
cells
obscuring
most
of
the
skeleton.
Spicules
are
(Fig.
4B-C,
Table
1):
styles,
straight
or
slightly
sinuous
(320-460
x
4-11
pm)
often
with
mammiform
tips
or
with
swellirlgs;
oxeas
(60-340
x
4-10
pm),
less
frequent
than
styles
or
absent;
may
occur
in
two
size
categories.
Smaller
oxeas
straight,
curved,
angulated
or
twisted.
Large
numbers
of
endosymbiotic
bacteria
present
through-
out
the
sponge.
Etymology.
-
The
species
is
named
after
Maria
Cristina
Diaz
in
recognition
to
her
contribution
to
the
knowledge
of
the
biology
of
the
genus.
Distribution.
-
Belize,
Jamaica;
subtidal
on
man-
groove
roots,
0.5
m.
Remarks.
-
The
species
differs
from
Svenzea
zeal
mainly
in
shape
and
consistency.
It
has
all
the
diag-
nostic
characters
of
the
genus,
including
the
pres-
ence
of
granular
cells
and
skeletal
organisation.
Other
characters
common
to
dictyonellids
such
as
the
dominance
of
styles
and
the
spicules
with
vari-
able
endings
are
also
present
in
this
species.
Svenzea
devoogdae
sp.
nov.
(Figs.
5-6,
Table
1).
Material
examined.
-
Hoiotype,
ZMA
17097,
Indonesia,
SW
Sulawesi,
Spermonde
Shelf,
Bone
Lola,
05°03'145"S
119°21'207"E,
11
m,
Nicole
de
Voogd,
#NV/BL/080900/l29,
8
Sep
2000.Other
material:
RMNH
Por.1353,
Indonesia,
Bali,
N
side
Nusa
Penida,
off
Desa
Ped,
08“40'28"
S
115°30'50"
E,
24
m,
coll.
Nicole
de
Voogd,
NV/Bal.26/180401/217,
18
April
2001.
Description.
-
Massive
with
lobes
and
oscular
mounds.
Maroon-black
externally,
beige
choano-
some.
Surface
smooth,
infested
with
small
black
zoanthids.
Oscula
up
to
1
cm
in
diameter
on
top
of
lobes
and
mounds.
Ectosomal
skeleton
specializa-
tion
absent.
Choanosomal
skeleton
is
a
disorganised
reticu-
lation
of
uni-paucispicular
spicule
tracts
connected
at
the
nodes
with
very
little
spongin
and
obscured
by
granular
cells
(Fig.
6A).
Spicules
are
styles
and
less
frequent
oxeas
(Table
1,
Fig.
6B).
Styles
are
spp.
Measurements
(in
pm)
are
ranges
of
25
spicules
or
the
number
indicated
in
brackets,
with
means
±
standard
deviation
in
parenthesis.
Table
I.
Spicule
dimensions
for
Svenzea
Species
Styles
Oxeas
Svenzea
zeai
(Alvarez,
Van
Soest,
Riitzler)
Colombia
(Ilolotype)
length
220-290
(270,4±17.7)
270-330
(301.6±15.5)
width
7.5-12.5
(1
l.l±1.5)
7.5-12.5
(10.4±
1.2)
Belize
(CRC97-BAG7)
length
205-270(237.9±l6.8)
210-320(276.4±21.3)
width
7-12(9.5±1.5)
7-12(9.4±1.)
Svenzea
cristinae
n.
sp
Belize
(NTM
Z4155)
length
320-450(405.6±27.9)
60-200(131.6±42.3)
width
6.25-17.5(11.05±27)
3.75-10(6,17±2.1)
Jamaica
(ZMA
16613)
length
310-460(375.8±45.3)
120-340(212.5±102.4)[4]
width
4.25-11.25(7.97±1.5)
6.25-8.25(7.15±0.9)[5]
Svenzea
devoogdae
n.
sp
Sulawesi
(ZMA
17097)
length
160-270(218.4±25.1)
200-290(243.2±23.7)[14]
width
6.25-11.25(9.15±1.4)
7.5-12.5(9.7±1.4)[14]
Bali
(RMNH
For.
1363)
length
130-270(221±38.8)
130-240(
182.5±47.9)[4]
width
8.25-13.75(11.2±1.4)
7.5-12.5(10±2,0)[4]

Contributions
to
Zoology,
71(4)
-
2002
175
flexuous
or
straight
some
with
bases
slightly
bent
or
reflexed
(130-270
x
6-14
pm).
Oxeas
are
straight
or
angulated,
some
with
hastate
points
(130-290
x
8-13
pm).
