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409
J. Paleont., 82(2), 2008, pp. 409–414
Copyright 䉷2008, The Paleontological Society
0022-3360/08/0082-409$03.00
THE EARLIEST ENDOSYMBIOTIC MINERALIZED TUBEWORMS FROM THE
SILURIAN OF PODOLIA, UKRAINE
OLEV VINN
1
AND
MARI-ANN MO
˜TUS
2
1
Institute of Geology, University of Tartu, Vanemuise 46, 51014 Tartu, Estonia, ⬍olev.vinn@ut.ee⬎;
2
Institute of Geology, Tallinn University of
Technology, Ehitajate str. 5, 19086 Tallinn, Estonia, ⬍motus@gi.ee⬎
A
BSTRACT
—The earliest endosymbiotic tubeworms have been discovered within skeletons of the tabulate coral Heliolites sp. from the Silurian
(Ludlow) of Podolia, Ukraine. The new tubeworm species has a maximum diameter about 1 mm, a slightly conical tube, a smooth lumen in
the tube and a lamellar wall structure. The tube wall is 0.05–0.10 mm thick. The new endosymbiotic tubeworm Coralloconchus bragensis n.
gen. and sp. shares zoological affinities with the tentaculitids (incertae sedis) and is assigned to the Family Cornulitidae (Tentaculita, Cornulitida).
INTRODUCTION
T
UBES OF
endosymbiotic worms in the various invertebrates,
such as tabulate corals (Tapanila, 2004) and rugose corals
(Elias, 1986), are already common in the Ordovician (Tapanila,
2002, 2005), but they lack preserved mineralized shell. However,
if there were pre-Silurian endosymbiotic worm tubes of aragonitic
composition they would have been dissolved on the sea floor un-
less recrystallized early into low-Mg calcite (Cherns and Wright,
2000; Palmer and Wilson, 2004). Similarly, worm tubes of chi-
tinous composition would have disappeared. The exact biological
affinities of producers of worm-shaped Paleozoic bioclaustrations
are not known, but their morphology and ecology resemble mod-
ern polychaetes and vermetid gastropods (Tapanila, 2005). The
earliest known completely endosymbiotic animals with their own
exoskeleton are lingulid brachiopods from Heliolites corals in the
Silurian of Gotland (Richards and Dyson-Cobb, 1976), and from
the heliolitids, sarcinulid corals and stromatoporoids in the Silu-
rian of Anticosti Island (Tapanila and Copper, 2002; Tapanila and
Holmer, 2006). The ecology of the Ordovician gastropods en-
crusted by bryozoans (McNamara, 1978) and symbioses between
Silurian gastropod Semitubina sakoi Kase, 1986, and a favositid
coral (Kase, 1986) was different in that the host was a gastropod
on which bryozoan or coral larvae settled. In contrast to the Si-
lurian endosymbiotic lingulates, extensively encrusted Paleozoic
gastropods probably remained mobile and they definitely were not
endosymbiotic during their juvenile growth stage. The earliest
hitherto known endosymbiotic tubeworm-like fossils are Tor-
quaysalpinx Plusquellec, 1968 from the Middle Devonian stro-
matoporoids and tabulates (Stel, 1976; Zhen, 1996; Plusquellec,
1968; Tapanila, 2005) and Streptindytes Calvin, 1888 from the
Middle Devonian rugose corals (Calvin, 1888; Tapanila, 2005)
and stromatoporoids (Clarke, 1908; Tapanila, 2005). Darrel and
Taylor (1993) have discussed macrosymbioses in fossil corals.
