Three New Loxosomella (Entoprocta: Loxosomatidae)
from Coral Reef Shore in Okinawa, Ryukyu Archipelago,
Japan
Author: Iseto, Tohru
Source: Zoological Science, 18(6) : 879-887
Published By: Zoological Society of Japan
URL: https://doi.org/10.2108/zsj.18.879
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ZOOLOGICAL SCIENCE 18: 879 –887 (2001)
© 2001 Zoological Society of Japan
Three New Loxosomella (Entoprocta: Loxosomatidae) from
Coral Reef Shore in Okinawa, Ryukyu
Archipelago, Japan
Tohru Iseto*
Department of Chemistry, Biology and Marine Science, Faculty of Science,
University of the Ryukyus, Nishihara, Okinawa 903-0213, Japan
ABSTRACT—Three new species of solitary entoprocts of the genus Loxosomella, L. monocera sp. nov., L.
lappa sp. nov., and L. aloxiata sp. nov., are described from coral reefs in Okinawa Island and Sesoko Island,
Ryukyu Archipelago, Japan. This is the first report on entoprocts from this archipelago. Whereas most of
Loxosomella species are known to be epizoic, the three species were found on stones and shell remains in
the shallow reef flat or on the slide glasses settled there. It is thus probable that the present species do not
depend on host animals unlike most other congeneric species.
INTRODUCTION
Entoprocts (=Kamptozoans) are solitary or colonial,
sessile animals which occur in intertidal zone to deep seas of
more than 500 m depth. Only one species, Urnatella gracilis,
is reported from fresh waters. They are known to be phylogenetically enigmatic animals. Recent molecular studies suggest that this phylum is related to Cycliophora, Annelida,
Mollusca, and other groups having trochophore larvae (Mackey
et al., 1996; Zrzavy et al., 1998). To date, a total of approximately 160 species are recognized for this phylum, of which
ca. 50 species are colonial (the order Coloniales) and the remainder are solitary (the order Solitaria). The taxonomical
study of this group is still at an early stage, and there seem to
be a number of entoprocts yet to be described (e.g. Nielsen,
1982).
From Japan, 29 entoprocts have been reported, of which
17 (10 Loxosomella and 7 Loxosoma) are solitary and 12 (2
Pedicellina, 7 Barentsia, 1 Pseudopedicellina, 1 Loxosomatoides, and 1 Urnatella) colonial (Oka, 1895; Toriumi, 1949,
1951; Yamada, 1956; Konno, 1971, 1972, 1973, 1974, 1975,
1976, 1977a, b; Ikeda et al., 1977; Oda, 1982). All these species are reported from around the main islands of Japan: no
entoprocts have ever been reported from the Ryukyu Archipelago, a chain of over 100 islands ranging between Kyusyu
of Japan and Taiwan, surrounded by coral reefs. This area
enjoys tropical climate, which contrast to temperate climate in
most of the rest of Japan.
In other part of the world, Entoprocta have been reported
from tropical waters. For example, 19 species including both
* Corresponding author: Tel. +81-98-895-8880;
FAX. +81-98-895-8576.
E-mail: k998551@sci.u-ryukyu.ac.jp
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solitary and colonial ones are known from the Indo-Malayan
region (Harmer, 1915), and three solitary species of the genus Loxosoma from Thailand (Nielsen, 1996). It is thus likely
that entoprocts actually occur in the Ryukyus, and that the
absence of records of this group there merely reflects an absence of surveys in the Ryukyus by its taxonomist.
I have surveyed entoprocts in shallow waters of Okinawa
Island, the largest island of the Ryukyu Archipelago, and of
an adjacent small island. As a result, three forms of solitary
entoprocts, each representing an undescribed species, have
been collected from their reef flats as below.
MATERIALS AND METHODS
Sampling was carried out in Mizugama (Okinawa Island) and
Sesoko (Sesoko Island) (Fig. 1). Slide glasses were immersed in reef
flats, and were collected after two months. Stones and shell remains
were also collected in these reef flats. Epifaunas on these objects
were examined under a binocular stereomicroscope. Some of these
stones and glasses, as well as shells, were examined immediately
after collection, whereas others were kept in a bucket of seawater
with glass plates for 1– 2 months in order to obtain asexually proliferated individuals attached on the glass plates.
