Table Of ContentThe Beagle, Records of the Museums and Art Galleries of the Northern Territory, 1998 14: 149-158
PRISTINA PROBOSCIDEA AND PRISTINELLA OSBORNI (OLIGOCHAETA,
NAIDIDAE) FROM A FRESHWATER CREEK
NEAR DARWIN, NORTHERN TERRITORY, AUSTRALIA,
WITH DESCRIPTIONS OF THE GENITAL ORGANS
OF BOTH SPECIES
CHRISTER ERSEUS' AND REINMAR GRIMM"-
'Department of Invertebrate Zoology, Swedish Museum of Natural History,
Box 50007, SE-104 05 Stockholm, Sweden
^Zoologisches Institut und Zoologisches Museum, Universitdt Hamburg,
Martin-Luther-King-Platz 3, DE-20146 Hamburg, Germany
ABSTRACT
Naidid oligochaetes from a freshwater creek feeding into West Arm, Darwin Harbour, were collected in July
1993. The majority of the worms were in different stages of sexual maturity. They represent two species,
Pristina prohoscidea Beddard, 1896, and Pristinella osborni (Walton, 1906). In this paper, full descriptions
of both somatic and genital features of these taxa are given. Taxonomic and phylogenetic aspects are
discussed.
KEYWORDS: Naididae, Oligochaeta, Pristina, Pristinella, taxonomy, zoogeography
INTRODUCTION records of microdrile Oligochaeta from
inland waters of Australia (Finder and
The Naididae, with a total of about 175 Brinkhurst 1994) includes 31 species of
species in about 25 genera worldwide, Naididae, but it can be expected that this
comprise a considerable part of the number will increase when additional
freshwater oligochaete fauna of all habitats have been investigated.
continents except Antarctica. Several genera The wide distribution of the Naididae is
and species of this family are more or less largely due to their opportunistic strategy of
cosmopolitan, occurring in virtually all asexual reproduction as the normal case. For
kinds of freshwater habitats. There is a many naidid species, reproductive organs
surprising coincidence in the number of have never been described, which is
species found on the different continents: 54 unsatisfactory, as the classification and
species are known from Europe, 81 from phylogenetic assessment of oligochaetes are
Asia, 51 from Africa, 55 from North largely based on the position and
America, and 52 from South America morphology of these structures.
(Brinkhurst and Jamieson 1971; Brinkhurst While participating in the Sixth
1986; Harman et al. 1988; Grimm 1987; International Marine Biological Workshop
Sememoy and Timm 1994). The high score on the Marine Flora and Fauna of Darwin
of Asian species mainly depends on the fact Harbour, at Mandorah, near Darwin
that Lake Baikal, with its many endemic (Northern Territory, Australia) in July 1993,
taxa, has been extensively studied by the first author collected a few naidid
Russian workers (Semernoy and Timm oligochaetes from a freshwater creek feeding
1994). Some of these species are, however, into West Arm, Darwin Harbour. The
briefly described. A recent compilation of majority of these worms were in different
149
C. Erseus and R. Grimm
stages of sexual maturity. The material specimens (one mature, plus the immature
represents two species, Pristina proboscidea worm) both 3.5 mm long (contracted),
Beddard. 1896, and Pristinella osborni consisting of a total of 46 and 35 segments,
(Walton, 1906), both widespread in the respectively. Anterior zooid of immature
tropics, but never before reported as sexually wonn with 21 segments. Prostomium with
mature. In this paper, full descriptions of proboscis (Fig. 3A). Clitellum extending
both somatic and genital features of the two over VIII-IX, when developed. Dorsal setae;
taxa are given, and some taxonomic and hairs one to four per bundle, serrate, about
phylogenetic aspects of these findings are 200 pm long, none especially elongated;
di.scussed. needles one to four per bundle, straight,
Other material of (largely marine) single-pointed with fine tips, and without
oligochaetes collected during the Darwin nodulus, 42-49 pm long. Ventral setae three
Harbour workshop has been published to seven per bundle in Il-lII, five to seven
separately (Coates and Stacey 1997; Erseus per bundle in IV, four to eight per bundle in
1997; Healy and Coates 1997). V, up to seven per bundle in VI of asexual
specimens, up to nine per bundle in
following segments, decreasing to mostly
METHODS five per bundle further towards the posterior.
