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1319
Accepted by Jones: 15 Aug. 2006; published: 25 Sept. 2006
29
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334 (online edition)
Copyright © 2006 Magnolia Press
Zootaxa 1319: 29–42 (2006)
www.mapress.com
/zootaxa/
Pyrgopsella (Cirripedia: Balanomorpha: Pyrgomatidae)
is not a sponge-inhabiting barnacle
YAIR ACHITUV* & NOA SIMON-BLECHER
The Mina & Everard Goodman Faculty of Life Sciences, Bar Ilan University, Ramat Gan 52900, Israel
*Corresponding author:E-mail: achity@mail.biu.ac.il
Abstract
A new species of coral-inhabiting barnacle, Pyrgopsella youngi, is described. It was found in a
colony of the coral Symphyllia radians Milne Edwards & Haime, 1849 from Sulawesi, Indonesia.
The barnacles were suspended in the coral tissue and were easily detached. A unique feature of
Pyrgopsella is its membranous basis; in P. youngi the calcareous basis is reduced to a vestige by
which the barnacle is attached to the coral. Pyrgopsella has been regarded as a genus of sponge-
inhabiting barnacle, thus unique in the otherwise coral-associated family Pyrgomatidae, but our
findings confirm that this genus too comprises coral-inhabiting barnacles. We propose a new genus,
Pyrgospongia, to accommodate the sponge-inhabiting barnacle originally described as Pyrgopsella
stellula Rosell, 1973. The relationships of both Pyrgopsella and Pyrgospongia within the
Pyrogomatidae are discussed.
Key words: Barnacles; Pyrgomatidae; Corals; Pyrgopsella; Pyrgospongia
Introduction
Coral-inhabiting barnacles of the family Pyrgomatidae are obligatory symbionts of
scleractinian corals, hydrocorals and sponges. The shell of these barnacles is composed of
either four or two calcareous wall plates or a unified calcareous shell wall and a basis. The
shell wall is flat to low conical and projects above the host coral’s surface. The basis is
usually cup-shaped or tubular, tapering and usually embedded deeply in the coral skeleton.
The presence of a fully calcified basis is one of the dominant features of the pyrgomatids,
except in Pyrgopsella (the unique genus of the tribe Pyrgopsellini), with a nearly entirely
membranous basis, and the members of the tribe Hoekini, in which the wall is separated
from the calcified part of the basis by a narrow membranous zone.
Gruvel (1906; 1907) found in the collections of the Indian Museum in Calcutta three
specimens of a pyrgomatid barnacle, dredged from a depth of 90 m in the Andaman
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Islands, Bay of Bengal (11º49’30’’N, 92º55’55’’S). He assigned these to a new genus and
species, Pyrgopsis annandalei. Most subsequent authors have attributed this taxon to
Gruvel (1907), but the species name was actually made available in his preliminary report
of 1906. The genus name, however, proved to be an invalid junior homonym and was later
replaced with the new name Pyrgopsella by Zullo (1967). The most unusual characteristic
of P. annandalei was the membranous basis. The host was unknown, but the absence of a
calcareous basis led Baluk & Radwanski (1967) to the conclusion that Pyrgopsella should
not be assigned to the Pyrgomatinae. Ogawa & Matsuzaki (1992) also urged omission of
Pyrgopsella from among the coral-barnacles, owing to “unsolved ecological and
evolutional problems” that classification within the Pyrgomatidae would imply.
Since its description by Gruvel (1907) Pyrgopsella was not reported and already in
1938 it was not found in the collection of the Indian Museum in Calcutta (Nilsson-Cantell
1938). Rosell (1973; 1975) found another barnacle with a membranous basis embedded in
a sponge in the Philippines and described it as a second species of Pyrgopsella, P. stellula.
Rosell (1975) interpreted the membranous basis as an adaptation for living in sponges, a
situation analogous to that in the sponge-inhabiting barnacles of the archaeobalanid genus
Membranobalanus (Rosell 1975) and he suspected that Gruvel’s (1906; 1907) P.
annandalei also came from a sponge. Ross and Newman (1973) accepted this assumption
and it has become established in the scientific literature (e.g., Galkin 1986; Ogawa &
Matsuzaki 1991; Ross & Newman 1995; Newman 1996). The reduced shell plates of the
sponge-dwelling archaeobalanids of the genus Acasta and the plasticity of the shell in this
genus are also regarded as adaptations for living within sponges (Ilan et al. 1999).
