Invertebrate Biology 124(4): 332–343.

r 2005 American Microscopical Society, Inc.

DOI: 10.1111/j.1744-7410.2005.00031.x

Fine structure of the epistome in Phoronis ovalis:

significance for the coelomic organization in Phoronida

Alexander Gruhl,a Peter Grobe, and Thomas Bartolomaeus

Freie Universität Berlin, Evolution und Systematik der Tiere, 14195 Berlin, Germany

Abstract. The hypothesis of a common ancestry of the lophophorate taxa Brachiopoda,

Bryozoa, Phoronida, and the Deuterostomia can be traced back to the late 19th century when
Masterman recognized a tripartite organization of the body consisting of pro-, meso-, and
metasome, along with coelomic body cavities in each compartment, as characteristic for
Echinodermata, Pterobranchia, Phoronida, and Brachiopoda. This idea became quite pop-
ular under the name ‘‘archicoelomate’’ concept. The organization of the phoronids, and es-
pecially of their transparent actinotroch larva, has for a long time been used as a touchstone
for the validity of this concept. As a coelomic lining can reliably be recognized only on the
ultrastructural level, this technique has been applied for adults of Phoronis ovalis, which is
assumed to be a sister species to all other phoronids. Phoronis ovalis contains only two co-
elomic compartments, a posterior coelom inside the trunk (metasoma), occupying the space
between the trunk epidermis and the digestive epithelium, and an anterior lophophoral co-
elom inside and basal to the tentacular crown (mesosoma). There is no coelomic cavity inside
the epistome (prosoma). This part of the body is filled with myoepithelial cells, which are
continuous with the epithelial lining of the lophophore cavity. These cells form a lumenless
bilayer and possess long, tiny myofilamentous processes, which are completely embedded in
an extracellular matrix. A comparison with data on P. muelleri shows that there is no need
to assume three different coelomic cavities in Phoronida, in contrast to the predictions of
the archicoelomate concept. At least for this taxon, a correspondence to the situation in
deuterostomes can hardly be found.
Additional key words: ultrastructure, coelom, phylogeny

