<ooloyzca/ Journal of the Lznnean ,EociPly (1983'1,77: 175-187. With 1 figure
An examination of the Phascolosoma subgenera
Ant illesoma, Rueppellisoma and Sat onus
(Sipuncula)
E. B. CUTLER
AND
N. J. CUTLER
LJtica College of Syracuse Uniuersip, lJtica, J e w York 13502, U.S.A.
.Icreptpdfnr publication July 1982
Three of the subgenera in the sipunculan genus Phascolosuma are reviewed, based on examination of
type material. The presumed difference between the subgenera Antillemma and Rueppellisoma (number
ofretractor muscles) is shown to be invalid and the taxa are therefore synonymous. T h e 15 species are
either consideredjunior synonyms o f f . antillarum (eight), transferred to other species or genera (six), or
considered inccrtae srdis (one).Phascolotnma antillarum is redescribed. Of the eight species in thc subgenus
Satonlo, only P . pectinaturn remains valid: the others are considered either to belong to other taxa (four)
or to be incertae m i i s (three).
KEY WORDS:- Sipuncula
-
Pllascoiosoma
.I\ntillesoma
Rueppellisoma
Satonus.
~
CONTENTS
Introduction .
. . . . . . . .
Subgenus Antillesoma Stephen & Edmonds, 1972
Phascolosoma antillarum Grube & Oersted, 1858 .
Subgenus Satonus Stephen & Edmonds, 1972
.
Acknowledgements.
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References.
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175
176
182
184
186
187
INTRODUCTION
Stephen & Edmonds (1972) divided the genus Phascolosoma into four new
subgenera-Phascolosoma s.s., Satonus, Antillesoma and Rueppellisoma-based largely
on information in the literature. However, examination of type material and
specimens of representative taxa have led us to believe the criteria used for this
separation were erroneous.
This paper deals with only the subgenera Antillesoma, Rueppellisoma and Satonus.
The species names considered herein are those found in Stephen & Edmonds
(1972) without consideration of changes made before or after that work except for
two new species described by Murina (1975). The literature section for each
species contains only those citations which relate to original collections, not
revisionary or review papers. Proposed taxonomic changes are summarized in
Table 1.
0024-4082/83/020175+ 13f03.00/0
175
0 1983 The Linnean Society of London
E. B. CUTLER AND N. J . CUTLER
176
Table 1. Summary of proposed taxonomic changes
Original name
Pharcolosoma (Antillemna) antillarum Grube & Oersted, 1858
Phascolosoma (Antillesoma) asser (Selenka, DeMan & Bulow, 1883)
Phascolosoma (Antillesoma) horsti (ten Broeke, 1925)
Phascolosoma (Antillesoma) microdentigerum (ten Broeke, 1925)
Phascolosoma (Antillesoma) minutum (ten Broeke, 1925)
Phascolosoma (Antillesoma) pelmum (Selenka et al., 1883)
Phascolosoma (Antillesoma) pelmum sensu Fischer, 1922
Phascolosoma (ilntillesoma) pelmum sensu Cutler, 1977
Phascolosoma (Antillesoma) schmidti Murina, 1975
Phascolosoma (Rueppel~isomu)gaudens (Lanchester, 1905)
Pha.rcolosoma (Rueppelli~oma)golikoui Murina, 1975
Phascolosoma (Rueppellisoma) nahaense (Ikeda, 1904)
Pha.rcolosoma (Ru~ppelli.ioma)onomichianum (Ikeda, 1904)
Pha.ha.lci)losoma(Rueppellisomni rurppelii Grube, 1868
Phascolotoma iRurppdlnsomar ruepprlii sensu Stephen, 1941
Phascoloioma (Rueppdlisoma) seu’elli (Stephen, 1941)
Phascolosoma (Rueppellisoma) simili (Chen & Yeh, 1958)
Phascolosoma (Rueppellisoma) weldoni (Shipley , 1892)
Phascolosoma (Satonus) demanni (Sluiter, 1891)
Phascolosoma (Satonus) duplicigranulatum (Sluiter, 1896)
Phascnlosoma (Satonw)fhlcidentatum (Sluiter, 1882)
Phascolosoma (Satonus) hebes (Sluiter, 1902)
Phascolosoma (Satonw) maculatum (Sluiter, 1886)
Phascolosoma (Satonus) mauritaniense (Herubel, 1924)
Phascolosoma (Satonw) ni.grztorquatum (Sluiter, 1882)
Phascolosoma (Satonus) pectinatum (Keferstein, 1867)
Proposed status
PhaJcolosoma (Antillesoma) antillarum
Phascolosoma (Antillesoma) antillarm
Phascolosoma (Phascolosoma) nigrescens
Phascolosoma (Phascolosoma) perlucens
Phascolosoma ( Phamlosoma ) nigrescens
Phascolosoma (Antillesoma) a n t i l l a m
ThemiSte sp.