While examining a colony of the coral Symphyllia radians Milne Edwards & Haime in
a tropical fish shop in Israel, the authors noticed the presence of coral-inhabiting barnacles
suspended in the coral tissue. Barnacles detached from the coral were found to be similar
to Pyrgopsella annandalei. When the coral was partly dried with a few barnacles still
attached, it became evident that the membranous basis is attached to the coral at its
proximal tip by a small, vestigial, calcareous disc or ring. The close similarity between our
material to the description of P. annandalei poses the question whether P. annandalei is a
sponge-inhabiting barnacle, or rather a coral-inhabiting barnacle with a reduced calcareous
basis. In the present report we describe these specimens as a new species of Pyrgopsella.
We also compare our material to paratypes and other material of Pyrgopsella stellula. On
the basis of differences in the opercular plates and other morphological features and the
different host phylum we propose here a new genus, Pyrgospongia, to accommodate this
sponge-inhabiting pyrgomatid barnacle.
Materials and methods
Fifteen specimens of the new species were examined, partly by light microscopy and
partly by scanning electron microscopy (SEM), as noted in detail below. In addition, three
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paratype specimens of Pyrgopsella stellula from the Philippines were obtained on loan
from the National Museum of Natural History, Smithsonian Institution (USNM 141633);
one of these was used for SEM of the shell and opercular plates. Three other specimens of
this species from Japan were obtained on loan from Mark J. Grygier (personal collection);
these had been collected by the late Tatsunori Itô on 18 June 1989 at 5 m depth off Tôhima
Rock, Tanabe Bay, Japan, from a sponge, and are part of the material reported by Grygier
(1992). Three additional paratypes of P. stellula, as well as additional Japanese specimens
collected by Itô, all of which had been housed in the Benthic Invertebrates Collection of
the Scripps Institution of Oceanography, were lost in transit from California to Israel.
The barnacles were removed from the host coral, examined, photographed and
dissected under dissecting microscope. The cirri and mouth parts were removed, mounted
in glycerin jelly on microscope slides and the slides sealed with nail varnish. The slides
were examined and photographed using a Vanox Olympus microscope. The wall plates
and opercular valves of the barnacles were separated from the coral, immersed for ~two
hours in household bleach and rinsed in tap water followed by distilled water. The
specimens were examined under a dissecting microscope. Adhering chitin was removed
using needles and fine forceps and the valves were dried on a small hot plate at 80ºC. The
shell of the paratypes of Pyrgopsella stellula was dehydrated through an ascending series
of alcohols and acetone and mounted on an SEM stub; the opercular valves of Pyrgopsella
stellula were treated as those of the coral barnacles. The dried samples were mounted on
brass stubs, coated with gold and examined with a JOEL scanning electron microscope at
25 kV. Images were copied and stored using Autobeam software.
Specimens have been deposited in the National Museum of Natural History, Naturalis
(RMNH), Leiden, The Netherlands, and the Zoological Museum, Tel Aviv University,
Israel (TAU).
Taxonomy
Pyrgopsella Zullo 1867
Pyrgopsis Gruvel 1907
Pyrgopsella Zullo 1967
Diagnosis: Walls sub-conical, totally concrescent. Basis membranous with short peduncle.
Opercular plates separate, scutum transversally elongated, tergum triangular.
Type species: Pyrgopsella annandalei Gruvel 1907.
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Pyrgopsella youngi sp. nov. Achituv
(Figs 1–6)
Material examined: Fifteen specimens, six with soft parts, were obtained from a partly
bleached colony of Symphyllia radians Milne-Edwards and Haime, found in a tropical fish
shop in Israel. The source of the coral was Sulawesi, Indonesia, exact location unknown.