All setae bifid, with distal prong slightly
The sand from the creek was repeatedly
longer than proximal (Fig. lA-C). Ventral
stirred with habitat water and the organic
setae of II-V 54-60 pm long, posterior
suspensions decanted into a fine-mesh sieve
ventral setae 48-58 pm long. One specimen
(250 pm). Live worms were sorted under a with ventral setae in II and III longer (63-78
dissecting microscope and fixed in Bouin’s pm) and thicker than the rest. In VI of
fluid. After about one day, they were specimens in an advanced stage of .sexual
transferred into 70% ethanol. All specimens maturity, ventral setae (genital setae; Figs
were stained in alcoholic paracarmine and ID; 3B: gs) about 70 pm long, bifid with
mounted whole in Canada balsam. Material prolonged straight, parallel, teeth comprising
of both species is deposited in the Museum one third of setal length, and with nodulus at
and Art Gallery of the Northern Territory one third from proximal end (Fig. ID). One
(NTM). Some specimens of Pristina specimen of full sexual maturity with one
proboscidia are also lodged in the Swedish slightly modified bifid seta representing each
Museum of Natural History (SMNH), ventral bundle in VIII (Fig. 1C). In yet
Stockholm. another mature specimen (depicted in Fig.
In the descriptions, segment numbers are 3B), ventral setae of VIII lacking. Male and
denoted by Roman numerals. spermathecal pores paired, in line with
ventral .setae; male pores anterior to middle
of VIII, spermathecal pores immediately
Pristina proboscidea Beddard, 1896
posterior to furrow between VI and VII.
(Figs 1, 3)
Pharyngeal glands in Ill-V.
Coelomocytes few, spherical, each with a
Material studied. NTM Wo 0121-0127,
distinct nucleus surrounded by weakly
7 whole-mounted specimens, and SMNH
granulated cytoplasm. Transition from
Main coll. 1615-1618, 4 whole-mounted
oesophagus to stomach inconspicuous,
specimens; all from small freshwater creek
somewhere behind VII (marked by presence
feeding into Stephens Creek, West Arm.
of gut content). Male genitalia (Fig. 3B)
Darwin Harbour, Northern Territory,
paired. Sperm funnel indistinct, position and
Australia, I2°38’S, l3r43’E, sand with
shape (indicated in Fig. 3B) deduced from
roots and debris along a short stretch of
adhering spermatozoa and position of vas
pools and rapids; coll. C. Erseus, 15 July
deferens. Vas deferens and atrium a
1993 (CE station no. 35).
continuous tubular male duct, at least about
Description. Ten specimens in various
375 pm long. Proximal end of male duct
stages of sexual maturity, and one immature
(Fig. 3B: vd), a short, naked part of vas
chain with two zooids. Two complete
150
Naidid oligochaetes from a freshwater creek near Darwin
Figs 1-2. Pristina pmboscidea: lA, elongate ventral seta in segment II; IB, normal posterior ventral seta;
1C, single ventral seta in VIII; ID, genital seta in VI; IE, dorsal needle seta. Pristinella oshornr. 2A.
posterior ventral seta; 2B, genital seta in VI; 2C, dorsal needle seta. Scale: 30 pm.
deferens, up to about 15 pm wide. Middle Remarks. The first description of
part of male duct, i.e. remaining part of vas Pristina proboscidea was given by Beddard
deferens, embedded in continuous mass of (1896). It was based on two South American
tightly packed, heavily granulated, prostate sexually immature specimens and is rather
cells (pr); this part of duct ciliated, incomplete. Sperber (1948) rede,scribed the
somewhat coiled, but very difficult to follow species as completely as it was then possible.
throughout its length. Distal part of male The specimens of the present study fit
duct (a) a naked atrium, about 170 pm long, Sperber’s description with the exception of
12-18 pm wide, terminally opening to the slight differences in the number of ventral
exterior through inconspicuous pore; atrium setae in the anterior segments, which,
appears non-ciliated. No part of male duct however, lie within the variability usual for
with notable muscular layer. Spermathecae naidids.