Phascolosoma (Phascolosoma) scolops
Phascolosoma (Antillesoma) antillarum
Phascolosoma (Antillesoma) antillarum
Phascolosoma (Phascolosoma) japonicum
Phascolosoma (Phascolosoma) scolops
Phascolosoma (Antillesoma) antillarum
incertae sedis
Phascolosoma (Phascolosoma) scolops
GolJingia (Nephasoma sp.)
Phascolosoma (Antillesoma) antillarum
Phascolosoma (Antillesoma) antillarum
incertae sedis
Phascolosoma (Phascolosoma) cf. nigrescens
zncertae sedis
Siphonosoma (Hesperosiphon) cumanenst
Phascolosoma (Phascolosoma) sp.
incertae sedis
Phascolosoma (Phascolosoma) cf. scolops
Phascolosoma (Satonus) pectinatum
SUBGENUS .4 1 IILLESO,Z-IA STEPHEN & EDMONDS, 1972
Phascolosoma su bg . Rueppellisoma syn. nov.
Diagnosis: Trunk with conspicuous papillae. Longitudinal musculature divided
into separate bands which anastomose and are not always visible through the
epidermis. Introvert hooks absent and tentacles which are arranged around the
nuchal organ dorsal to the mouth numerous (more than 30 in adults). Four
retractor muscles present but there may be extensive fusion between the two
muscles on either side giving the appearance of a single pair with a notch or split in
the broad origin. The spindle muscle is attached posteriorly and contractile vessel
carries many digitiform villi.
Remarks: The single character Stephen & Edmonds (1972) used to separate the two
subgenera Antillesoma and Rueppellisoma was the number of introvert retractor
muscles: two pairs verses one pair. In most sipunculans this situation is clear and
unambiguous but not in this complex. The high degree of fusion along much of the
length of each pair (right and left) makes it appear in some individuals as though
there is only one muscle on each side, each with a small fracture near the origin
(base). Discussing this issue in Phascolosoma antillarum, Fisher (1952) stated : “The
two muscles of each side soon unite. I n a strongly contracted specimen there is
hardly any separation and the animal appears to have only two retractors arising
near the posterior end of the body. T h e degree of separation is like that of
Phascolosoma asser (Selenka, Deman & Bulow, 1883, fig. 97).” Fischer (1914) has
also remarked on the unclear nature of this feature.
EXAMINATION OF PHASCOLOSOMA4 SUBGENERA
177
Figure 1. Phascolosoma antillarum retractor muscles. A, Specimen showing distinctly separated dorsal
and ventral retractors. B, A specimen with very little separation (indistinct) between dorsal and ventral
origins. C, Th e origin of the left pair of retractor muscles in an individual showing three subdivisions in
the ventral muscle and very little separation between the dorsal and ventral muscles (see text for
discussion).
The degree of fusionlseparation of these muscles is quite variable (Fig. 1). Rice
(pers. commn) has suggested that one should look for the presence of gonads which
are found along the bases of the ventral retractors only: this is a helpful feature
when the worms are in the proliferative phase of the gametogenic cycle. This
variability evidently led to erroneous interpretations by a few earlier workers
including Cutler & Cutler (1981). T h e reality of one pair of retractors has never
been critically examined but was used to erect the subgenus Rueppellisoma, said to
have only a ventral pair of muscles. We consider this an invalid separation and that
the subgenera Antillesoma and Rueppellisoma are synonymous, Rueppellisoma being
junior since based on a later published species.
Confusion may also result from an ontogenetic anomaly in which the loss (nondevelopment or fusion) of one pair of retractor muscles in a few individuals occurs.
Species erected on the basis of one or two specimens which are identical to a
common sympatric population except in having only one pair of retractor muscles
are particularly suspect.
Of the seven species originally designated to the subgenus Antitlesoma, four (P.
antillarum, P. asser, P. pelmum and P. schmidti) lack hooks and three (P.
microdentigerum, P. minutum and P. horsti) do not, but all are said to have contractile
vessel villi or tubules. T h e three species with hooks in fact do not have villi. What
ten Broeke (1925) was referring to as small villi are not true villi but bulbous
vesicles on the contractile vessel. These three species are clearly members of the
subgenus Phascolosoma and are not distinct entities.
The difficulty in distinguishing between P. pelmum and P. asser is evident in
Stephen & Edmonds (1972 : 278,282). Our comparisons of these two with each
other and with P. antillarum has led us to conclude that there are no significant
differences. T h e number of rows of platelets on the trunk papillae, number of
longitudinal muscle bands or intestinal coils is neither species-specific nor
biologically meaningful. For similar reasons P. schmidti (a single worm) is
considered conspecific with P. antillarum.