In the majority of recent textbooks on zoology and
developmental biology, members of the Phoronida
are considered to have a tripartite body, each part
equipped with its own secondary body cavity (e.g.,
Zimmer 1997; Nielsen 2001; Ruppert et al. 2004). The
same is assumed for Bryozoa and Brachiopoda,
which are traditionally grouped with Phoronida in
the taxon Lophophorata (Hyman 1959). Among oth-
er characters, trimeric coelomic organization is re-
garded as apomorphic for a taxon Radialia, uniting
lophophorates with deuterostomes (Jefferies 1986;
Ax 1989). Recent ultrastructural and developmental
studies, however, have failed to show such a distri-
bution of body cavities (Lüter 2000; Bartolomaeus
2001). Furthermore, molecular data argue for posi-
tioning the lophophorate taxa together with spiral-
a
Author for correspondence.
E-mail: agruhl@zoosyst-berlin.de
ians in the Lophotrochozoa (e.g., Halanych et al.
1995; Peterson & Eernisse 2001).
The idea of a tripartite body was influenced by the
so-called archicoelomate concept, introduced by
Masterman (1898), and revived and extended by
Remane (1950), Ulrich (1950, 1972), Siewing
(1973b, 1980), and Herrmann (1997) in the 20th cen-
tury. These authors emphasized tripartition as a pri-
mary feature in coelomate organization and argued
that such a condition could have already been found
in the bilaterian ancestor. They proposed that the
hypothetical bilterian ancestor was the end-point of a
transition from diploblastic medusae to triploblastic,
coelomatous Bilateria, a transition during which
gastrovascular pouches like those found in certain
cnidarian species became coelomic cavities by enter-
ocoely. Despite mixing different levels of ontogenesis
with assumptions on the course of evolution, this
idea is hardly in accordance with recent data sup-
porting the acoelomate organization of earliest bila-
terians (Rieger 1986; Ax 1996; Rieger & Purschke
Phoronis ovalis epistome fine structure
2005). But even within the framework of the arch-
icoelomate theory, the most controversial aspect is
whether the first body section of lophophorates, the
epistome, actually contains a coelom as predicted by
the theory; an epistome coelom is not univocally de-
scribed in phoronids (Caldwell [1882] versus Master-
man [1896], de Selys-Longchamps [1907] versus
Siewing [1974] and Herrmann [1980] for Phoronis
muelleri, Cori [1937] for Phoronida in general,
Zimmer [1964] for P. vancouverensis and Emig &
Siewing [1975] for P. psammophila). The reported
existence of a coelom became a touchstone for the
validity of the archicoelomate concept (Siewing 1980;
Herrmann 1997). All these studies were undertaken
at the light microscopic level. Clear discrimination
between different body cavities, however, generally
requires electron microscopic techniques, as the co-
elomic cavity is lined by an epithelium, often only
B50 nm thick, whereas a primary body cavity is lined
by an extracellular matrix (ECM).
The proposed tripartition interpreted in many light
microscopic studies has not been corroborated by in-
vestigations at the ultrastructural level. In the species
of Brachiopoda (Lüter 1998, 2000) examined to
date, no independent protocoel was found. In the
phylactolaemate Bryozoa, the open connection be-
tween epistomal cavity and lophophoral coelom
(see Mukai et al. 1997) has been confirmed (T.
Bartolomaeus, unpubl. data). Bartolomaeus (2001)
demonstrated that in P. muelleri in the preoral lobe of
the larva, as well as in the epistome of the adult, there
is no such cavity and a voluminous ECM can be
found here instead.
To elucidate whether this character is autapomor-
phic for P. muelleri or can be assumed for the ground
pattern of Phoronida, methodically similar investiga-
tions have to be undertaken for additional phoronid
species. The most recent phylogenetic analyses of
relationships within Phoronida argue either for
P. ovalis WRIGHT 1856 being the adelphotaxon of
all other phoronid species (Bartolomaeus & Grobe
2001), or at least for P. ovalis and P. muelleri to be
representatives of the highest-level sister taxa within
Phoronida (Emig 1985). In both cases, this would
mean that similar conditions in both species are most
parsimoniously interpreted as a ground-pattern char-
acter of the Phoronida.
Methods
Specimens
Adults of Phoronic ovalis were collected during
October 2001 in the Gullmar Fjord, West coast of
Sweden. Preparation and fixation were accomplished
333
at Kristineberg Marine Biological Station. Shells
from dead molluscs were dredged from 35 m depth
at 58113.000 N; 11124.410 E. After keeping them a few
hours under cold running seawater, individuals of P.
ovalis displayed their tentacle-crowns. The specimens
were removed by carefully breaking the shells.
Specimens were fixed in 2.5% glutaraldehyde buff-
ered in 0.1 M sodium cacodylate at 41C for 60 min.
Ruthenium red was added to the fixative to stain cer-
tain components of the ECM. After fixation, the
specimens were rinsed in the same buffer and stored
therein until they were postfixed in 1% OsO4 buff-
ered in 0.1 M sodium cacodylate at 41C for 45 min.
For transmission electron microscopy (TEM),
specimens were dehydrated in an acetone series and
embedded in araldite via propylene oxide. Serial sec-
tions of B60 nm thickness were made using diamond
knives on a Reichert ultramicrotome (Leica Micro-
systems, Wetzlar, Germany). Sections were collected
on copper, formvar-covered single-slot grids, and au-
tomatically stained with uranyl acetate and lead cit-
rate in a Leica EM Stain processor. They were
examined on a Philips CM 100 TEM (Phillips Elec-
tron Optics, Eindhoven, The Netherlands).
For scanning electron microscopy (SEM), speci-