Recently, all the Paleozoic endosymbiotic fossils have been
classified as ichnofossils (Tapanila, 2005), including those pos-
sessing their own skeleton such as Torquaysalpinx and Streptin-
dytes. However, the latter two do not meet the definition of an
ichnofossil (Bromley, 1970, 1996) because they contain fossilized
parts of the actual animal and thus are classical body fossils. Their
modern serpulid tubeworm analogues are similar, such as Spiro-
branchus giganteus (Pallas, 1766) in corals (Nishi and Nishihira,
1996) and Hydroides spongicola Benedict, 1887 in sponges,
which not only produce a trace (e.g. bioclaustration, see Palmer
and Wilson, 1988), but also secrete their own tube inside the host
skeleton. Their tubes are similar to those of substrate-cemented
and free-living serpulids (personal obs. O.V.). Several other Pa-
leozoic endosymbionts have previously been described as the
body fossils (Howell, 1962), such as Chaetosalpinx Sokolov,
1948 and Helicosalpinx Oekentorp, 1969. However, they lack
their own skeleton and are indeed bioclaustrations as interpreted
by Tapanila (2005).
In spite of striking morphological similarity to the modern ser-
pulid tubeworms (Weedon 1994; Taylor and Vinn, 2006), none of
the Paleozoic tubeworms such as sphenothallids (van Iten et al.,
1992; van Iten et al., 1996), cornulitids (Vinn and Mutvei, 2005),
microconchids (Weedon, 1990, 1991; Taylor and Vinn, 2006), try-
panoporids (Weedon, 1991) possess any of the key characters of
a serpulid tubeworm, such as the characteristic ultrastructure or
juvenile tube opened from both ends (Weedon, 1994; Vinn and
Mutvei, 2005; Taylor and Vinn, 2006). Most likely the earliest
true calcareous polychaete tubeworms appeared in the Mesozoic
(ten Hove and van den Hurk, 1993; Fischer et al., 2000; Vinn et
al., in press). Their characteristic tube structures and ultrastruc-
tures have been described from various Mesozoic worm fossils
(Senowbari-Daryan, 1997; Weedon, 1994; Vinn 2005a; Vinn et
al., in press).
The Silurian rocks (Wenlock-Pridoli) of Podolia (Ukraine) are
exposed in an approximately 80 km wide area along the river
Dniester and its tributaries (Fig. 1). The Silurian deposits were
formed in changing conditions from the normal-marine to lagoon
facies (Tsegelnjuk et al., 1983). The massive coral-stromatopo-
roid-algal bioherms (Grytsenko, 2007) from Lower to Upper Lud-
low of Podolia are characteristic of a shallow shelf environment.
The aim of this paper is to describe the earliest endosymbiotic
tubeworm and to discuss its biological affinities. This paper also
deals with the ecological and evolutionary implications of the
endosymbiosis of skeletal invertebrates.
MATERIAL AND METHODS
The coral specimens with the endosymbiotic tubeworms occur
in the locality ‘‘Zhavanets 39,’’ which is located on the left bank
of the River Dniester between the villages of Zhvanets and Braga
(Fig. 1). The Grinchuk Member of the Rykhta Formation is ex-
posed above the water level of the river (Tsegelnjuk et al., 1983).
The Grinchuk Member corresponds to the Uncinatograptus cau-
datus–Monograptus balticus Zone, upper Ludlow (Tsegelnjuk et
al., 1983) (Fig. 2). Heliolites sp. with Coralloconchus bragensis
n. gen. and sp. was collected from the layer of marl in the nodular
limestone approximately one meter above the water level from
the Grinchuk Member. The tabulate corals in the Grinchuk Mem-
ber at Zhavents 39 are represented by Squameofavosites incredi-
bilis (Tsegelnjuk et al., 1983), Syringoheliolites and at least five
species of Heliolites. Tesakov (1971) and Sokolov and Tesakov
(1984) described Favosites gothlandicus and Cystihalysites mir-
abilis from the Grinchuk Member near the village of Zhvanets.