Of the eight species originally in the subgenus ~ u e ~ p e ~ ~ i s three
o m a , (P. gotikovi, P.
nahaense and P. rueppelii) have hooks. Phascolosoma nahaense was determined to be
conspecific with P. scolops (Cutler & Cutler, 1981). Using the same logic we
17H
E. B. CUTLER AND N. J. CUTLER
consider P . golikovi a junior synonym of P.japonicum. The type of P . ruejpelii cannot
be located and the only museum specimens we found were P. scolops, lacking villi.
Therefore, i t is appropriately designated incertae sedi.r. The holotype of P. sewelli
lacks longitudinal muscle bands and in other respects clearly belongs to Goljingia
(.Nephasoma).The remaining four species (P. gaudens, P. onomichianum, P. simile and
P. meldoni) are indistinguishable from one another or from P. antillarum and are
therefore considered conspecific. The status of each of these species will now be
discussed in more detail.
Phascolosoma asser (Selenka et al., 1883)
Ph_ymosoma asser Selenka et al., 1883: 59 60, pl. 1, fig. 3, pl. 7, figs. 97 -101; Sluiter,
1886:503; 1891: 118.
Physcosoma asser: Fischer, 1895: 12; 1914: 5 ; 1922: 10; Sluiter, 1902: 11; Shipley,
1903: 133, 174; Leroy, 1936: 424.
Matprial examined: Stockholm-Fischer’s from Timor, Java ( # 1%). Edinburghunpublished material from Loyalty Island, New Caledonia ( # 1959.55.11).
Hamburg-Fischer’s from China (V-8843).
The type specimen of Selenka et al. (1883) had been in Berlin but had dried up
and been discarded. All of the other specimens we examined are indistinguishable
from Phascolosoma antitlarum.
Phascolosoma horsti (ten Broeke, 1925)
Physcosoma horsti ten Broeke, 1925: 89, text-fig. 1 1.
Material examined: Amsterdam-type
of ten Broeke (1925) (V.Si. 108), five
specimens from the West Indies.
These five worms have contractile vessels that are bulbous and folded but lack
true villi. The hooks have the internal structure of Phascolosoma nigrescens and
strongly resemble that species in other ways. We suggest that P. horsti should be
treated as a junior synonym of P. nigrescens.
Phascolosoma microdentigerum (ten Broeke, 1925)
P h p m o m a microdentigerurn ten Broeke, 1925: 88 -89, text-figs 8-10; Stephen, 1960:
517 518.
Material examined: Amsterdam - type of ten Broeke (V.Si. 11 1) and eight other
specimens from Curacao.
This material is in poor condition; it does not exhibit villi on the contractile
vessel but simply bulbous vesicles. As pointed out by Stephen (1960) ten Broeke
divided this species from P . dentigerum ( =P. perlucens) by rather small differences
and “an examination of comparative material might show that in reality these two
species should be combined”. Rice (1975 :37) also observed, “Specimens in my
collections from Spaansche Water, Curacao, in close proximity to the type locality
of P. microdentigerum could not be distinguished from P. perlucens found at other
localities throughout the Caribbean region”. We propose that P. microdentigerum be
considered conspecific with P. perlucens.
EXAMINATION OF PH,4SCOLOSOMA SUBGENER/1
179
Phascolosoma minutum (ten Broeke, 1925)
Physcosoma minutum ten Broeke, 1925: 87-88, test-figs 6-7
minutum Keferstein, 1863 : 40 = GolJingia minuta (Kef.)).
(not Phascolosoma
Material examined: Amsterdam-type of ten Broeke (V.Si. 117) from Curasao.
This single, incomplete, damaged worm lacks true contractile vessel villi. T h e
hooks and other features strongly resemble a small Phascolosoma nigrescens. We
propose that P. minutum be treated as a junior synonym of P. nigrescens.
Phascolosoma pelmum (Selenka et al., 1883)
Phymosoma pelma Selenka et al., 1883: 60, pi. 1, fig. 4, pl. 7, fig. 102; Sluiter, 1886:
504; 1891 : 118; Augener, 1903: 31 1-312.
Physcosoma pelma: Sluiter, 1902: 12; Shipley, 1902: 134; Fischer, 1922: 15; 1923:
23-24; Sato, 1935: 311-312, pl. 4, fig. 11.
Phascolosoma pelmum: Cutler, 1977: 150, fig. 11.
Material examined: Leiden-two specimens identified as type (probably co-type) of
Selenka et al. from Java. Sendai-three specimens identified by Sato (#3-15).