The barnacles were suspended in the coral tissue and easily detached from the coral. Part
of the host coral (catalogue number RMNH Coel. 33023) and the barnacle holotype
(RMNH C. 2602: two slides with cirri and mouth parts, SEM stubs of shell and opercular
valves) and two paratypes (RMNH C 2603 and RMNH C 2604: shell and opercular valves,
and SEM stubs with shell and opercular valves, respectively) have been deposited in the
RMNH. The other part of the host coral (catalogue number TAU Co32349), the remaining
paratypes, shell and opercular valves of four specimens prepared for SEM analysis, slides
of two specimens with cirri and mouth parts, are deposited in the TAU (catalogue number
TAU Ar27804). Two specimens were used for DNA extraction.
Etymology: The specific epithet honors the late Paulo Young in recognition of his
contribution to our knowledge of cirripede biology.
Diagnosis: Wall concrescent, ribs on shell absent. Basis, membranous, with inferior
end calcareous. Scutum and tergum not fused, scutum transversally elongated, tergum
triangular with internally directed tooth.
Description: Shell (Fig. 1C, 2A, 3A, B) white but nearly transparent, concrescent,
low-conical, thin, oval, maximum carino-rostral diameter 8 mm, maximum lateral
diameter 4 mm. Radiating rib absent; shallow, concentric growth lines on outer surface.
Shell tubiferous, tubes wide, penetrating 2/3 of shell; inner shell white; sheath with
concentric growth lines, reaching margins of shell; short spines on sheath perimeter
located at margins of lateral septa, number of spines equal to number of septa, with no
denticulation on margins of septa. Orifice oval, located at carinal end of shell; carino-
rostral diameter 1/3 carino-rostral diameter. Tergoscutal flaps banded purple on white
cream ground. Basis (Fig. 1B) membranous, sometimes with basal part elongated and
peduncle-like, basal tip connected to small, white, calcareous disc, latter usually attached
to septum of coral calyx (Fig. 3A,B).
Scutum and tergum white, separate. Scutum (Fig. 2C, E) transversally elongated, thin,
total length (including tergal tooth) ~five times maximal width; basal margins slightly
sinusoidal; adductor muscle pit shallow, distinct; adductor ridge small, low; lateral
depressor muscle pit indistinct; width of tergal tooth ~1/2 width tergal margin, located
closer to occuludent margin then to basal margin; growth lines on outer surface; narrow,
oblique furrow beginning at tergal margin halfway between tergal tooth and occludent
margin, ending near occludent basal angle. Tergum (Fig. 2D, F) triangular, growth lines on
outer surface; short, spur barely distinct; external groove running from middle of scutal
margin to basi-carinal apex; small notch in middle of scutal margin; basi-scutal angle
pointed; thin, pointed, inward-projecting tooth on spur; notch in middle of carinal margin;
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inner side with depression accommoding tergal tooth of scutum; growth lines visible
inside depression.
FIGURE 1. A. Symphyllia radians, the host coral of Pyrgopsella youngi; ~15 barnacles were
removed from this colony; note the absence of pits, owing to the barnacle’s near absence of a
calcified basis. B. Pyrgopsella youngi, paratype RMNH C 2604 from Symphyllia radians. C.
Pyrgospongia stellula, shell of paratype USNM 141633, Bauang Dakula Reef, Tawitawi, Sulu
Archipelago, Philippines, collected and identified by N. C. Rosell. D. Pyrgospongia stellula,
Tanabe Bay, Japan, coll. T. Itô, det. M. J. Grygier, arrow indicates basi-occuldent angle of scutum.
Scale bars: A = 5 cm; B = 2 mm; C = 1 mm; D = 1 mm.
Labrum (Fig. 4A, E) rounded with deep median notch, very short, fine setae on
margins and outer surface; palps (Fig. 4A) elongate, oval with upper margins straight, long
setae on outer surface, longest at distal end, inner margins with shorter setae. Mandible
(Fig. 4C) with five teeth along cutting edge, distances between teeth unequal, upper two
teeth occupying 1/2 length cutting edge, second and fourth teeth bifid, lower angle with
short spines; face and margins of mandible bearing setae, lower margins with combs,
conical spines on lower angle. Maxillule (Fig. 4D) with cutting edge unnotched, armed
with row of ~ten unequal spines, tuft of smaller, hair-like setae at distal angle; setae
scattered along upper part of maxillule, row of dense setae on lower part; some combs of
hairs on face of maxillule projecting beyond cutting edge. Maxilla (Fig. 4B) oval-elongate,
long setae distally, short setae on inner side, few simple setae on upper and lower margins,
distal surface with two rows of short, sharp setae; lower angle with two to three spines.