We studied a collection of 185 tabulate corals from the Muksha
Member to the Grinchuk Member (Lower-Upper Ludlow). It in-
cludes seven species of Favosites, five species of Heliolites, one
species of Paleofavosites, 10 species of Cystihalysites, four spe-
cies of Cladopora, and two species of Syringopora.Riphaeolites,
Angopora,Stelliporella and Syringoheliolites are rare in collection
and are represented by only one species each. Material for study
410 JOURNAL OF PALEONTOLOGY, V. 82, NO. 2, 2008
F
IGURE
1—Map of Podolia (Ukraine), showing the location of outcrop
Zhavanets 39 from which Coralloconchus bragensis n. gen. and sp. was col-
lected.
F
IGURE
2— Stratigraphic context of the sample locality (after Tsegelnjuk
et al., 1983).
was collected from 13 localities. Two thin sections (one transverse
and one vertical section) were made of each coral specimen.
Thin sections containing tube worms were then polished and
etched with dilute (one per cent) acetic acid for 10 minutes and
coated with gold for SEM photography carried out with a Zeiss-
SEM at the Institute of Geology, University of Tartu.
Tubes of Recent calcareous annelids were selected for com-
parison with Coralloconchus bragensis n. gen. and sp. using the
database (unpublished, O.V.) of tube ultrastructures of 50 calci-
fying polychaete species. Examined Recent tubes of Glomerula
piloseta (Perkins, 1991) (Sabellidae), Pyrgopolon ctenactis
(Mo¨rch, 1863) (Serpulidae) and Dodecaceria coralii Leidi, 1855
(Cirratulidae), were embedded in epoxy resin, ground in longi-
tudinal and transverse direction, polished and etched with 1%
acetic acid for two minutes prior to SEM examination. Scanning
electron microscopy was carried out with a Hitachi S-4300 at the
Swedish Museum of Natural History, Stockholm.
All the specimens figured here are deposited at the Institute of
Geology, University of Technology, Tallinn (GIT 534-1-1, GIT
534-1-2, GIT 534-1-3, GIT 534-2-1).
SYSTEMATIC PALEONTOLOGY
Phylum I
NCERTAE SEDIS
Class T
ENTACULITA
Boucˇek, 1964
Order C
ORNULITIDA
Boucˇek, 1964
Family C
ORNULITIDAE
Fisher, 1962
Genus C
ORALLOCONCHUS
new genus
Type species.⎯Coralloconchus bragensis n. gen. and sp.
Diagnosis.⎯A genus of Cornulitidae with small, slender, irreg-
ularly curved conical tubes with slowly increasing diameter.
Tubes have thin walls and a smooth lumen. Tube wall has a la-
mellar microstructure. Tubes are devoid of septa and vesicles in
the adult part and are not spirally coiled.
Other species.⎯Monotypic thus far.
Etymology.⎯Corallum ⫽Latin for ‘‘coral’’; concha ⫽Latin for ‘‘shell’’;
the shell of the new genus is embedded in coral.
C
ORALLOCONCHUS BRAGENSIS
new species
Figures 3, 4, 5.1–5.4
Description.⎯Small, irregularly curved, slightly conical tubes with thin
walls (0.05–0.10 mm thick) embedded into host skeleton (coral) in their whole
length. Angle of divergence of tube about 10 degrees. Tube lumen smooth,
lacking septa at the tube diameters 0.5–1.05 mm. Exterior of tube mostly
smooth, rarely covered with variously developed perpendicular ridges. These
perpendicular ridges 50–60 m high, their inter val 0.1–0.3 mm. Shape of
ridges triangular in longitudinal section of tube. Tube wall up to twice as thick
at top of highest ridges as at interspaces between ridges. Tube wall lamellar;
lamellae 3.0–3.5 m thick, formed by calcite plates parallel to tube wall.
Types.⎯Holotype GIT 534-1-1, and three paratypes (specimens GIT 534-
1-2, GIT 534-1-3, GIT 534-2-1) from the Silurian (Ludlow) of Zhavanets 39,
Podolia, Ukraine.
Etymology.⎯After Braga village near the type locality of Zhavanets 39 in
Podolia.