(Fig. 3B: s) small and club-shaped, each Pristina proboscidea has been considered
with duct, about 35 pm long, and about 28 to be closely related to the highly variable P.
pm wide, and an oval ampulla, about 60 pm longiseta Ehrenberg, 1828 (e.g., Rodriguez
long, and about 45 pm wide. In one 1987). To date, the only important
specimen (Fig. 3B), spermathecal ampullae taxonomic difference between these two taxa
containing loose masses of stained material has referred to the possession of elongated
reminiscent of sperm, but these hair setae in segment 111 in P. longiseta', in P.
‘spermatozoa’ much shorter than those proboscidea the hair setae of this segment
adhering to sperm funnel. are as long as those of the neighbouring
151
C. Erseus and R. Grimm
segments. Typically, the hairs of both P. proboscidea in the list of African naidid
longiseta and P. proboscidea are serrated, species. Its possible African occurrence has
but the elongate hairs of P. longiseta are been ba.sed on a single individual found on
normally smooth. Sometimes, however, the Zanzibar (Michaelsen 1905), but the
setal characteristics of P. longiseta overlap affiliation of this specimen with P.
with those of P. proboscidea. In African proboscidea is more than doubtful. On the
material, Grimm (1990) found (1) specimens basis of setal morphology, the question may
with elongate, smooth hair setae in segments even arise whether P proboscidea is merely
II and III; (2) a large variation in the a variety of the widespread P. longiseta with
serration of the hair setae (even the elongate reduced hair setae in segment III.
hairs in segment III were sometimes However, the present material of P.
serrated), which has been used to separate probo.scidea has revealed that the genital
subspecies in P longiseta (Sperber 1948: organs are different in the two taxa. In P
237; Brinkhurst and Jamieson 1971: 403); longiseta, the atria are clearly wider and
and (3) specimens in which only the hairs of have thicker walls than the vasa deferentia,
III, or those of both II and III, were reduced and although the epithelia of both the atria
in length or even completely lacking, i.e. and vasa deferentia are glandular, prostate
animals that according to an identification glands are nowhere present (Piguet 1906;
key would be P. probo.scidea. Absence of Figs 22, 24-25; Sperber 1948). This differs
elongate hairs may in some cases be due to considerably from the slender, thin-walled
loss, probably in handling the specimens. atria, and heavily prostatic va.sa deferentia in
Because of these taxonomic uncertainties, P. probo.scidea (Fig. 3B). Furthermore, in P.
Grimm (1990) did not include P. longiseta, the spermathecae are not small
Fig. 3. Prisma proboscidea. A, prostomium with proboscis; B, lateral view of segments VI-VIII, showing
genital seta, spermatheca and male duct of one side of worm. Abbreviations: a. atrium; gs. genital seta; nip,
male pore; pr, prostate gland; s, spermatheca; .sg. setal gland; sc, somatic seta; sf, sperm funnel; vd, vas
deferens.