Number of articles of cirri of three randomly selected individuals given in Table 1.
Cirrus I (Fig. 5A) highly setose; anterior ramus ~twice length of posterior ramus; proximal
articles of anterior ramus and all segments of posterior ramus with protuberances bearing
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setae. Cirrus II (Fig. 5B) with rami ~equal length; articles slightly protuberant, bearing
long setae. Cirrus III with anterior ramus somewhat longer than posterior ramus; articles
with protuberances bearing tufts of long, plumose setae (Fig. 5C). Cirri IV (Fig. 5D) to VI
(Fig. 6A) long and slender; each article with four pairs of setae of different lengths
arranged along anterior margin, generally each pair accompanied by short, proximal seta;
setae at distal end of articles longer, ~three times article width; two or three short setae at
posterior articulation of each article (Fig. 6B); basal articles with a row of pairs of setae.
Penis (Fig. 6A) long, annulated, scattered thin setae along length; distal end modified,
more slender then rest of penis, not annulated, few setae (Fig. 6C).
FIGURE 2. Pyrgopsella youngi, shell wall and opercular valves of holotype (RMNH C. 2602). A.
Shell wall, outer view. B. Shell wall, inner view. C. Scutum, inner view; D. Tergum, inner view.
E. Scutum, outer view. F. Tergum, outer view. Scale bars = 1 mm.
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TABLE 1. Cirral counts of three randomly selected specimens of Pyrgopsella youngi from
Symphyllia radians, expressed as a–b for numbers of articles in anterior (a) and posterior (p) rami,
respectively, of each cirrus; + indicates a broken ramus, R = right cirrus, L= left cirrus.
FIGURE 3. Pyrgopsella youngi. A. Shell wall of a specimen and its vestigial basis attached to the
septum of a coral calyx. B. Disc-like basis (arrow pointing to basis) attached to the septum of a
coral calyx. C. Internal view of tergum tilted on its basal margin, showing pointed scuto-basal
margin and internal tooth. Scale bars: A, B = 1 mm; C = 500 µm.
Remarks: The barnacles from Symphyllia radians differ from those described by
Gruvel (1907) in two striking aspects: the tergum as depicted and described by Gruvel
carries a truncate internal tooth, whereas the inward projecting tooth found in our
specimens is thin and pointed and located on the short spur. The second difference
concerns the distal end of the penis. In Pyrgopsella annandalei it is ornamented with short
spines, which are missing in our material, and the modified end of the penis in Gruvel’s
material is pear-like, while in the present material this part of the penis is elongate. On the
basis of these differences, we conclude that our material is not referable to P. annandalei,
but should be assigned to a new species, named by us as P. youngi.
In most coral-barnacles the shell is overgrown by the coral and calcareous material is
deposited on the barnacle surface. In many cases the coral forms spines or perturbations
over the surface of the shell plates and in some cases even an entire corallite. However, in
Pyrgopsella, soft coral tissue covers the shell with no deposition of calcareous material
Cirrus I II III IV V VI
Specimen 1 R 15–8 10–8 12–9 30–15+ 29–26 29–24+
L 15–7 9–8 12–9 32–23 28–23 28–28
Specimen 2 R 13–8 8–7 10–8 20–18 18–+ 5+-23
L 16–7 9–7 10–8 23–20 +-26 20–24
Specimen 3 R 16–8 8–8 12–12 23–4+ +-22+ 30–26
L 17–7 7–7 10–9 26–22 23–24 13+-25
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over it and the barnacle comes to be suspended in the coral tissue. A more remarkable
difference between Pyrgopsella and other pyrgomatids is the membranous basis with a
vestigial, calcareous part. In all other coral-inhabiting barnacles the basis is calcareous
and, in most cases, it is cone-like and tapered, embedded in the coral skeleton. A particular
feature of the coral Symphyllia radians is the thick, fleshy tissue in which the barnacles are
embedded. This thick tissue supports the barnacles, thus making a calcareous basis
superfluous.