mens were dehydrated in an alcohol series, automat-
ically critical-point dried with a Bal-Tec CPD 030,
coated with gold in a Bal-Tec SC 005 sputter coater,
and examined in a Hitachi S 450 SEM (Hitachi
Instruments, San Jose, CA, USA).
Terminology
Because of the complicated development, the no-
menclature of body axes is confusing. Many anatom-
ic features are denoted variably by different authors.
In this study, the nomenclature summarized by Emig
(1975) is used. However, the terms ‘‘trunk’’ and ‘‘lop-
hophore’’ were chosen instead of Emig’s ‘‘meta-
some’’ and ‘‘mesosome,’’ and ‘‘trunk coelom’’ and
‘‘lophophore coelom’’ instead of ‘‘metacoel’’ and
‘‘mesocoel.’’ This is followed because the traditional
terms imply as yet unproven correspondence to the
coelomic cavities found in deuterostomes. The term
‘‘secondary body cavity’’ or ‘‘coelom’’ refers to a
body space lined completely with an epithelium, as
defined in Ruppert (1991) and Bartolomaeus (2001).
Results
The epistome is situated between the mouth and
the indented portion of the tentacle ring, which lines
the lophophoral concavity. This pillow-shaped organ
is situated at the base of the tentacle ring. Being
closely attached to it, the epistome has a bean-
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vol. 124, no. 4, fall 2005
334
Fig. 1. Phoronis ovalis. SEM. Tentacles are partly removed
to allow visualization of the epistome (ep). Mouth opening
(arrow) is covered by food and debris particles. loc,
lophophoral concavity.
like structure when viewed from above (Fig. 1). Its
convex side faces the mouth opening. The lateral ex-
tent of the epistome is 60–70 mm and it measures 20–
30 mm in length and in height. Its surface is ciliated
and noticeably spongy, and seems to consist of many
gland cells. The epistome is composed of peripheral
epidermal cells that rest on a basal lamina, which en-
closes mesodermal cells containing myofilaments.
Epidermis
At the ultrastructural level, the epidermis of the
epistome is composed of a very high epithelium
whose cells are B20 mm in height (Fig. 2A). Basally
scattered neural processes were found. The cells rest
on a 100-nm-thick basal lamina (Fig. 4). Three dis-
tinct areas of epistome epidermis can be differentiat-
ed, mainly because of cytological attributes. They are
situated on the abfrontal side, at the tip, and on the
frontal side of the epistome (Fig. 2A).
Most cells of the epistome’s abfrontal side are
slightly prismatic, with one cilium each. The elec-
tron-transparent cytoplasm contains a few organ-
elles, with the oval nuclei located basally. The
apical cell membrane extends into numerous 0.5-
mm-long microvilli, which bear a glycocalyx (Fig.
2B–D). Zonulae adhaerens, attached to a zone of ac-
tin filaments, occur in the apical regions of the cells.
The cells bear ciliary basal structures including a ver-
tical and a horizontal striated rootlet (Fig. 2C,D).
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vol. 124, no. 4, fall 2005
Gruhl, Grobe, & Bartolomaeus
The accessory centriole, indicating the direction of
the effective ciliary stroke, is located opposite to the
horizontal rootlet, connected by a basal foot both to
the basal body on one side and to the vertical rootlet
on the other side. The position of the centriole and,
accordingly, the direction of the horizontal rootlet
differ between the median and the lateral cells. The
accessory centrioles are oriented toward the tip of the
epistome in median cells, and are directed laterally in
peripherally situated cells. The axoneme has the usu-
al 9
212 pattern of microtubule arrangement.
The epidermal cells at the tip of the epistome are
higher and much narrower (Fig. 2A). They form a
boundary between the abfrontal and frontal sides.
The most apically situated cells are spindle shaped
and only a few of them are connected to the basal
lamina via a narrow process each. They bear deviant
cilia whose basal structures include three striated ver-
tical rootlets running basally along the core. An ac-
cessory centriole is located somewhere between two
of these rootlets (Fig. 2D). The cilium is surrounded
by eight enlarged microvilli, forming a collar and car-
rying a thickened glycocalyx. Basal to these cells, the
epithelial nerve plexus is more pronounced. Howev-
er, neural contacts to these cells could not be found.
In the frontal apical region, the cells are of elon-
gated shape. Because of a high concentration of free
ribosomes, the cytoplasm appears electron dense
(Fig. 2A). Furthermore, many specialized gland cells
reside between the other epidermal cells (Fig. 2B).
They are characterized by a unique fine structure of
their vesicles, which is granulous and electron opa-
que, and has oblong and loopy core structures. Their
cilia are composed of an axoneme and a basal body
lacking any rootlets. These gland cells clearly differ
from gland cells found in the epidermis of the trunk
and lophophore (Fig. 3A). Here, four types of gland
cells can be distinguished. All of them do not possess
cilia and their vesicles bear no resemblance to those
of the epistomal gland cells (Fig. 3C). The other ep-
idermal cells bear two orthogonally arranged striated
rootlets, the horizontal one pointing toward the tip of
the epistome, with the accessory centrioles oriented
toward the mouth opening (Fig. 2).
Basally in the epistome, slightly abfrontal to the
median axis, a concentration of neural processes is
located (Figs. 4A,B; 5A,B). They belong to a circular
nerve running interior to the bases of the tentacles
and passing the epistome at its abfrontal side.
Musculature
The muscular system of the epistome is underneath
the subepidermal basal lamina and composed of
Phoronis ovalis epistome fine structure 335