Discussion.⎯The irregularly curved tube, smooth tube lumen
and shape of the external perpendicular ridges of the new species
are similar to Conchicolites Nicholson, 1872 (Tentaculita, Cor-
nulitida, Cornulitidae) (Vinn and Mutvei, 2005), but it differs in
being embedded into the coral skeleton and in having lamellar
tube ultrastructure. The lamellar wall structure and lack of pseu-
dopuncta of the new species is similar to Septalites Vinn, 2005b
(Tentaculita, Cornulitida, Cornulitidae), but Coralloconchus bra-
gensis has much thinner walls and lacks the septa. The new spe-
cies is also somewhat similar to Trypanopora Sokolov and Obut,
1955 (Tentaculita, Trypanoporida) and Torquaysalpinx (Tentacu-
lita, Trypanoporida), but it differs in lacking vesicular wall struc-
ture and septa. The new species slightly resembles the endosym-
biotic tubeworm Streptindytes acervulariae Calvin, 1888
(Tapanila, 2005, p. 93, fig. 1.) from the Middle Devonian rugose
corals of Iowa, but C. bragensis differs in having an irregularly
curved tube instead of the regularly coiled, inverted cone-shaped
shell of Streptindytes. Unfortunately, the morphology of the ju-
venile growth stage of Coralloconchus bragensis is not known
and cannot be compared with the cornulitids, microconchids (Ten-
taculita, Microconchida) and trypanoporids (Tentaculita, Trypan-
oporida). The zoological affinities of Streptindytes need revision,
but most likely it is either a microconchid or trypanoporid tube-
worm.
DISCUSSION
Calcareous annelid tubes.⎯In polychaete annelids, calcareous
tubes occur in serpulids (? Triassic-Recent) (ten Hove and van
den Hurk, 1993; Taylor and Vinn, 2006; Vinn et al., in press),
sabellids (Jurassic-Recent) (Ja¨ger, 2004; Vinn et al., in press), and
cirratulids (Oligocene—Recent) (Fischer et al., 2000). The fossil
and Recent calcareous annelid tubes possess a variety of ultra-
structures, but they are all different from that of Coralloconchus.
Their tubes usually are not lamellar (Fig. 5.6). Even in those
which are lamellar or contain lamellar layers, such as Recent
Glomerula piloseta (Sabellidae), Pyrgopolon ctenactis (Serpuli-
dae), and Dodecaceria coralii (Cirratulidae) (Fig. 5.5), the tube
lamellae have an aragonitic spherulitic prismatic ultrastructure
(Vinn et al., in press; Vinn, 2007) different from that of platy
homogeneous lamellae of Coralloconchus bragensis. Among cal-
careous polychaetes, only some Recent serpulids are thus far
known to be endosymbiotic (Nishi and Nishihira, 1996).
Vermiform gastropod shells.⎯Some Recent and fossil gastro-
pods (vermetids and siliquariids) have worm-shaped shells with
411VINN AND MO
˜TUS—ENDOSYMBIOTIC SILURIAN TUBEWORMS FROM THE UKRAINE
F
IGURE
3—1, 2, Coralloconchus bragensis n. gen. and sp. in Heliolites sp., Upper Ludlow, Silurian of Podolia, Ukraine, Scale bar 0.3 mm. 1, Longitudinal
section in plane-polarized light, paratype GIT 534-1-2; 2, cross section in plane-polarized light, holotype GIT 534-1-1, Scale bar 0.35 mm.