FIGURE 4. Pyrgopsella youngi, trophi. A. Labrum with overlying palps. B. Maxillae. C.
Mandible. D. Maxillule. E. Half of labrum. Scale bars: A–D = 500 µm; E = 200 µm.
Rosell (1973; 1975) reported the existence of a barnacle similar to Pyrgopsella
annandalei embedded in a sponge collected in the Philippines. He described this barnacle
as a new species, P. stellula. Grygier (1992) reported the occurrence of P. stellula in Japan,
extending the geographical distribution of this species, and noted an additional species of
Pyrgopsella in a sponge from Indonesia. Rosell (1973; 1975) suggested that the genus
Pyrgopsella consists of sponge inhabiting barnacles and assumed that the host of P.
annandalei was also a sponge. This view has been widely accepted, but our new find
suggests that Pyrgopsella is a coral-barnacle and that Gruvel’s species, which is similar to
ours, most probably was detached somehow from a coral.
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FIGURE 5. Pyrgopsella youngi, cirri. A–D. Cirri I to IV, respectively. Scale bars = 250 µm.
It is surprising that for a century, ever since its description by Gruvel (1907),
Pyrgopsella was never reported from corals. Both P. annandalei and P. youngi are rather
large pyrgomatids, reaching a carino-rostral diameter of 8 mm, and can hardly be missed.
The lack of records until now can be explained by the way in which coral-inhabiting
barnacles are studied. Mostly, the barnacles are extracted from dry corals kept in museum
collections. Moreover, since identification of corals is based on the structure of the calices
and septa, in many cases biologists remove the coral tissue using household bleach or an
alkali solution, whereby Pyrgopsella with its membranous basis would be lost.
Examination of the dried and bleached skeleton of the coral from which the present sample
was removed showed no conspicuous evidence of the presence of the barnacles. Only
detection of the vestigial calcareous basis could show that barnacles had formerly lived on
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the coral. In the collection of Naturalis, Leiden, there is a colony of Symphyllia radiance
from S.W. Sulawesi (RMNH Coel 24745) with vestigial calcareous basis. Our new
findings underline the importance of examining live corals, or corals preserved with their
tissue, for the presence of barnacles, a technique that might reveal additional kinds of
coral-inhabiting barnacle.
FIGURE 6. Pyrgopsella youngi, details. A. Cirri VI and penis. B. Articles of cirrus IV. C. Tip of
penis. Scale bars: A = 500 µm; B, C = 250 µm.
The main feature that Pyrgopsella stellula (Figs. 1C; 7A, B), P. annandalei (Pl. 2 Fig.
7b in Gruvel, 1907) and P. youngi (Fig. 2A, B) share is a single-plate shell, but otherwise it
is difficult to accept that these species all belong to the same genus. In addition to being
symbionts of hosts from different phyla, there are striking morphological differences
between these species. Pyrgopsella stellula lacks the peduncle that is present in P.
annandalei (Pl. 2 Fig. 7a in Gruvel, 1907) and in P. youngi (Fig. 1B, D). In our material,
and one may perhaps assume also in P. annandalei, the basal tip of the peduncle anchors
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the barnacle to the septa of a coral calyx. The shell of P. stellula is thin and flexible and
only partly calcified (Fig. 7 A, B) The calcareous area is in the form of thin spokes
radiating from the operculum and a flexible membrane connects those spokes. In P. youngi
and P. annandalei, the shell is totally calcified. In P. stellula there is no distinct sheath,
whereas it is distinct in P. annandalei and P. youngi (Fig. 2B). The sickle-shaped, barbed
bristles and star-like spines (Fig. 7G) that are arranged in concentric rows over the outer
shell membrane in P. stellula are absent in P. youngi. The shape of the opercular valves of
P. stellula (Fig. 7 C, D, E, F) and those of P. annandalei (Pl. 2 Figs 9,10 in Gruvel, 1907)
and P. youngi (Fig. 2 C, D, E, F) are different. The scutum of P. stellula is elongated, rather
big and lacks a tergal tooth (Fig. 7D); in situ it projects beyond the rostral margins of the
shell (Fig. 1D; see also Rosell 1975: Fig. 2c). The tergum is missing the internal tooth that
is found in the other species of Pyrgopsella (Figs 2D, 7C). Cirrus III of P. stellula carries
small conical spines that are not found in P. youngi and cirrus IV of P. stellula is armed
with diagonally arranged conical spines (Rosell 1975: Fig. 2n) that are absent in P. youngi.