Stockholm-Fischer’s
specimen from Java ( # 1 12). Copenhagen-Cutler’s
Galathea Sts. 31 1, 452,454 from Makassar and Ganges; also unpublished material
collected by Th. Mortensen from Koh Chang on the Thailand/Cambodia border.
Selenka et al. (1883, pl. 7) illustrate papillae of P. pelmum, P. asser, and P.
antillarum. It is the arrangement of platelets on these papillae which they used to
differentiate these species. Our observations suggest that the minor differences in
these epidermal structures are insufficient grounds for establishing separate species.
The overwhelming similarities far outweigh these presumed differences.
The worm collected by Fischer (Stockholm # 112) is partially dried but clearly
is a Themiste not a Phascolosoma. Cutler’s Galathea worms are really Phascolosoma
scolops. The remainder of the specimens do belong to this complex but, with our
present appreciation of the normal variation in external appearance within this
population, they fit easily into the present concept of P. antillarum; therefore P.
pelmum becomes a junior synonym of P. antillarum.
Phascolosoma schmidti Murina, 1975
Phascolosoma schmidti Murina, 1975: 57 -59, fig. 3.
Material examined: None.
This single, 75 mm worm with a 60 mm introvert from Okinawa is housed in the
Leningrad Museum but is unavailable for examination. Murina compared it to P.
antillarum, P . asser and P . pelmum and said it differed from these by the degree of
separation of the retractor muscles, having three (not one) fixing muscles, shorter
nephridia attached for only one-quarter their length, and longitudinal muscle
bands which do not anastomose.
The large size of this individual is striking (others in this group are commonly
15-30 mm) and may well account for the differences. The nature of these
differences in this one worm are, from our experience, probably not indicative of a
reproductively isolated population (species) but rather at one end of a continuum
within a very plastic species. A recent collection of 118 members of this genus from
180
E. B. CUTLER AND N. J . CUTLER
Okinawa did not include any animals of this size (Cutler & Cutler, 1981). If other
specimens can be collected, a subspecific rank might be warranted but we consider
this to be a large, atypical member of and a junior synonym to P. antillarum.
Phascolosoma (Rueppellisoma) gaudens (Lanchester, 1905)
Physcosoma gaudens Lanchester, 1905: 38, pl. 2, fig. 1 1.
Material examined: London-Lanchester’s type; three specimens from Malay.
These three specimens are small (about 15 mm) and have been dried out at
some point. One dissected specimen clearly shows the two separate retractor
muscles on the right side but on the left side they appear almost totally fused. These
are all clearly referable to P . antillarum.
Phascolosoma (Rueppellisoma) golikovi Murina, 1975
Phascolosoma golikovi Murina, 1975 : 54-55, fig. 1.
Material examined: None.
This single worm has a body less than 3 mm long and is clearly a juvenile
specimen. The type was unavailable for examination so our conclusions are based
on the published description. Murina compared this species (individual) to P.
nahaense and P . rueppelii; it differed from both in the form of the hook. However, it is
also said to lack a posteriorly attached spindle muscle which would suggest it might
belong in the subgenus Satonus.
Since it apparently has only two retractor muscles, Murina did not compare it to
P. japonicum, a very common species in that region (2-4 m in Sea of Japan). The
anomalous, occasional loss/fusion of retractor muscles has been discussed elsewhere
(Gibbs, 1973; Cutler & Cutler, 1981). As noted in the section on Satonus, the
apparent absence of a posteriorly attached spindle muscle, especially in one very
small worm, may not be a solid basis for erecting a new species. To base a new
species on such an individual which in all other ways is like P.japonicum seems
imprudent. Our analysis suggests that P. golikovi should be considered a junior
synonym of P. japonicum.
Phascolosoma (Rueppellisoma) nahaense (Ikeda, 1904)
Phymosoma nahaense Ikeda, 1904: 29-31, text-fig. 8, 59-62.
Phascolosoma nahaense: Cutler & Cutler, 1981 : 85-86.
Material examined: None.
Ikeda’s two specimens from Naha, Okinawa have been lost. Cutler & Cutler
(1981) discussed this species in their review of Ikeda and Sato’s species and
concluded that i t should be treated as two Phascolosoma scolops with aberrant
retractor muscles (one pair failing to develop or having fused with the other as
happens in some populations). We herein reaffirm that conclusion, i.e. P. nahaense
is a junior synonym of P . scolops.
EXAXfIUATION OF PHAS(.OLOSOhIA SUBGENERA
181
Phascolosoma (Rueppellisoma) onomichianum (Ikeda, 1904)
Phymosoma onomichianum Ikeda, 1904: 26-28, text-fig. 7, 56-58.