F
IGURE
4—1–3, Coralloconchus bragensis n. gen. and sp. in Heliolites sp.,
Upper Ludlow, Silurian of Podolia, Ukraine, through the light microscope;
large black circle on the photo is pyritized Trypanites boring. 1, cross section,
holotype GIT 534-1-1; 2, oblique section, paratype GIT 534-1-3. 3, longitu-
dinal section, paratype GIT 534-1-2. The arrows point to the specimens. Scale
bar 2.5 mm.
the irregularly coiled free whorls. Their oldest fossils are known
from the Mesozoic (Savazzi, 1996; Bandel and Kowalke, 1997;
Bandel and Kiel, 2000; Rawlings et al., 2001). The siliquariids
are endosymbiotic and live in sponges (Savazzi, 1996). The tubes
of vermetid gastropods have three aragonitic layers (Wrigley,
1950) which differ from the single-layered lamellar calcite tube
wall of Coralloconchus, the siliquariid tube wall having crossed
lamellar structure (see Savazzi, 1996, p. 169, text-fig. 10 D). Their
tubes also have slits different from the complete tube wall of
Coralloconchus bragensis. Moreover, all unequivocal vermiform
gastropods are geologically too young (Bandel and Kowalke,
1997; Bandel and Kiel, 2000) to be candidates for Early Paleozoic
endosymbiotic tubeworms.
Tentaculitid tubeworms.⎯A diverse fauna of tubeworms of
tentaculitid affinities is known from the Ordovician (Weedon,
1991; Weedon, 1994; Vinn, 2005b; Vinn and Mutvei, 2005; Tay-
lor and Vinn 2006; Vinn 2006a, 2006b; Vinn and Isakar, 2007).
Their diversity presumably reached a maximum by the Devonian.
The cornulitids range from the Ordovician (Caradoc) to the Car-
boniferous (Richards, 1974); trypanoporids are known only from
the Devonian (Weedon, 1991), microconchids range from the Or-
dovician (Ashgill) to Middle Jurassic (Taylor and Vinn, 2006;
Vinn and Taylor, 2007). The exact zoological affinities of tenta-
culitid tubeworms are not known, but most likely they represent
a group of extinct calcifying phoronids (Vinn, 2005b; Taylor and
Vinn, 2006). The earliest hitherto known endosymbiotic tenta-
culitid tubeworm is Torquaysalpinx from the Devonian. The ten-
taculitid tubeworms are characterized by lamellar tube structure
and closed, often bulbous, tube origins. Pseudopunctae are also
common in the tentaculitid tubeworms. The lamellar tube struc-
ture of C. bragensis closely resembles that of the other tentaculitid
tubeworms such as cornulitids, microconchids, trypanoporids.
The systematic position of C. bragensis within Tentaculita is also
well supported by the stratigraphic range of the new genus, which
coincides with the range of high diversity of the tentaculitid
worms.
Paleoecology of Coraliconchus.⎯Coralloconchus bragensis is
found only in Heliolites sp. of the Silurian of Podolia. It is a
relatively rare endosymbiont in the tabulate corals and occurs in
only two Heliolites sp. specimens of 34 specimens studied. In
contrast, the traces of soft bodied worms, such as Chaetosalpinx
siberiensis are common in other corals and are much more abun-
dant. The restricted occurrence of C. bragensis in Heliolites sp.
presumably indicates host specificity of the tubeworm. Corallo-
conchus bragensis infested living hosts and caused slight changes
in the growth of the surrounding coral skeleton (Figs. 4.1). How-
ever, the tubeworm was probably not parasitic, because changes
in the coral skeleton are in the geometrical arrangement of coe-
nechymal tubuli around the worm tube rather than in the diameter
of the tubuli (Fig. 5.1). Among the Recent coral endosymbionts,
serpulid tubeworms could ecologically be most similar to C. bra-
gensis.
Evolutionary implications.⎯The calcareous exoskeletons of
probable ancestors of C. bragensis supported their soft tissues and
hence biological function. In endosymbiosis, the exoskeleton of
a tubeworm loses its original protective and supporting function.