152
Naidid oligochaetes from a freshwater creek near Darwin
and club-shaped (as in P. probascidecr. Fig. bitld with nodulus at half to one third from
3B), but long, with ampullae extending distal end. Teeth of needles clearly visible,
through the whole length of segment VII equally long (about 2.5 pm) and diverging at
(Piguet 1906: Figs 22, 25). Finally, Piguet a wide angle (Fig. 2C). Needles 39 pm long
reported that, in R loiigisefa, the glands in II, 34 pm in III, decreasing towards 30 pm
associated with the genital setae in VI do not in posterior segments. Ventral setae four to
envelop the setae but are situated separate live per bundle in II, five per bundle in
from and behind these setae (Piquet 1906: III-IV, six per bundle in V, one per ‘bundle’
Figs 22, 25). in VI-VII, four per bundle in VIII, then
With regard to the prostatic vasa decreasing to three per bundle posteriorly.
deferentia and the slender, thin-walled atria, Most anterior ventrals with distal tooth
P. proboscidea resembles P. americana slightly longer than proximal, and nodulus
Cernosvitov, 1937 (Cernosvitov 1937: Fig. proximal; posterior ventrals with equally
10) more closely than P. Umgiseta. long teeth and nodulus median to slightly
Distribution. To date, Pristina distal (Fig. 2A). Ventral setae not measurable
proboscidea has been recorded from North in anterior segments; setae more than 32 pm
(Brinkhurst 1986) and South America long, decreasing to about 30 pm in the
(Harman et al. 1988), South (Ali and posterior segments. In segments VI and VII,
Rashiduzzaman 1976) and East Asia genital setae modified, about 60 pm long,
(Brinkhurst et al. 1990), Australia (Binder without nodulus, bifid with prolonged
and Brinkhurst 1994; present paper) and straight, parallel, teeth comprising one third
New Zealand (Bayly 1989). Its occurrence in of setal length (Figs 2B. 4: gs). Each genital
continental Africa is less certain (see above), seta associated with large gland (Fig. 4: sg)
but it has been reported from the irrigation histologically resembling prostate glands.
system of Tenerife (Grimm 1978). A record Male and spermathecal pores paired, in line
from the Iberian Peninsula (Rodriguez and with ventral setae; male pores somewhat
Armas 1983) is the only known European anterior to middle of VIIl, spermathecal
occuirence so far. The species thus appears pores immediately posterior to furrow
widespread, but it may not be cosmopolitan. between VI and VII.
Rodriguez (1987) considered the special Pharyngeal glands in III-V. Coelomocytes
geographical situation of the Iberian numerous, spherical, about 5-10 pm wide,
Peninsula to favour recent introductions of each with distinct nucleus surrounded by
American naidid species. This could apply weakly granulated cytoplasm. Stomachal
also to Tenerife. dilatation not distinct. Male genitalia (Fig. 4)
paired. Sperm funnel indistinct, but revealed
by adhering spermatozoa. Male duct a
Pristinella osborni (Walton, 1906) continuous, ciliated, tube, at least about 180
(Figs 2, 4) pm long, about 10-12 pm wide; transition
between vas deferens and atrium not
Material studied. NTM Wo 0128, 1 obvious. Proximal part of duct naked, middle
whole-mounted specimen from small part surrounded by continuous mass of
freshwater creek feeding into Stephens tightly packed, heavily granulated, prostate
Creek, West Arm, Darwin Harbour, Northern cells. Distal part of male duct naked and
Territory, Australia, I2°38’S, I3I°43’E, terminally opening to exterior through
sand with roots and debris along a short inconspicuous pore. No part of male duct
stretch of pools and rapids; coll. C. Erseus, with any notable muscular layer.
15 July 1993 (CE station no. 35). Spermathecae (Fig. 4: s, so) small and club-
Description. Length (contracted) 1.9 mm. shaped, each with somewhat bulbous duct,
Sexually mature specimen without budding- about 20 pm long, about 15 pm wide, and
zone, with 26 segments. Pro.stomium oval ampulla, 30-35 pm long, about 25 pm
without proboscis, rounded. Clitelium not wide. Spermathecal ampullae containing
developed. Dorsal setae: hairs one per spermatozoa, somewhat loosely arranged in
bundle, smooth; needles, one per bundle. small bundles.
