Journal of Foraminiferal Research, v. 30, no. 2, p.

135–142, April 2000

PRELIMINARY SURVEY OF ARCELLACEANS (THECAMOEBIANS) AS

LIMNOLOGICAL INDICATORS IN TROPICAL LAKE SENTANI,
IRIAN JAYA, INDONESIA

A. P. DALBY1, ARUN KUMAR,1 J. M. MOORE2, and R. T. PATTERSON1

ABSTRACT pollution (Asioli and others, 1996; Patterson and others,

1996; Reinhardt and others, 1998).
Arcellacean (thecamoebian) assemblages recovered Previous studies have focused mainly on faunas found in
from Lake Sentani, a large tropical lake southwest of higher latitude temperate zone lakes (Medioli and Scott,
Jayapura, Irian Jaya, Indonesia, are characterized by 1988). The minimal research that has been carried out in
low diversity and low abundances. Dominated by Cen- tropical regions has been primarily descriptive, with scant
tropyxis aculeata and Arcella vulgaris, this fauna is sim- attention paid to limnological associations. Examples of this
ilar to those indicative of stressed environments (brack- research include: faunas from lakes in Java and Sumatra,
ish conditions, high levels of industrial contaminants) in Indonesia (Hoogenraad and Groot, 1940, 1946; van Oye,
temperate regions. However, neither condition exists in 1949); lakes in Malaysia (Sudzuki, 1979); brackish and
Lake Sentani. Previous work has determined that the freshwater lakes and ponds on the islands of Bombay (Cart-
lake is oligomictic, characterized by weak circulation er, 1856a, 1856b, 1864, 1865); the Sokoto River in Nigeria
with turnover occurring only every few years. Prolonged (Green, 1963) and the wetlands of central Brazil (Green,
isolation of the lake bottom produces progressively re- 1975).
duced oxygen levels and results in reduced productivity The purpose of this study is to add to this database by
among benthic organisms. The feeble stratification that documenting the arcellacean fauna found in tropical Lake
exists here creates reduced oxygen levels at depth pro- Sentani, Irian Jaya, Indonesia, and variations in arcellacean
viding a likely explanation of the stressed arcellacean assemblages to changes in measured limnological parame-
fauna. The oligomictic conditions observed here and the ters.
resultant fauna are widespread and are characteristic of
a large proportion of tropical lakes around the world.
As the low bottom water oxygen conditions will have a LAKE PHYSIOGRAPHY
serious impact on most benthic organisms in these lakes, Lake Sentani occurs in the northeastern corner of the
other limnological signals including anthropogenic con- Province of Irian Jaya, Indonesia, southwest of Jayapura
tamination will be masked. This is a disappointing result (Fig. 1). It is a large oligomictic lake (28 km east-west by
as the utility that has been developed for the group as a 19 km north-south; area 10,400 ha), situated at an elevation
limnological indicator in temperate lakes does not ap- of 73 m in a fault-controlled depression. The bedrock base-
pear to apply in a significant proportion of low latitude ment consists mainly of Mesozoic mafic and ultramafic
lakes. rocks of the Cyclops Ophiolite Belt (Moore and others,
1995). The lake is fed by a catchment area of about 600
INTRODUCTION km2 and has only one outlet via the Jafuri River to the
nearby Pacific Ocean near the Papua New Guinea border.
Arcellaceans (thecamoebians) are freshwater protozoans Average annual rainfall around the lake is high, averaging
with agglutinated or autogenous tests that occur abundantly about 2 m. Due to seasonal variations in fluvial inflow, the
in most Quaternary lacustrine sediments (Medioli and Scott, lake level fluctuates about 0.4 m (Howard, 1987). The lake
1983; Scott and Medioli, 1983). Distinct associations of ar- is oligomictic (infrequent turnover events) with oxygen
cellacean species and strains (infraspecific morphotypes) stratification occurring in the western basin at 22 m depth
can be correlated with a variety of climatic and other en- (Canadian International Development Agency, 1985; How-
vironmental factors such as oxygen levels, minimum aver- ard, 1987; Table 1; Fig. 1).
age water temperature, level of organics and substrate type, Measurements taken in this and previous studies indicate
clastics, and pollution levels (Scott and Medioli, 1983; Pat- lake temperatures ranging from 29⬚C to 31⬚C. Surface water
terson and others, 1985; Medioli and others, 1990; Collins pH is near neutral with values measured between 6.2 and
and others, 1990; McCarthy and others, 1995; Patterson and 7.2. Due to limited circulation, this oligomictic lake is quite
others, 1996; Asioli and others, 1996). Recent studies car- turbid, particularly in the westernmost basin. Plankton levels
ried out in lakes of northeastern Ontario, Canada and north- are low at 1–2 mg/L, although periodic algal blooms often
ern Italy have further identified a positive relationship be- result in high levels of fish mortality (Canadian International
tween arcellacean infrasubspecific strains and heavy metal Development Agency, 1985).