On the basis of these differences we suggest that P. youngi sp. nov. and P. annandalei do
not share the same evolutionary line as P. stellula and that the latter should be assigned to
a different genus. Therefore, we propose a new genus, Pyrgospongia, to accommodate the
sponge-inhabiting pyrgomatid Pyrgospongia stellula (Rosell 1973).
Pyrgospongia gen. nov.
Diagnosis: Wall concrescent, thin, made of radiating rods connected by chitinous material,
elliptical in outline; ribs on shell absent; heath absent. Basis membranous. Scutum and
tergum not fused, scutum transversally elongated, tergum triangular.
Type species: Pyrgopsella stellula Rosell, 1973.
Etymology: From the Greek pyrgos (tower), found in the family name Pyrgomatidae of
the coral-inhabiting barnacles, and the Latin (from Greek) spongia (sponge), referring to
the host group.
Remarks: Based on the morphology of the shell and the opercular valves, Ross &
Newman (1973) suggested that Pyrgopsella annandalei is an offshot of the “Savignium
line” of pyrgomatids. Anderson (1992) suggested that P. annandalei is a specialized
member of the line of Savignium crenatum (Sowerby). However, the opercular valves of
our material, as well of the material depicted by Gruvel (1907), are closer in form to those
of Trevathana dentata (Darwin); a tergal tooth and a thin, pointed, inward-projecting tooth
located on the spur are characteristic of T. dentata but missing in Savignium crenatum. The
position of Pyrgospongia is more ambiguous. The opercular valves, elongated scutum, and
especially the tergum of Pyrgospongia stellula, without any sign of an internal tooth,
appear to be more similar to those of a different pyrgomatid line comprising those species
of Cantellius with transversally elongated scuta, such as C. brevitergum (Hiro) or C.
transversalis (Nilsson-Cantell). It might be supposed that Pyrgospongia was derived from
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FIGURE 7. Pyrgospongia stellula, SEM micrographs of paratypes, USNM 141633. A. Shell with
chitinous external cuticle of rostral part removed to expose radial calcareous rods within (partly
deformed due to drying during preparation for SEM examination). B. Shell of another specimen,
inner view. C. Tergum, outer view. D. Scutum, outer view. E. Tergum, inner view. F. Scutum, inner
view. G. Piece of shell-region outer cuticle showing concentric rows of barbed bristles and star-
shaped spines. Scale bars: A–F = 1 mm; G = 200 µm.
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within Cantellius. On the other hand, being a sponge-inhabiting rather than coral-
inhabiting barnacle and having rather pleisomorphic opercular valves, the Philippine and
Japanese P. stellula may not be a pyrgomatid at all. Molecular analysis using DNA
sequences may shed light on this enigma.
Acknowledgments
This study was supported by the Israel Scientific Foundation (grant no. 430/03-3). Visit of
YA to Naturalis, Leiden, was supported by the European Commission's Research
Infrastructure Action via SYNTHESYS project (NL-TAF 2088). We acknowledge Mr.
Assi Arbiv, Mr. Saar Ayache and Mr. Oren Orgil of ASEA Holon, Israel, who donated the
coral with the barnacles. Dr. Yaacov Langsam’s help with the preparation of the SEM
micrographs is greatly appreciated. Dr. Frank D. Ferrari and Dr. Cheryl Bright arranged
the loan of specimens from the Smithsonian Institution. We thank Prof. W.A. Newman for
his critical reading of and comments on an early draft and for preparing a loan of
specimens that went astray. Dr. M. J. Grygier, besides providing specimens on loan,
assisted us greatly in improving the manuscript based on his unpublished studies of the
morphology of sponge-associated pyrgomatids (cf. Grygier, 1992). We thank Dr C.H.J.M.
Fransen, Dr B. Hoeksema and Dr L.P. van Ofwegen of Naturalis for help and hospitality.
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