153
C. Erseus and R. Grimm
Fig. 4. Pristinella oshorni, somewhat obliquely lateral view of segments Vl-Vlll, showing genital setae and
male duct of one side of worm, plus spermathecae of both sides. Abbreviations: gs, genital seta; nip, male
pore; pr, prostate gland; s, spermatheca; sf, spenn funnel; so. spermatheca of other side; vd, vas deferens.
Remarks. This species, one of the According to Brinkhurst and Jamieson
smallest naidids known, was first described (1970), there should be a sudden stomachal
by Walton (1906), as Naidium oshorni, from dilatation in VII or VIII, but none is visible
Ohio, North America. Stephenson (1914) in the present material.
described Naidium mimitiim from Lahore, As de.scribed above, each male duct of P.
India. Sperber (1948) placed both species in oshorni is a ciliated tube without a clear
the genus Pristina as P. ininuta Stephenson, demarcation of an atrium, and it is not
1914. Brinkhurst and Jamieson (1971) stated conclusive whether or not a proper atrium
that the separation of Naidium oshorni from (an invaginated, ectodermal, part of the male
N. mi nut urn was based on such slight duct) is pre.sent.
differences that it had to be neglected, and Distribution. Pristinella oshorni appears
correctly used the oldest name, Pristina more or less cosmopolitan. It is the most
oshorni (Walton, 1906). Following abundant Pristinella species in Europe,
Brinkhurst (1985), it is now called where it is often found in freshwater
Pristinella oshorni (see Discussion). meiobenthos and in groundwaters. It has
The new specimen fits the description been reported from southern Europe
given by Sperber (1948). Its needle setae are (Martinez-Ansemil 1984; Sambugar 1986)
simply bifid and not pectinate. Intermediate up to the River Rhine basin in the north
teeth as reported by Grimm (1990) and (Lafont and Durbec 1990; Wiegand and
Finder and Brinkhurst (1994) are not visible Matthess 1993). In America, the
(very few needle setae are in a suitable northernmost report comes from Lake Erie
position for detailed observation). (Spencer 1977), the southernmost from
154
Naidid oligochactes from a freshwater creek near Darwin
Bolivia (Juget and Lafont 1994). It i.s also Thus, with regard to genital features, P.
recorded from Africa (Grimm 1990), Asia osborni fits Brinkhurst's definition ol
(Naidu 1961) and Australia (Finder and Pristina belter than that of Pristinella.
Brinkhurst 1994; present paper). Pristina and Pristinella contain about ten
species each, but they are often difficult to
discriminate because of a wide overlap of
DISCUSSION taxonomic characteristics. Several additional
nominal species in the literature are regarded
Pristina Ehrenberg, 1828, and Pristinella as dubious (e.g., four species described by
Brinkhurst, 1985, originally treated as a Botea 1983) or as junior synonyms of other
single genus (Pristina), are currently the taxa.
only genera in the subfamily Pri-stininae The male ducts are now known for six of
sensu Sperber (1948), a ta.xon subsequently the Pristina species: P. longiseta (type
revised to become a tribe, Pristinini, within species), P. breviseta Bourne, 1891, P.
the Naidinae (Nemec and Brinkhurst 1987). phnnaseta Turner, 1935, P. leidyi Smith,
This group is ‘curiously homogeneous' 1896, P. americana and P. proboscidea. The
(Sperber 1948: 247), differing from all other atria are generally slender (short in P.
Naididae by (1) its dorsal setae commencing phnnaseta), and when present, the prostate
from segment II, (2) its continuously tubular glands are associated with the vasa
male ducts, and (3) the relatively high deferentia, not with the atria. Prostates have
number (seven as opposed to four to five) of not been observed in P. longiseta and P.
segments formed at the anterior end in a leidyi. but the inner ends of the male ducts
budding zone. At least the first two of these have somewhat glandular walls in these two
features may be plesiomorphic, but for the species (Piguet 1906; Smith 1896).