1 Ottawa-Carleton Geoscience Centre, Department of Earth Sciences, MATERIALS AND METHODS

Carleton University, Ottawa, Ontario, K1S 5B6, Canada.
2 Department of Earth Sciences, Simon Fraser University, Burnaby, Eighteen sediment-water interface samples were collected
British Columbia, V5A 1S6, Canada. using an Eckman box grab sampler in September of 1994

135
136 DALBY, KUMAR, MOORE, AND PATTERSON

FIGURE 1. Lake Sentani Map showing sample localities. Numerical localities represent samples collected during 1994 and 1995, and alphabetical
localities from 1997.

and May of 1995. Water depth, surface water temperature, Before shipment to Canada for processing, a 30% methyl
water temperature at depth, and a relative measure of oxy- alcohol solution was added to each sample to inhibit decay
gen concentration at depth were recorded for each sample of organic material. Rose Bengal (acid red 94) protoplasmic
locality. Oxygen and temperature measurements were re- stain was then added to indicate whether the specimens were
stricted to 15 m or less by limitations of the equipment. In alive at the time of collection. After standing for several
August of 1997, pH and salinity readings were recorded hours, the samples were screened with a 1000 ␮m sieve to
from ten additional localities (Table 1; Fig. 1). remove coarse particles, and a 43 ␮m screen to retain ar-
The geographical location of each sample site was deter- cellaceans, and to remove silts and clays. Finally, all sam-
mined using a Magellan Global Positioning System receiver. ples were treated with isopropyl alcohol and refrigerated.
A commercial sonar device (fish finder) was used to deter- The samples had an average volume of 18 cm3 and were
mine the depth of each sample station. In 1997, lake bottom
entirely processed.
water samples were obtained using a WildCo 1510TT water
The samples were then subdivided into aliquots for anal-
sampler. Measurements of pH of water samples were cali-
ysis using a wet splitter (Scott and Hermelin, 1993). Al-
brated against a pH 7.01 buffer standard. Salinity was mea-
sured with an automatic temperature compensating salinity though the low numbers of specimens found in this study
refractometer calibrated with distilled water, and was below did not really warrant splitting, low abundances could not
the detection limit in all samples. be determined prior to counting, which was subsequently
Samples with significant populations of arcellaceans were done using a binocular microscope.
found at muddy bottom sites with a large admixture of fine Electronically stored scanning electron micrographs were
organic material. Sites characterized by winnowed sandy obtained using a JEOL 6400 scanning electron microscope
substrates generally yielded small allochthonous arcellacean at the Carleton University Research Facility for Electron
assemblages. Some attempted sample stations were found Microscopy (CURFEM). The plate was electronically pro-
to be floored by exposed rock, providing no samples for duced on an Apple Macintosh computer using Adobe Pho-
analysis. toshop and output to a Linatronic printer.
 ARCELLACEANS FROM LAKE SENTANI INDONESIA 137

TABLE 1. A. Sample locations, physical characteristics of Lake Sentani, specimen counts and their percentages. Field data collected in September
of 1994 and May of 1995 by J.M. Moore. B. Sample locations and pH measurements of water samples. Salinity was below detection limit in all
samples and thus was not presented on this table. Field data collected in August of 1997 by J.M. Moore.

RESULTS AND DISCUSSION and strains were very low as well, rendering more substan-
The arcellacean fauna of Lake Sentani is quite impover- tive multivariate cluster analyses unfeasable. Therefore,
ished in terms of species numbers and Shannon Diversity only a qualitative assessment of the data was carried out.
Index scores (Table 1). Total counts of individual species All samples were dominated by two species, Arcella vul-
138 DALBY, KUMAR, MOORE, AND PATTERSON

PLATE 1
Scanning electron micrographs of Lake Sentani arcellaceans. 1 a–e Incerta sp. A. Agglutinated sphere of unknown affinity. 2 a–b Centropyxis
constricta ‘‘aerophila’’. Aperture forms an angle with the fundus. 3 Difflugia protaeiformis ‘‘claviformis’’. Test made from coarse grains and is
 ARCELLACEANS FROM LAKE SENTANI INDONESIA 139