To observe budding modes and liberated buds, some animals,
removed from surfaces of these objects, were temporally cultured in
petri dishes. When the specimens had large buds, the buds grew up
and dropped off from the mother within a few days. When the specimens did not have large buds, prolonged culture was carried out by
adding sufficient amount of microalgae (Marine Chlorella 100,
Marine-bio Co., Japan) to each dish one time a day. Within 30 min
after each treatment, stomach of each animal was filled with the algae. Then, the seawater of the dish was exchanged. The buds grew
up on mother animals and were liberated after one week or so. In
some species, the newly liberated buds metamorphosed and settled
on the dishes within a few days.
0.37M of MgCl2, up to equal volume to seawater, was added to
the sample for narcotization, and then, the specimens were fixed in
880
T. Iseto
Fig. 1. Map of Okinawa Island and its vicinity, showing geographic locations of Mizugama and Sesoko Island, the type localities of the species
described here.
2% formalin in seawater. All the drawings were prepared on the basis
of fixed specimens under a light microscope equipped with a camera
lucida. Type specimens are deposited in the National Science
Museum, Tokyo (NSMT).
DESCRIPTION
Family Loxosomatidae Hincks, 1880
Genus Loxosomella Mortensen, 1911
Loxosomella monocera sp. nov.
(Figs. 2 and 3)
Material examined.
Holotype: An adult (NSMT-Ka 20), derived from budding
from an individual found on a shell remain from the reef flat at
a depth of ca. 1 m in front of the Sesoko Station of the Tropical Biosphere Research Center (University of the Ryukyus),
Sesoko Island, Okinawa, Japan (26°38’N, 127°52’E) on 12
July 1999.
Paratypes: Five adults: NSMT-Ka 21, the mother individual of the holotype; NSMT-Ka 22, sibling individuals of the
holotype derived from NSMT-Ka 21 by budding. Two liberated buds: NSMT-Ka 23, derived from NSMT-Ka 21; NSMTKa 24, derived from an individual collected from reef flat in
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Mizugama (estuary of Hija River), Okinawa Island, Okinawa,
Japan (26°21’N, 127°45’E), on 12 September 1999.
Etymology: The specific name “monocera” is composed
of two Greek words, “monos”, meaning “single”, and “keras”
meaning “horn”, in reference to the characteristic single appendage on the abfrontal side.
Diagnosis.
Adult: Total length (from basal tip of stalk to uppermost
part of tentacle membrane) up to 420 µm. Tentacle number
invariably 10. Stalk short: about half as long as calyx. No foot
in adult. Lateral sense organs absent. Lateral lobe of stomach absent. One conspicuous appendage, ca. 50 µm in length,
on abfrontal midline of calyx. Liberated bud: Slug-like
appearance with no boundary between calyx and stalk or between stalk and foot. Total length 350 µm in NSMT-Ka 23,
420 µm in NSMT-Ka 24. Tentacles folded. Foot groove
present. Lateral sense organs absent. Appendage present at
posterior, outer edge of lophophore, probably homologous to
adult abfrontal appendage.
Reproduction: Buds emerge from the frontal area of
calyx at level of the lower side of the stomach, attaching themselves to their mother’s body by tips of their feet; as growing
Three New Loxosomella from Okinawa
881
Fig. 2. Holotype of Loxosomella monocera (NSMT-Ka 20) with three buds (b). All buds are still in early stage. Arrows indicate the abfrontal
appendage characteristic for this species. A–C: Drawings of frontal, lateral and abfrontal views, respectively. D, E: Photographs of frontal and
lateral views, respectively. F: Lateral view of abfrontal appendage in greater magnification. Bars =100 µm.
Fig. 3. A liberated bud of Loxosomella monocera (NSMT-Ka 24, paratype). Arrows indicate an appendage which may be homologous to the
abfrontal appendage in adults. Bars=100 µm.
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T. Iseto
larger, they extend leftward or rightward. A maximum of three
buds were observed simultaneously on a single mother. Nevertheless, only one bud grew large at a time. No larvae were
found.
Remarks: From about 80 species hitherto described for
the genus Loxosomella, L. monocera is distinct in having the
characteristic abfrontal appendage. Also, morphology of the
liberated bud of the present species is unique among the congeners. The liberated buds do not expanded their tentacles
before metamorphosis into adult form, and thus they never
swam using the tentacular cilia as the liberated buds of other
species do.