This is different from soft bodied worms (corresponding trace
fossils e.g., Helicosalpinx;Chaetosalpinx ferganensis Sokolov,
1948; C. siberiensis Sokolov, 1948), which achieved their primary
protection against predators and supporting functions via adap-
tation to the endosymbiotic lifestyle. The Silurian (Wenlock–Lud-
low) cornulitids bear abundant shell repair marks (Vinn and Mut-
vei, 2005), which could be caused by predation. Similar shell
repairs occur in tubeworm-like Anticayltraea calyptrata (Eich-
wald, 1860) (Tentaculita) from the Silurian (Ludlow) of Baltos-
candia (Vinn and Isakar, in press). There are unambiguous ex-
amples of selective predation on a problematic colonial metazoan
412 JOURNAL OF PALEONTOLOGY, V. 82, NO. 2, 2008
F
IGURE
5—1–4, Coralloconchus bragensis n. gen. and sp. in Heliolites sp., Upper Ludlow, Silurian of Podolia, Ukraine, SEM figures. 1, Cross section,
holotype GIT 534-1-1, showing the changes in the growth pattern of tubuli around the specimen; outer ring is penciled around the specimen for SEM
orientation; 2, close-up of the lower part of photograph 1, the arrows point to the external and internal surface of the tube wall of the specimen; 3, GIT 534-
1-1, close view of the cross section of the specimen’s tube wall, the arrows point to the external and internal surface of the tube wall of the specimen; 4,
GIT 534-1-1, detailed view of the lamellar tube wall of the specimen, showing the platy nature of the calcite lamellae forming the tube wall; 5, Dodecaceria
coralii Leidi, 1855, Recent, Mexico, cross section of the tube, showing the spherulitic prismatic ultrastructure of the aragonitic lamellae; 6, Pyrgopolon
ctenactis (Mo¨ rch, 1863), Recent, Bonaire, Netherlands Antilles, Caribbean; cross section of the outer nonlamellar layer of the tube, showing the semi-irregularly
oriented spherulitic prismatic crystals.
413VINN AND MO
˜TUS—ENDOSYMBIOTIC SILURIAN TUBEWORMS FROM THE UKRAINE
hederellid from the Devonian (Wilson and Taylor, 2006), and Ko-
walewski et al. (1998) have showed a significant increase in dril-
ling predation during the Devonian. The Devonian is also char-
acterized by the occurrence of two endosymbiotic tubeworm
genera, e.g., Torquaysalpinx and Streptindytes in tabulate corals,
rugose corals and stromatoporoids (Tapanila, 2005). One of pos-
sible reasons for the endosymbiosis of Coralloconchus could be
evolving boring predation capable of eliminating most of the pro-
tective advantages of the worm tube. The endosymbiotic C. bra-
gensis may also have been less endangered by overgrowth of the
other encrusting skeletons such as bryozoans as compared to en-
crusting cornulitids (probable ancestors).
ACKNOWLEDGMENTS
We are grateful to J. Aruva¨li, Institute of Geology, University of Tartu, for
help with SEM and H. A. ten Hove, Zoological Museum, University of Am-
sterdam, for providing us with the identified tubes of Recent serpulids. Vinn
is grateful to H. Mutvei, Swedish Museum of Natural History, for help with
SEM and advice on the methodology for studying skeletal ultrastructures. We
are grateful to B. Pratt, L. Tapanila and anonymous reviewer for the useful
comments on the manuscript. V. Gritsenko is thanked for the help in collecting
material in Podolia. Vinn acknowledges financial support to projects NL-TAF-
111 and SE-TAF-1520 by SYNTHESYS, a program financed by the European
Commission under the Sixth Research and Technological Development
Framework Program ‘‘Structuring the European Research Area’’ and the Es-
tonian Science Foundation for the grant No. 6623 ‘‘Tube formation and bio-
mineralization in annelids, its evolutionary and ecological implications.’’ Mo˜-
tus research was supported by the Estonian target funding programme No.
0332524s03 and the Estonian Science Foundation grants Nos JD05-41 and
6127. M. Wilson read earlier versions of the Manuscript.
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