Physcosoma onomichianum Sato, 1934: 247; 1939: 397-398, pl. 22, fig. 14, pl. 23, fig.
43.
Phascolosoma onomichianum: Wesenberg-Lund, 1959 : 67 ; 1963: 127-1 28 ; Murina,
1967: 44; Cutler & Cutler, 1981: 86-88.
Material examined: Japan-eight
specimens collected at Onomichi by Cutler &
Cutler, 1981. Tokyo-one specimen identified by Ikeda from Itoman, Okinawa.
Sendai-many identified by Sato (#2-15, 2-21, 2-28, 3-5, 7-24, 8-13, 8-15).
Copenhagen-one specimen from Mauritius in Wesenberg-Lund ( 1959).
This species was discussed at length in an earlier paper wherein the retractor
muscles were mistakenly interpreted as a ‘single pair’. “Each of the muscles is
subdivided near its origin from the trunk wall into two short roots” (Cutler &
Cutler, 1981 :87). The similarity to other species in this complex was pointed to,
along with the need for a careful re-examination. Wesenberg-Lund (1959) also
alluded to the uncertainties here. One action, overlooked by subsequent authors
including ourselves, was that of Fischer ( 1914) wherein he placed P. onomichianum
in the synonymy of P. asser. This observation is particularly interesting in this
context as he referred to Ikeda’s description and noted that the only apparent
difference was the two retractors. He concluded that this may have been the result
of abnormal fusion of the original four retractor muscles or from Ikeda having
missed the significance of the division near the base. Fischer also notes Selenka’s
comment about P. asser that the small dorsal easily unites with the ventral muscle.
It is now clear to us that P. onomichianum is conspecific with P. antillarum which also
includes P. asser.
Phascolosoma (Rueppellisoma) rueppelii Grube, 1868
Phascolosoma rueppelii Grube, 1868: 643, pl. 8, fig. 2 ; Wesenberg-Lund, 1957 : 12.
Phymosoma rueppellii: Selenka et al., 1883: 82-83.
Pttyscosoma rueppellii: Shipley, 1902 : 135; Stephen, 1941 : 406-407.
Material examined: London-Stephen’s six specimens from the Murray Expedition
off Arabia. Edinburgh-seven specimens identified by Stephen from Somalia but
never published ( # 1958.23.107 & 108).
This particular species has not survived well in that most of the collections listed
above have been lost, including the type. Both of Stephen’s collections appear to be
small Phascolosoma scolops. In view of these data we are hereby suggesting this name
3e placed in incertae sedis. If the type is located, we suspect it can probably be
referred to P. scolops.
Phascolosoma (Rueppellisoma) sewelli (Stephen, 1941)
”hyscosoma sewelli Stephen, 1941 : 405-407.
Material examined: London-Stephen’s type from Gulf of Oman ; one specimen
iom Maldive Islands.
There are two specimens bearing this name. The designated type, with a 60 mm
runk is in good condition (St. 59) but the second worm has dried out (St. 162).
I82
E. B. CUTLER AND N. J . CUTLER
Neither worm has longitudinal muscle bands (the dried one exhibits some fractures
in this layer). The spindle muscle is unattached posteriorly, the contractile vessel is
without villi, and in all respects these are clearly referable to the subgenus Goljngia
(. Vephasoma).
Phascolosoma (RueFpellisoma) simile (Chen & Yeh, 1958)
Physcosoma similis Chen & Yeh, 1958: 274-276, text-figs 3, 4.
Material examined: None.
IVhile we have been unable to locate this material, we are nevertheless, based on
a comparison of the descriptions and distributions, treating this species as
conspecific with Phascolosoma antillarum.
Phascolosoma (Rueppellisoma) weldonii (Shipley, 1892)
Phpcosoma weldonii Shipley, 1892: 77 -78.
Material examined: London-Shipley’s
type from Bahamas. C a m b r i d g e t w o
specimens which we believe to be co-types. Paris-one specimen with this name
but no other datum in vial (V-26).
Shipley’s three worms are in good condition and exhibit two pairs of retractor
muscles that quickly join to form a single muscle on each side. There are 17 34
longitudinal muscle bands, not 10-12 as Shipley stated. These three worms are
conspecific with Phascolosoma antillarum. The specimen found in Paris had no data
attached to it except this name. It is clearly a Themiste.
PHASCOLOSOMA A.NTII.LAR1JM GRUBE & OERSTED, 1858
Phascolosoma antillarum Grube & Oersted, 1858: 117- 1 18; Diesing, 1859 : 762 ;
Keferstein, 1863; 40, pl. 3, figs 2, 1 1 ; 1865: 435, pl. 31, fig. 11, pl. 33, fig. 37;
Ikeda, 1904: 24-25; Fisher, 1952: 434-436, pl. 39, figs 8, 9 ; Longhurst, 1958:
85; Rice, 1975: 42; Rice & McIntyre, 1972: 42; Cutler, 1977: 150; Tarifeno &
Rojas, 1978: 119-121, fig. 13.