The function of the abfrontal appendage remains unknown. It does not seem to be comparable to the lateral sense
organs found in many other species of the genus, because
these organs, invariably located on both of the lateral sides of
the calyx, are considerably shorter than the abfrontal appendage of L. monocera.
Fig. 4. Holotype of Loxosomella lappa (NSMT-Ka 25) with a small bud (b). A, B: Drawings of laterofrontal and abfrontal views, respectively. C–
E: Photographs of laterofrontal, abfrontal and lateral views, respectively. F: Lateral sense organ. Arrows in A–F indicate lateral sense organs. G:
Longer (thick arrow) and shorter (thin arrow) appendages at base of calyx. Bars =100 µm (A–E), or 50 µm (F and G).
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Three New Loxosomella from Okinawa
This species was easily cultured in a petri dish. The liberated buds metamorphosed, settled on the substrata and generated the buds of the next generation.
Loxosomella lappa sp. nov.
(Figs. 4–6)
Material examined.
Holotype: An adult (NSMT-Ka 25) collected on 4 April
2000 from a glass plate that had been stored for two months
in the laboratory with stones collected from the reef flat at a
depth of ca. 1 m in Mizugama (estuary of Hija River), Okinawa
Island, Okinawa, Japan (26°21’N, 127°45’E). Paratypes: 53
adults (NSMT-Ka 26), one young specimen (NSMT-Ka 27),
and one liberated bud (NSMT-Ka 28) all collected from the
glass plate described above.
Etymology: The specific name “lappa” is a Latin word
meaning “bur”. This refers to the characteristic appendages
covering the body of this species.
Diagnosis.
Adult: Total length (from basal tip of stalk to uppermost
part of tentacle membrane) up to 480 µm. Tentacle number
14 in holotype, but obscure in paratypes due to severe contraction of lophophore during fixation. Stalk as long as calyx.
No foot in adult. A pair of lateral sense organs at margin of
calyx rim. Rim of the calyx expanded, forming a disc-shaped
calyx. Several transparent granules, ca. 30 µm in diameter, in
calyx. Appendages at margin of calyx rim, back of calyx, and
883
stalk. Some of appendages irregularly long, ca. 100 µm. Appendages smaller in number and shorter in young specimens
than large individuals. Muscle fibers, running to many directions, conspicuous in stalk. Liberated bud: Total length (from
posterior tip of foot to anterior-most point of tentacular membrane between two abfrontal tentacles) 330 µm. Tentacle number 10. Stalk very short. Foot with foot groove present. A pair
of lateral sense organs present. Appendages comparable to
those in adult specimens absent.
Reproduction: Buds emerge from laterofrontal areas of
the calyx at level of the middle stomach, attaching themselves
to their mother’s body by tips of their feet. No larvae were
found.
Remarks: The most characteristic feature of the present
species lies in its possession of many appendages covering
the body surface. Although some Loxosomella species are
reported to have some appendages, such long appendages
as in the present species are reported only in L. cirrifera
(Harmer, 1915), L. circularis (Harmer, 1915), and L. velata
(Harmer, 1915). The former two of these species are distinct
from L. lappa in lacking expanded rim in calyx. The calyx rim
of L. velata is much more expanded than that of L. lappa,
reaching to the upper half of the stalk. In general shape, L.
lappa resembles L. ditadii Marcus and Marcus, 1968, but the
long appendages are not described for L. ditadii (see Marcus
and Marcus, 1968).
This species was found on glass plates immersed in a
bucket with stones collected from the reef flat. Thus, the origi-
Fig. 5. Drawings of a young specimen of Loxosomella lappa (NSMT-Ka 27, paratype) with a very small bud (b). Note that the appendages are
smaller in number and shorter than older holotype specimen (Fig. 4). A: Frontal view. B: Abfrontal view. Arrows indicate lateral sense organs.
Bars=100 µm.
Fig. 6. Liberated bud of Loxosomella lappa (NSMT-Ka 28, paratype). Drawing (A) and photograph (B) of lateral view. Arrows indicate lateral
sense organs. Bars=100 µm.