Sustantial
numbers
of
endosymbiotic
bacteria
present.
Distribution.
-
Sulawesi,
Bali,
Indonesia;
11-20
m
Etymology.
-
The
species
is
named
after
Nicole
de
Voogd,
(University
of
Amsterdam
/
National
Mu-
seum
Naturalis,
Leiden)
who collected
the
speci-
mens
examined
here.
Remarks.
-
Both
megascleres
appear
in
a
wide
size
range
(160-250
pm
length)
in
the
specimen
from
Bali;
the
smaller
generally
thicker than
the
larger
ones.
This
feature
was
not
so
obvious
in
the
speci-
men
from
Sulawesi
and
suggests
that
intraspecific
variation
of
spicule
size
might
be
high.
The
spe-
cies
is
remarkably
similar
to
Svenzea
zeai
differ-
ing
only
in
that
spicules
can
be
sinuous,
rhabdose
(styles)
or
angulated
(oxeas).
As
in
the
Caribbean
species,
S.
devoogdae
is
quite
common
within
its
known
geographical
range
but
with
a
patchy
dis-
tribution
(N.
de
Voogd
personal
communication)
Discussion
A
possible
fourth
species
of
the
new
genus
is
Pseu-
daxinella?
flava
Lehnert
&
Van
Soest,
1999
from
76
m
on
the
fore
reef
off
N
Jamaica.
It
shares
fea-
tures
with
above
described
species,
especially
with
S.
zeai
such
as a
reticulation
of
styles
of
244-380
x
2-11
pm
and
lack
of
ectosomal
specialization.
However,
the
reticulation
in
this
species
is
aniso-
tropic,
with
multispicular
ascending
tracts,
and
more
importantly,
the
characteristic
granular
cells
and
large
embryos/larvae
were
not
observed,
preclud-
ing
a
definite
assignment
to
Svenzea.
The
higher
taxonomic
position
of
Svenzea
re-
mains
unclear.
Based
merely
on
skeleton
and
spi-
cules
an
unambiguous
assignment
to
a
family/or-
der
is
not
possible.
In
these
features,
the
genus
shows
affinities
with
members
of
both
Halichondrida
and
Haplosclerida.
We
assign
the
genus
to
Halichon-
drida;
Dictyonellidae
for
it
shares
microanatomical
and
developmental
features
with
members
of
the
dictyonellid
genus
Scopalina
Schmidt,
particularly
the
granulous
cells,
certain
spicule
features,
and
larval
development,
morphology,
and
aspects
of
behaviour
(Riitzler
et
ah,
in
press,
Uriz
1982).
On-
going
research
using
molecular
data
is
expected
to
find
independent
evidence
of
the
phylogenetic
re-
lationships
for
Svenzea
and
to
help
define
its
posi-
tion
in
the
higher
classification.
Recent
results
of
that
research
using
28S
rRNA
(McCormack
et
al,
in
press)
indicates
already
that
Svenzea
zeai
is
not
closely
related
to
haplosclerid
species.
Acknowledgements
We
are
grateful
to
Nicole
de
Voogd,
who
kindly
collected
and
provided
us
specimens,
photographs
and
information
of
the
Fig.
5.
Svenzea
devoogdae
n.
sp.
A-B,
holotype
ZMA
17097
(Sulawesi)
in
situ,
Sulawesi,
Indonesia.
Scales A, 50
mm,
B,
25
mm.
Fig.
6.
Svenzea
clevoogdae
n.
sp. A,
SEM
of
the
choanosomal
skeleton.
B,
spicules
(scales
100
pm)

176
B.
Alvarez
et
al.
-
Svenzea,
new
sponge
genus
from
tropical
reefs
Indonesian
species.
We
thank
Maria
Cristina
Diaz,
Sven
Zea,
and
Kate
Smith
for
their
valuable
comments
and
discussions
on
the
new
genus
and
forthe
collection
of
some
of
the
speci-
mens
included
in
this
work.
Helmut
Lehnert
kindly
donated
his
Jamaican
material
to
ZMA.
The
collection
of
most
of
the
speci-
mens
included
in
this
study
was
supported
by
the
Smithsonian
Institution’s
Caribbean
Coral
Reef
Ecosystems
Program
(Con-
tribution
no.
643)
J,
van
Arkel,
University
of
Amsterdam,
provided
graphic
support,
Rob
van
Soest
acknowledges
support
from
the
EC
MASS
CT97-0144
grant
‘SYMBIOSPONGE’,
which
allowed
Raquel
Gomez
and
Mario
J. De
Kluijver
to
collect
the
Curasao
material.
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Received:
7
August
2002