Phymosoma antillarum: Selenka et al. 1883: 57, pl. 7, figs 93-96; Fischer, 1895: 12.
Phycosoma antillarum: Gerould, 1913: 420-421, pl. 62, figs 19,20; Fischer, 1922: 11 ;
Leroy, 1936: 424; Sato, 1939: 394.
Phascolosoma fuscum Keferstein, 1862 : 67.
Phascolosoma nigriceps Baird, 1868: 90, pl. 11, figs l-1A.
Phascolosoma aethiops Baird, 1868: 90.
Yhymosoma asser Selenka et al., 1883: 59-60, pl. 1, fig. 3, pl. 7, figs 97-101 ; Sluiter,
1886:503; 1891: 118.
Physcosoma asser Fischer, 1895: 12; 1914: 5; 1922: 10; Sluiter, 1902: 11; Shipley,
1903: 133, 174; Leroy, 1936: 424.
Phymosoma pelma Selenka, et al. 1883: 60, pl. 1, fig. 4, pl. 7, fig. 102; Sluiter, 886 :
504; 1891 : 118; Augener, 1903: 31 1-312.
Physcosoma pelma Sluiter, 1902: 12; Shipley, 1902: 134; Fischer, 1922: 15; 923 :
23-24; Sato, 1935: 311-312, pl. 4, fig. 11.
Phascolosoma pelmum Cutler, 1977: 150, fig. 11.
Physcosoma weldonii Shipley, 1892 : 77-78
EXAMINATION OF PHASCOLOSOMA SUBGENERA
I83
PhFosoma onomichianum Ikeda, 1904: 2&28, text-figs 7, 56-58.
Physcosoma onomichianum Sato, 1934: 247; 1939: 397-398, pl. 22., fig. 14, pl. 23, fig.
43.
Phascolosoma onomichianum Wesenberg-Lund, 1959: 67 ; 1963: 127-1 28 ; Murina,
1967: 44; Cutler & Cutler, 1981 : 86-88.
Physcosoma gaudens Lanchester, 1905: 38, pl. 2, fig. 1 1.
Physcosoma similis Chen & Yeh, 1958: 274-276, text-figs 3-4.
Phascolosoma schmidti Murina, 1975: 57-59, fig. 3.
Material examined: Berlin-Grube’s from Barbados ( #6058), St. Croix ( # 1015)
and Puntarenas ( # 1014). What is listed as type ( # 1011, 1012 & 1013) are in
catalogue but specimens are missing. Stockholm-Fischer’s from St. Thomas
( # 4 4 ) and Pacific side of Panama (#30, 132 & 178). Hamburg--211
1 from
West Indies (no other data). Edinburgh-one specimen from Gold Coast
( # 1948.23.1310) ; two specimens from Inhaca (=Phascolosoma cJ stephensoni
unpubl.) ( # 1960.48.8). London-Baird’s type of P. aethiops from St. Vincent and
P. nigriceps from St. Thomas. Washington D.C.-Gerould’s
specimens from
Florida.
This name is the senior synonym for this entire complex. The species is
redescribed below.
Description
This species commonly has a trunk 15-30 mm long but may be up to 85 mm and
a width equal to 1&25y0 of the length. In living animals the introvert is 65-75OL,
of the trunk length but in preserved material it may appear to be only 2(r25°/0 as
long as the trunk. The trunk is pale yellow-brown and bears many large, dark
papillae especially large and crowded around the anterior end. The introvert also
bears papillae on its proximal portion but the distal part is smooth and white. This
zone is marked off by a distinctive collar. O n the distal tip are numerous (3&200)
digitiform tentacles surrounding the dorsal nuchal organ. The number of tentacles
is correlated with size (Cutler & Cutler, 1981 : 86) and they have a violet (brown
when preserved) pigment distributed in patches or stripes along the tentacles. This
pigment usually extends onto the area around the nuchal organ. Hooks are absent.
Internally the longitudinal musculature is divided into numerous anastomosing
bundles. The four retractor muscles originate about 60-70y0 of the distance to the
posterior end of the trunk. The origin of the dorsal pair is very close to the ventrals,
and they usually remain separate for only a very short distance (1&20y0 of their
length). In some situations this appears to be a single pair, each having two large
roots. The contractile vessel is covered with many tubular villi along its length, the
large ones bifurcating or branching at their tips. The intestinal coil is anchored at
both ends by the spindle muscle and a small rectal caecum is usually present but
not often evident at first glance. The pair of nephridia open just posterior to the
anus, are about 4&50% of the trunk length (shorter in larger worms) and attach
by a connective tissue membrane to the body wall for 75-90y0 of their length
(occasionally less).