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884
T. Iseto
Fig. 7. Holotype of Loxosomella aloxiata (NSMT-Ka 29) with five buds (b) in various stages. A, B: Drawings of frontal and abfrontal views,
respectively. Thin arrows indicate tentacles bending inward after fixation. Thick arrows indicate lateral sense organs. C: Photograph of frontal
view. Bars =100 µm.
Fig. 8. Drawing (A) and photograph (B) of calyx of Loxosomella aloxiata (NSMT-Ka 31, paratype) in abfrontal view. Detritus originally covering
calyx was removed. Buds are omitted in A. Thin arrows indicate tentacles bending inward after fixation. Thick arrows indicate lateral sense
organs. Bars=100 µm.
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Three New Loxosomella from Okinawa
nal individuals are likely to have occurred on stones as components of their epifauna.
Loxosomella aloxiata sp. nov.
(Figs. 7–9)
Material examined.
Holotype: An adult (NSMT-Ka 29) on a slide glass, settled
on the reef flat for two months at a depth of ca. 1 m in Mizugama
(estuary of Hija River), Okinawa Island, Okinawa, Japan
(26°21’N, 127°45’E), collected on 5 July 2000. Paratypes: 14
adults (NSMT-Ka 30) collected with holotype specimen.
An adult (NSMT-Ka 31) on a slide glass, settled for two months
at the type locality, collected on 30 December 2000. Two liberated buds: NSMT-Ka 32, liberated from a specimen collected with holotype; NSMT-Ka 33, liberated from a specimen
collected at the type locality on 23 June 1999.
Etymology: The specific name “aloxiata” is a Greek word
meaning “not oblique”. This refers to the position of lophophore that is not oblique frontally and opens at upper most
part of the calyx, whereas the lophophores of most other solitary species lean frontally.
Diagnosis.
Adult: A large species, total length (from basal tip of stalk
to uppermost level of tentacle membrane) up to 1600 µm.
Whole body often covered with detritus except for lophophore.
Tentacle number 12 to 24. Stalk long: two to three times longer
than calyx. No foot in adult. A pair of lateral sense organs,
long enough to protrude from detritus covering, present. A
pair of lateral lobes of stomach present. A few abfrontal ten-
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tacles bending inward in fixed specimens; they expanding as
other tentacles in live specimens. Liberated buds: Total length
(from posterior tip of foot to anterior-most point of the tentacular membrane between two abfrontal tentacles) 470 µm
in NSMT-Ka 32, 400 µm in NSMT-Ka 33. Tentacle number 12
in both specimens. Stalk very short. Foot with foot groove
present. A pair of lateral sense organs present. Abfrontal part
of lophophore bending inward in fixed specimens.
Reproduction: Buds emerge from laterofrontal areas of
the calyx at level of the upper side of the stomach, attaching
themselves to their mother’s body by tips of their feet. A maximum of five buds were observed simultaneously on a single
mother. No larvae were found.
Remarks: This species resembles large individuals of L.
antarctica Franzén, 1973. The latter species, however, has a
total of 12 tentacles. Moreover, lateral sense organs are rudimentary and buds emerge from middle of the stomach in L.
antarctica (Franzén, 1973; Emschermann, 1993). Loxosomella
olei du Bois-Rymond-Marcus, 1957 also resembles L. aloxiata,
but is distinct from the latter in larger body size (up to 2600
µm long), much longer tentacles in adults, and presence of
14 tentacles in large buds (du Bois-Rymond-Marcus, 1957).
Loxosomella aloxiata is a large species, and the total
length often exceeded 1500 µm, reaching 1600 µm. Although
all of the type specimens were found on the slide glasses, this
species was also found on the stones and dead coral fragments several times. This indicates that the natural habitats
of this species include surfaces of stones and other non-living
objects. The budding of this species was easily induced by
feeding with microalgae. Nevertheless, most of the liberated
buds failed to attach themselves to the surface of culture dish,
leading long time culture in the laboratory to failure.
DISCUSSION
Fig. 9. Drawing (A) and photograph (B) of left lateral view of liberated bud of Loxosomella aloxiata (NSMT-Ka 33, paratype). Thick and
thin arrows indicate lateral sense organ and a part of lophophore bending inward after the fixation, respectively. Bars =100 µm.