sake of the following discussion it is For the taxa currently placed in Pristinella
assumed that the Pristininae/Pristinini are (.see Brinkhurst 1985), infomiation on male
monophyletic. Sperber’s (1948) description ducts is scanty. According to Sperber (1948),
of the Pristininae also mentions ‘the Pristinella ainplubiotica and P. idrensis have
characteristic stomach, with cells containing short vasa deferentia followed by atria that are
intra-cellular canals with special features', either ‘not differenlialed’ or ‘small’; prostate
but these canals were not observed in the cells are ‘probably’ absent. In his description of
present material. As pointed out by 'Pristina amphibiotica changtuensis', Liang
Brinkhurst (1985), they need to be re¬ (1963: fig. 2F) illustrated a more developed
investigated using electron microscopy. and differentiated atrium, the male duct,
Brinkhurst (1985) separated Pristinella however, still lacking prostate glands.
from Pristina sensu stricto, refening to a Pristinella osborni, on the other hand, has
number of morphological differences. prostate glands along the middle part of its
According to him, Pristina is characterised male ducts but lacks demarcated atria (Fig. 4).
by a pro.stomial proboscis, genital setae in at In fact, as the male ducts of P. osborni arc
least two different segments (varying ciliated throughout, this species may lack atria
positions), presence of spermathecae, and altogether; Sperber (1948: 49) claimed that
prostate glands on the vasa deferentia, naidid atria are non-ciliated. The previously
whereas in Pristinella, proboscis, prostate studied material of P. amphibiotica and P.
glands and spemiathecae are absent, and idrensis were possibly not as sexually mature
genital setae arc either absent, or present in as the pre.sent individual of P. osborni, which
VIII (based on the genital organs of only two could explain why prostate glands and
species, Pristinella amphihiotica (Lastockin, spennathecae were not observed in the two
1927), and P idrensis (Sperber, 1948). former taxa. Alternatively, P. osbonn could be
Pristina prohoscidea fils into this general regarded as representing a third (yet un-named)
pattern, whereas Pristinella osborni does genus in the Pri.stina/Pristinella complex. At
not. The latter possesses spermathecae as any rate, the distinction between Pristina and
well as prostates, and it has genital setae in Pri.stinella does not appear as clear as it was
both VI and VII, but not in VIII (sec Fig. 4). initially thought to be (see Brinkhurst 1985).
155
C. Erseus and R. Grimm
A prostomial proboscis does not occur Brinkhurst, R.O. 1986. Guide to the freshwater
only in Pristina, but also in the other naidid aquatic microdrile oligochaetes of North
genera Stylaria Lamarck, 1816, Arcteonais America. Canadian Special Publication of
Piguet, 1928, and Ripistes Dujardin, 1842, Fisheries and Aquatic Sciences 84: 1-259.
and Homochaeta Bretscher, 1896 (Grimm Brinkhurst, R.O. and JaEiiieson, B.G.M. 1971.
1985). The phylogenetic analysis by Ncmec Aquatic Oligochaeta of the world. Oliver &
and Brinkhurst (1987), however, indicates Boyd: Edinburgh.
that the three latter taxa are not closely Brinkhurst, R.O., Qi, S. and Liang, Y.L. 1990. The
related to the Pristina/Pristinella complex. aquatic Oligochaeta from the People’s
The (synapomorphic) possession of a Republic of China. Canadian Journal of
proboscis thus appears to support Zoology 68: 901-916.
monophyly of Pristina sensu stricto, i.e., Cemosvitov, L. 1937. Notes sur les Oligochaeta
assuming that the absence of a proboscis in (NaYdiddes et Enchytraeiddes) de I’Argentine.
Pristinella is not the result of a .secondary Anales de! Museo Argentino de Ciencias
loss of this character. Naturales Buenos Aires 39: 135-157.
The observations on the present material Coates, K. A. and Stacey, D. F. 1997. Enchytraeids
elucidate the importance of including genital (Oligochaeta: Annelida) of the lower shore and
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resolved. fauna of Danvin Harbour, Northern Territory,
Australia. Pp 67-79. Museums and Art
Galleries of the Northern Territory and the
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158