Fig. 2. Phoronis ovalis. TEM. Epistome. A. Sagittal overview showing the different components. epaf, abfrontal
epidermis; epf, frontal epidermis; mc, myoepithelial cells. B. Epistomal gland cell containing excretory vesicles (ve),
nucleus (nu), and an axoneme (an). Normal epidermal cells bear microvilli (mv). C. Ciliary basal structures (ba) of frontal
epidermal cell. The accessory centriole (ac) points toward the mouth opening. hr, horizontal rootlet. D. Ciliary basal
structures of epidermal cell at the epistome’s tip.

myoepithelial cells. In each cell, the myofilaments are
separated from the perinuclear part, which contains
the nucleus as well as the majority of the organelles
and cytoplasm. The myofibrils are inside a single long
tiny process, which extends laterally (Figs. 4A,C; 6).
These processes are embedded in a thickened lamina
fibroreticularis, which is continuous between the
frontal and abfrontal side of the epistome. The
cell somata are situated medially in the epistome
(Figs. 4B; 6). They are interconnected via zonulae

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336 Gruhl, Grobe, & Bartolomaeus

Fig. 3. Phoronis ovalis. TEM. A. Sagittal section of trunk epidermis. Normal epidermal cells (epc) contain apical secretory
vesicle (asv); specialized gland cells (glc1–4) bear different types of vesicles. Cells settle on an extracellular matrix (ecm).
B. Trunk musculature consists of myoepithelial cells (emc) whose filamentous processes (fp) are arranged either circularly
or longitudinally. They are connected to the ECM via hemidesmosomes (hd). C. Different types of vesicles in epidermal
gland cells. D. Diaphragm separating lophophoral and trunk coeloms. ECM layer (arrowheads) is surrounded by the
lining epithelia of the lophophore (llc) and the trunk coelom (ltc). epc, epidermal cell; gd, intestinal epithelial cell;
fp, filamentous processes of coelomic epithelial cells; glc, gland cell in the nerve ring.

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Phoronis ovalis epistome fine structure 337

Fig. 4. Phoronis ovalis. TEM. Epistome. A. Peripheral sagittal section showing myofilament-rich processes (fp) of the
inner epithelial cells completely embedded in an extracellular matrix (ECM) (arrowheads). B. Central sagittal section
showing perikarya of the inner epithelial cells, which are continuous with the lining of the lophophore coelom as indicated
by apical adherens junctions (arrow). C. Cross section. Intraepistomal cells bear myofilamentous processes that extend
laterally into the ECM. epaf, abfrontal epidermis of the epistome; epf, frontal epidermis of the epistome; fp, fibrous
processes of coelomic epithelial cells; lcoe, lophophore coelom; llc, lining of trunk coelom; ne, neural processes.