Distribution
In the Western Atlantic this has been reported from Jamaica, the Bahamas, Key
West, Florida, Cuba, several West Indies Islands, Brazil, Venezuela and Panama.
12
184
E. B. CUTLER AND N . J . CUTLER
In the Eastern Atlantic it has been found at Sierra Leone and the Gold Coast, then
into the Indian Ocean at Durban, South Africa, Mozambique, Madagascar,
Mauritius, Maldive and Laccadive Islands and Ceylon. I n the Indo-West Pacific
it is reported from Indonesia, Malaysia, Philippines, Korea, Formosa, China and
southern Japan, then out to New Caledonia, the West Caroline Islands and
Hawaii. In the eastern Pacific it has been recorded from Panama, Costa Rica (not
Chile) and Baja, California. I n summary, this is a cosmopolitan species found in
tropical and subtropical, shallow water.
Remarks
One point that has been confusing over the years is whether or not this species
has ever been collected in Chile. The basis for that is the place name Puntarenas as
in Grube & Oersted (1858). This was later changed to Punta Arenas by other
authors. The former (original) is in Costa Rica; the second is in Chile. It is clear
that the Chile location is a m i s t a k e i t has never been found that far south
(Tarifeno & Rojas, 1978).
SUBGENUS SATO.VUS, STEPHEN & EDMONDS, 1972
I n general, we found that the original material had been inadequately described
or that the presence of a posteriorly attached spindle muscle has been overlooked.
Those original errors had never been corrected and therefore the foundation of this
taxon is very different from what Stephen & Edmonds (1972) supposed. Six of
these species were described by Sluiter ; Herubel and Keferstein each described
one.
The species originally designated as subgenus Satonus will be discussed, their
proposed status being summarized in Table 1. We discuss the one species without
hooks first; then those with hooks.
This analysis does not include PhaJcolosoma (Satonus)pectinatum (Keferstein), the
only species which may fit the definition of this taxon and which is represented by
numerous specimens in several collections. A separate paper will address this
problem.
Phascolosoma (Satonus) hebes (Sluiter, 1902)
Physcosoma hebes Sluiter, 1902: 13-14, pl. 1, figs 5, 6.
type (V. Si. 128/4).
This single worm is incomplete, the posterior end is missing and the circular
muscles have contracted to partially close off that damaged end. The gut, which
was described as being in loops, not coils, is clearly only the anterior one-third of
the gut and therefore not in its normal coil. There are dissepiments and skin bodies
present, and there is no doubt that this is a specimen of Szphonosoma cumanense with
its posterior two-thirds missing, not a Phascolosoma of any type. Therefore, this
name is placed in synonymy with Siphonosoma cumanense.
Material examined: Amsterdam-Sluiter's
Phascolosoma (Satonus) demanni (Sluiter, 1891)
Physcosoma demanni Sluiter, 1891 : 121-122, pl. 2, figs 15, 16.
EXAMINATION OF PH.~lSCOLOSO.UA SUBGENERA
185
Material examined: Amsterdam-Sluiter’s type (V. Si. 99).
This single specimen is hardened as the result of having dried out at some point.
It has been dissected and the intestinal coil is not all there. The hooks have an
internal structure like Phascolosoma nigrescens but given the poor condition of the one
and only specimen it is impossible to verify the true nature of the spindle muscle. It
is our judgement that P. demanni must be considered incertae sedis.
Phascolosoma (Satonus) duplicigranulatum (Sluiter, 1886)
Phymosoma duplicigranulatum Sluiter, 1886: 501-502.
Physcosoma duplicigranulatum: Shipley, 1899: 155; Sluiter, 1902: 13.
Material examined: Amsterdam-SIuiter’s
type (V.Si. 102). London-one
specimen, possible co-type of Sluiter’s. Cambridge-two specimens identified by
Shiple y .
Both of Sluiter’s worms have been damaged and the one in London was allowed
to partially dry out at some time. One can still see in the Amsterdam worm the
broken spindle muscle attached to the posterior end of the body. The hooks in this
material very strongly resemble Phascolosoma nigrescens. Shipley’s two worms had
not been opened. Both show an intact, normal, posteriorly attached spindle
muscle. All of these are clearly referable to the subgenus Phascolosoma sensu strzcto
and probably are not distinct species.
Phascolosoma (Satonus)falcidentatum (Sluiter, 1882)
Phymosoma falcidentatus Sluiter, 1882: 150, pl. 1, figs 2, 7, 12; 1891: 117.