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This is the first report on Entoprocta from the Ryukyu
Archipelago. Even in other parts of the world, the entoprocts
in coral reefs have not been well studied taxonomically, and it
is thus not surprising that all specimens examined here represented undescribed species.
Nielsen (1964) divided the genus Loxosomella (sensu
lato) into two subgenera, Loxosomella and Loxomitra, on the
basis of differential budding modes: buds of the nominotypical
subgenus are attached to their mother’s body by the tip of
their feet, whereas those of the subgenus Loxomitra by
the back of stalks or calyxes. Soule et al. (1987) elevated the
subgenus Loxomitra to the rank of full genus. I agree with this
change, but since there are no other characters to distinguish
these two genera, observation of budding modes is essential
for a correct generic assignment of a given species. Therefore, if the specimen has no buds, culturing is required to
induce budding. In the three species described here, buds
were invariably attached to their mother’s body by foot tips,
and it is thus obvious that all these species belong to the
genus Loxosomella (sensu stricto).
It is uncommon to collect entoprocts using settlement
886
T. Iseto
panels. Ryland and Austin (1960) immersed panels in a
dock in Swansea, U. K., and found Loxomitra kefersteinii
(Claparéde, 1867) and Loxosomella antedonis Mortensen,
1911 attached on the panels some time later, and, to my knowledge, this is the only case of application of this method for the
sampling of entoprocts in the field. In the present study,
Loxosomella aloxiata was found abundantly on settled slide
glasses. It is probable that this method enables us to discover
additional undescribed species in future studies.
Solitary entoprocts are usually known to inhabit on other
animals as epizoics or commensals (Hyman, 1951; Ruppert
and Barnes, 1994), and a variety of animal groups (e. g., polychaetes, sipunculans, bryozoans, and sponges) are known to
be their hosts (summarized in Nielsen, 1964; Soule and Soule,
1965). Only a few solitary entoprocts have been found from
non-living objects. In Loxosomella, for example, only L. olei
du Bois-Reymond-Marcus, 1957 lives on stones (du BoisReymond-Marcus, 1957), and L. hispida Marcus and Marcus,
1968 and L. shizugawaense (Toriumi, 1949) on algae (Toriumi,
1949; Marcus and Marcus, 1968; Konno, 1971). Furthermore,
only one species, L. isolata Salvini-Plawen, 1968, is known to
be non-epizoic among 25 species of the genus Loxosoma
(Salvini-Plawen, 1968, 1986; Nielsen, 1996), and in Loxomitra,
only L. kefersteinii (Claparéde, 1867) is found on non-living
substrata, as well as on other animals (Claparéde, 1867;
Nitsche, 1875; Ryland and Austin, 1960; Nielsen, 1966). In
this regard, the present species seem to be rather exceptional,
because all of them were found only on non-living substrata. I
examined some of the host candidates, such as polychaetes,
sipunculans, and sponges collected from the type localities of
the present species, but any solitary entoprocts were not found
on them.
Nielsen (1964) suggested that the solitary entoprocts not
associated with other animals “may hardly be regarded as
quite natural”, because they were found under more or less
artificial conditions, such as in aquaria or on settlement plates.
However, because Loxosomella monocera and L. aloxiata
were found on stones and shell remains directly collected from
the reef flat, they probably have no host animals even in
nature. Although it is possible that these species inhabit not
only on non-living substrata but also on other animals, the
non-living substrata seem to be common habitats for solitary
entoprocts at least in the Ryukyus. It is probable that solitary
entoprocts without host animals will be also found in shallow
waters in other tropical areas.
ACKNOWLEDGEMENTS
I wish to express my gratitude to Emeritus Professor Mayumi
Yamada (Hokkaido University), Professor Shunsuke F. Mawatari
(Hokkaido University), Professor Teruaki Nishikawa (Nagoya University), and Dr. Euichi Hirose (University of the Ryukyus), for their valuable advice and encouragements. Thanks are also due to Hidetoshi
Ota (University of the Ryukyus) for his critical reading of the manuscript and helpful suggestions on the zoological nomenclature. This
study was supported in part by the Sasagawa Scientific Research
Grant from The Japan Science Society and a subvention to graduate
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students by Research Institute of Marine Invertebrates.
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(Received March 1, 2001 / Accepted May 10, 2001)
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