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vol. 124, no. 4, fall 2005
338 Gruhl, Grobe, & Bartolomaeus

Fig. 5. Phoronis ovalis. Schematic

illustration of sagittal sections of
the lophophore region with mouth
opening (mo) and epistome (ep).
Rectangles indicate the position of
TEM micrographs in Figs. 3 and 4.
A. Peripheral sagittal section. Pro-
cesses of coelomic epithelial cells
containing myofilaments invade
the subepidermal ECM (black) of
the epistome. B. Central sagittal
section. Coelomic epithelium
forms a continuous lining inside
of the lophophore coelom and the
epistome. epi, epidermis; gd,
gastrodermis; lcoe, lophophore
coelom; rn, ring nerve; tcoe,
trunk coelom.

adhaerens and are continuous with the myoepithelial
lining of the lophophoral coelom. Basally they ad-
here to the matrix by dense plaques and hemidesmo-
somes.
Coelom
The lining epithelia of trunk and lophophore co-
eloms are exclusively composed of myoepithelial cells
that are interconnected via apical adherens junctions
(Fig. 3A,B). They form either a distinct circular mus-
cle layer or single transverse muscle strands crossing
the coelomic space. The lophophoral coelom fills the
lophophore and extends into the tentacles (Fig. 5).
Muscle cells with longitudinally arranged myofila-
ments predominate in the tentacles (Fig. 7). The lu-
men of the lophophoral coelom is nearly completely
filled with amoeboid coelomocytes that also occur in