Physcosoma falcidentatus: Sluiter, 1902 : 13.
Material examined: None.
A search of the collections in Amsterdam where the rest of Sluiter’s specimens
from that report are housed proved futile. It is evident that this single worm has
been lost and therefore this species will be placed in incertae sedis.
Phascolosoma (Satonus) maculatum (Sluiter, 1886)
Phymosoma maculatum Sluiter, 1886: 51 1-512, pl. 4,fig. 4; 1891; 118-1 19; Augener,
1903 308-3 10.
Physcosoma maculatum: Sluiter, 1902 : 1 1.
Material examined: Amsterdam-Sluiter’s type, two specimens (V.Si. 176 & 1 10).
These two worms are generally in good condition but # 110 is highly contracted
and its intestinal coil is missing. There is a white thread-like muscle from the
posterior end which is probably the remnant of the posteriorly attached spindle
muscle. V.Si. 176 is similar in that it appears to have a broken spindle muscle; one
that had been previously attached posteriorly.
On the basis of these observations it seems appropriate to move this species into
the subgenus Phascolosoma sensu stricto. Whether or not it represents a unique species
needs to be determined.
186
E. B. C U l L E R AND N. J . CUTLER
Phascolosoma (Satonus) mauritaniense (Herubel, 1924)
Physcosoma mauritaniense Herubel, 1924: 110-1 1 1, text-fig. 4.
Material examined: Paris-one specimen (V-2 1) named by Herubel.
Herubel based his description on two worms. However, the larger worm which
he described is now lost. What is in the museum is a 10 mm worm which has been
partly dried up, the posterior end is missing, the gut is protruding through the
body wall, and it had never been dissected. Its value as a reference specimen is nil
despite the fact that it may technically qualify as a paratype. It is impossible to
discern anything about its internal anatomy. No other specimen has been reported.
Given the above situation it seems best to move this species into incertae sedis as
there is little hope of ever being able to determine with certainty the nature of this
species.
Phascolosoma (Satonus) n i g r i ~ ~ r q u a t(Sluiter,
~m
1882)
Phymosoma nigritorquatum Sluiter, 1882: 151-152, pl. 1, figs 3, 8, 11.
Physcosoma nigritorquatum: Fischer, 1919: 280; 1921 : 4-5, figs 1, 2.
Material examined: Amsterdam-Sluiter’s
type, four specimens (V.Si. 80).
Hamburg-Fischer’s specimen from Shark’s Bay (V-9116).
None of these worms is in good condition, since the intestinal coil and other
internal organs have seriously deteriorated. Sluiter’s worms exhibit a poorly
defined remnant of a spindle muscle at the posterior end. Fischer’s worm has a well
defined posteriorly attached spindle muscle.
Edmonds (1980 : 61) discusses this species in his Australia paper. H e has not
found anything matching Sluiter’s description in recent years. It seems clear that
Fischer’s worms belong to the subgenus Phascolosoma sensu strict0 but Sluiter’s
material is in a condition that makes a positive statement impossible.
We consider this species to be incertae sedis, probably representing some form of
Phascolosoma (Phascolosoma) and possibly P. scolops.
ACKNOWLEDGEMENTS
This paper was stimulated by our work in Japan and recent visits to several
European museums and was made possible by the generous co-operation of the
following individuals and institutions : H. Terayama, Univ. Zool. Mus., Tokyo; Z.
Kawabata, Tohoku Univ. Zool. Inst., Sendai; A. Inaba, Mykaishima Mar. Biol.
Stat., Onomichi; S. van der Spoel, Inst. Taxonomische Zool., Amsterdam; J. van
der Land, Rijkemuseum van Nat. Hist., Leiden; J. Renaud- Mornant, Mus. Natl.
d’Hist. Natr., Paris; G. Hartwich, Mus. fur Naturkunde, Berlin, DDR; M.
Dzwillo, Univ. Zool. Inst. und Zool. Mus., Hamburg; J. B. Kirkegaard, Univ.
Zool. Mus., Copenhagen ; R. Olerod, Naturhistoriska Riksmuseet, Stockholm ; S.
Chambers, Roy. Scottish Mus., Edinburgh; K. A. Joysey, Univ. Mus. Zool.,
Cambridge; R. W. Sims, Brit. Mus. (Nat. Hist.), London. Consultation with M.
Rice (Washington) and T . Nishikawa (Nagoya) is gratefully acknowledged. P. E.
Gibbs (Plymouth) has helped in several ways including a critical reading of an
early draft of this article. Funding for the projects was provided by the U . S.
National Science Foundation ( I N T 7814554 and DEB 801 1121).
EXAM INATION O F PHAS(.’OI.OSOMA SUBGENERA
187
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