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Phoronis ovalis epistome fine structure
the processes extending into the tentacles. The co-
elomic epithelial cells can be distinguished from the
coelomocytes because of apical adherens junctions
that show their epithelial character.
The diaphragm separates lophophoral and trunk
coeloms (Figs. 3D; 5). It consists of a continuous
339
Fig. 6. Phoronis ovalis. Recon-
struction of the epistome, longi-
tudinal section. The epistome
is covered by a monociliated
epidermis (epi) (cilia not shown).
The epithelium of the lophophore
coelom (llc) is continuous with
the epistome’s myoepithelial cells.
Their basal processes (fp) contain
myofibrils and embedded within
voluminous subepidermal matrix
(ecm) of the epistome. lcoe,
lophophore coelom; mv, microv-
illi; za, zonula adhaerens.
central ECM layer surrounded by the myoepithelial
cells of both lophophoral and trunk coeloms. The
trunk coelom occupies the space between the diges-
tive tract and the trunk epidermis. Its coelomic epi-
thelium forms the pronounced trunk musculature
(Fig. 3B). Two types of muscle cells can be observed.
Fig. 7. Phoronis ovalis. TEM. Cross-
section of a tentacle. Coelom in the
tentacle is lined by myoepithelial
cells (emc) and filled with co-
elomocytes (cc). The coelom
encloses a blood vessel (bv). af, ab-
frontal side; f, frontal side; gc,
gland cell; ne, nerve process.
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340
The annular muscle cells reside under the subepider-
mal ECM and possess processes with circularly ar-
ranged myofilaments. These processes are embedded
in the ECM and so the musculature appears pseudo-
stratified (Fig. 3B). The longitudinal muscle cells are,
with their somata, integrated into the epithelium.
Their myofilamentous processes, which are much
thicker than those of the circular muscle cells, extend
into the coelomic lumen.
Discussion
Epidermis
The epistomal epidermis mainly consists of ciliated
cells. The distribution of cilia and the directions of
their rootlets, indicating the direction of the effective
stroke, gives evidence for the course of the created
water flow at the epistome. The water current at the
middle of the abfrontal side is directed upward, to-
ward the tip, whereas laterally it is directed around
the epistome, toward the frontal side. On the frontal
side, the cilia beat in the direction of the mouth open-
ing. The cells at the tip of the epistome show no clear
distribution of their accessory centrioles. However,
their fine structure shows some resemblance to pre-
viously described laterofrontal cells of the tentacles,
which are presumably sensory.
A unique type of gland cells was found in the fron-
tal epidermis of the epistome. Excretion of their ve-
sicular material was not observed, but it is likely that
it consists of mucous or tryptic substances, which are
added to the passing food current. Whereas the gland
cells of the trunk and lophophoral epidermis find
overall resemblance in previous ultrastructural ob-
servations on the epidermis in other phoronid species
(Pourreau 1979; Fernandez et al. 1991 for Phoronis
psammophila; Aguirre et al. 1993 for P. australis,
P. hippocrepia; Temereva et al. 2001 for Phoronopsis
harmeri), the epistomal gland cells have not been
found in any of these species.
Gilmour (1978) considers the function of the epis-
tome as to direct the incoming water current and to
reject inappropriate particles. Regarding the ciliation
and the distribution of gland and presumptive sen-
sory cells, it can be assumed that, at least in P. ovalis,
a second role of the epistome may be to add sub-
stances to the food current.
Musculature
The epistome is filled with a narrow process of the
coelomic epithelium of the lophophoral cavity. The
cells bear myofilamentous processes extending later-
Invertebrate Biology
vol. 124, no. 4, fall 2005
Gruhl, Grobe, & Bartolomaeus
ally. Thus, the epistome is able to accomplish con-
strictions along its lateral axis, meaning it can
become narrower. A constriction along its longitudi-
nal axis is not possible because of the absence of lon-
gitudinally arranged myofilaments. Movements of
the epistome itself can only be mediated by the mus-
culature of the lophophoral region. This has been
described in earlier studies, and was not the subject of
this investigation.
Coelom
In P. ovalis, the myoepithelial cells of the lophoph-
ore’s coelomic lining form a bilayer that extends into
the epistome like a tongue. There is no fluid filling
between the cells as in the lophophoral coelom. None-
theless, the cells are clearly epithelial, as they show a
strong polarity with apical adhaerens junctions and
basal hemidesmosomes. Cells facing each other are
never connected by any junctions. Such an arrange-
ment can potentially be expanded by fluid accumula-
tion. If this should actually occur, the resulting fluid-
filled cavity would be confluent with the lophophoral
coelom. As a septum is lacking, in neither case could
independent coelomic cavities occur in the epistome
and the lophophore–tentacle complex.
Most studies dealing with the epistomal cavity in
Phoronida do not provide strong evidence for a co-
elomic nature because they lack clear documentation
of an epithelial lining. Except for the study by
Siewing (1974), all were undertaken at the light mi-
croscopic level: Pross (1974) provides pictures of
muscle strands within the epistome but it is not clear
whether they are part of a coelomic lining; this is also
the case in Siewing (1973a). Siewing (1974) and Herr-
mann (1980, 1986) give morphological descriptions
of ontogenetic and metamorphic stages in P. mu-
elleri. However, it is not visibly clear whether the
‘‘septum’’ they show in their histological micro-
graphs (Herrmann 1980) is a real septum, which con-
sists of two coelomic epithelia resting base to base on
a common matrix, or if it is the lining of the lop-
hophoral coelom resting on a matrix that lines a vo-
luminous primary body cavity. The only TEM
micrographs Siewing (1974: figs. 6, 7) provides do
not show an epithelium. Zimmer (1978, 1980) give
detailed descriptions and documentation on the oc-
currence of mesoderm in the actinotroch preoral
hood, and regards it as homologous with the adult
epistome. Ultrastructural evidence of the coelomic
nature of these structures is absent in these studies.
Moreover, the strong confidence Siewing (1973b,
1980) and others laid in the archicoelomate theory
seemingly led to the misinterpretation of parts of the
Phoronis ovalis epistome fine structure
lophophoral coelom as an epistomal coelom. Despite
the descriptions they gave, Siewing (1974: fig. 21) and
Emig & Siewing (1975: fig. 6) showed the ganglion at
the anal face of the cavity they interpreted as an epi-
stomal coelom. The ganglion, however, is always ad-
jacent to the lophophoral coelom in phoronids.
Furthermore, the authors said that the epistomal co-
elom borders on the trunk coelom, which never oc-
curs in phoronids because the lophophoral coelom
acts as a hydroskeleton for the tentacles, so that it lies
orally to the trunk coelom. A further argument con-
cerns the position of the big vascular ring vessel,
which, according to Siewing (1974) and others, is sit-
uated in the epistomal coelom. This is clearly a mis-
interpretation, because several authors (Hyman
1959, Temereva & Malakhov 2003, this study) show
the ring vessel to be located at the bases of the ten-
tacles in the lophophoral coelom.
The present study shows that the epistome of
P. ovalis internally contains myoepithelial cells that
belong to the lining of the lophophoral coelom. This
result is in accordance with the observations made in
P. muelleri (Bartolomaeus 2001). An independent
epistomal coelom is lacking here, too, and muscle
cells are found inside the epistome. An ECM can be
found interspersed between them. Thus, even if there
should be a cavity within the epistome in any repre-
sentative phoronid, as implied in previous studies,
this could be nothing else than either fluid-filled
interstices within the matrix, i.e., a primary body
cavity, or a part of the lophophoral coelom.
Our study clearly shows that there is no evidence
for an epistomal coelom in P. ovalis. As this species is
sister taxon to the remaining phoronid species, this
result corroborates the assumption that a protocoel
or epistome coelom is primarily lacking in Phoronida
(Bartolomaeus 2001). According to Pross (1980), Sie-
wing (1980), and Zimmer (1997), brachiopods pos-
sess an unpaired protocoel as well as paired meso-
and metacoels, in accordance with the archicoelo-
mate theory. As in the Phoronida, electron micros-
copy changed this interpretation; Lüter (1998, 2000)
was able to refute the independence of the epistomal
coelom. In fact, there is continuity with the secondary
body cavities of the lophophore and median tentacle.
Lüter (1996) furthermore postulated a single co-
elomic anlage, which is incompletely divided later
during ontogenesis, for the ground pattern of Bra-
chiopoda. In Bryozoa, finally, only the limnetic phy-
lactolaemate species show a tripartite organization
with trunk, tentacle region, and epistome (Hyman
1959; Mukai et al. 1997; Zimmer 1997). Recent ul-
trastructural data show that their epistome contains a
coelomic cavity. Despite the predictions made by the
341
archicoelomate theory, however, this cavity is not re-
stricted to the epistome, but confluent with the trunk
coelom (T. Bartolomaeus, unpubl. data). Although
developmental studies have to confirm this, it seems
likely that the epistome has secondarily been invaded
by a protrusion of the trunk coelom. Within this
framework of knowledge on coelomic cavities in lop-
hophorate taxa, these findings have far-reaching con-
sequences.
A system of coelomic cavities consisting of an an-
terior protocoel, a pair of mesocoel cavities forming
the hydroskeleton of the tentacle apparatus, and a
pair of metacoel cavities has also been described for
hemichordate and echinoderm species. The tradition-
al assumption that such a trimeric coelomic organi-
zation supports the hypothesis of a common ancestry
of deuterostomes and the lophophorates can hardly
be held any longer. Whether these findings actually
support the Lophotrochozoa hypothesis, according
to which the lophophorate taxa are modified spiral-
ians, needs to be tested in a broader approach.
Among some other possible autapomorphies for a
taxon Radialia, consisting of Deuterostomia, Phor-
onida, Brachiopoda, and Bryozoa, there is no evi-
dence for a reduced spiral cleavage in lophophorates
(Zimmer 1997).
Acknowledgments. We thank the team from
Kristineberg Marina Forskningstation (Sweden) for help
with collecting the animals. This work was supported by
DFG BA 1520/3.
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