garis Ehrenberg 1830 and Centropyxis aculeata Ehrenberg this lake though, and the periodic fish-killing algal blooms
1832. In a few samples various difflugids and a possible suggest that some eutrophication, at least in surface waters,
new species of arcellacean, identified as Incerta sp. A, also could be occurring. However, eutrophication is not indicated
occur in appreciable numbers. by the benthic arcellacean fauna because Difflugia oblonga,
To evaluate the limnological significance of this arcella- one of the major indicator taxa of high organic content (Col-
cean assemblage, it is important to understand their ecology lins and others, 1990), is almost entirely absent from the
and distribution. Impoverished faunas similar to those found lake.
in Lake Sentani have been described elsewhere, and could The dominance of the arcellids and centropyxids coupled
be attributed to one or more of the distinct environmental with the dearth of difflugids may be a function of substrate
controls discussed below: brackish water conditions, low type. With the exception of the barren samples (S12A,
pH, anthropogenic contamination, substrate type, and envi- S14A) which contained a fair amount of mineral grains, all
ronmental conditions peculiar to tropical lakes. samples were rich in organic matter. Difflugids require min-
Moore and others (1995) proposed that Lake Sentani was eral grains to construct their tests and in the absence of such
an arm of the ocean well into the Holocene and that there will not be found in any appreciable numbers, if at all (Ha-
still might be saline wedges present at depth. Centropyxids, man, 1990; Scott and others, 1991). Arcellids produce their
primarily Centropyxis aculeata, are known to be opportu- tests autogenously, and centropyxids in Lake Sentani seen
nistic and capable of existing under hostile conditions (Pat- to be able to utilize diatom frustules to construct their tests
terson and others, 1996; Reinhardt and others, 1998) and (Plate 1, fig. 6e), so it is not surprising that they dominate
have been reported from brackish environments in coastal the organic-rich samples.
areas with salinities as high as 5‰ (Decloitre, 1953; Todd Other factors that control the geographic distribution of
and Bronnimann, 1957; Honig and Scott, 1987; Medioli and modern arcellaceans include the vegetation in and around
Scott, 1988; Hayward and others, 1996). Faunas similar to lakes and climatic conditions, which ultimately control wa-
those in Lake Sentani are found in cores from lakes in up- ter levels, chemistry, trophic levels, and the nature of a ther-
lifted coastal areas of New Brunswick, Canada, in intervals mocline, if present (Collins and others, 1990). The low di-
marking the transition from marine to freshwater conditions versity and dominance in all samples by Centropyxis acu-
(Patterson and others, 1985). However, water testing indi- leata and Arcella vulgaris indicate a stressed environment
cated no trace of salinity in any Lake Sentani sample. unlike any recorded in northern hemisphere temperate lakes.
Arcella vulgaris is an important component of the arcel- A characteristic of tropical oligomictic lakes like Lake Sen-
lacean faunas of Lake Sentani, dominating many samples. tani is very weak circulation with turnover occurring only
This species is commonly reported from boggy ponds in the every few years, thus resulting in diminished oxygen con-
Arctic and further south to Florida (Collins and others, tent (Wetzel, 1983).
1990), and is well adapted to the low pH typical of these Although our water temperature and oxygen data only
ponds. In James Lake of northeastern Ontario, A. vulgaris extend to 15 m depth, other work has shown ‘‘. . .that ox-
almost exclusively dominates low pH (2.0–5.5) environ- ygen stratification occurs in the western basin at 22 m
ments (Kumar and Patterson, 1997; Kumar and Patterson, depth’’ resulting in oligomictic conditions (Howard, 1987).
2000). However, in studies on lakes with varying pH in Although most of our samples are from the eastern region
northeastern Ontario it was found that centropyxids, so of the lake, the feeble stratification recorded by Howard
dominant in Lake Sentani, cannot tolerate low pH (⬍5.5) (1987) no doubt exists lakewide. This stratification ulti-
environments (Patterson and others, 1996; Reinhardt and mately results in the prolonged isolation of the lake bottom
others, 1998; Patterson and Kumar, 2000). Thus the abun- and progressively reduced oxygen levels. This would have
dance of centropyxids in Lake Sentani samples indicates in turn resulted in similarly reduced productivity among
that pH is not a controlling factor here, and the measured benthic organisms, as indicated by the stressed arcellacean
pH values of 7.1 to 7.2 for lake water at several depths in fauna. Qualitative reports on similar arcellacean faunas have
different localities corroborate this conclusion. been reported from Nigeria (Green, 1963) and Brazil
Centropyxid species found in the Cobalt area, northeast- (Green, 1975) suggesting that this fauna is widespread and
ern Ontario, such as Centropyxis aculeata often dominate characteristic of tropical lakes.
substrates contaminated by heavy metals and other indus- Although our data are sparse, the impact of prolonged
trial contaminants (Patterson and others, 1996; Reinhardt stratification on benthic organism productivity in these lakes
and others, 1998). No geochemical analysis was carried out is undoubtably strong enough to mask out most other lim-
on samples in Lake Sentani so this influence cannot be cat- nological signals, such as anthropogenic eutrophication.
egorically refuted. However, there is no major industry This could also provide a possible explanation for the dearth
there, and human habitation consists of only small villages of Difflugia oblonga in these samples, usually an excellent
along the shores and on the islands of Lake Sentani (Fig. indicator of eutrophication, despite other kinds of evidence
1). Sewage from houses in the villages goes directly into for lake eutrophication. Although it is dangerous to draw

←
opaque. 4 Difflugia oblonga ‘‘glans’’. Test ovoid with rounded fundus. 5 a Centropyxis aculeata ‘‘discoides’’. Same as ‘‘aculeata’’ but spines
absent. 5 b Arcella vulgaris. Test round, hyaline; aperture invaginate. 6 a–e Centropyxis aculeata ‘‘aculeata’’. 6 a–d This strain has a variable
number of spines. 6 e Enlargement of 6d showing imprints caused by diatoms, often used as xenogenous particles to construct arcellacean tests as
in the case of Lake Sentani centropyxids. All scale bars are 50 ␮m.
140 DALBY, KUMAR, MOORE, AND PATTERSON

sweeping conclusions based on results from a single lake Cucurbitella [sic.] constricta REINHARDT and others 1998, pl. 1, fig. 6
we suggest that, because of the serious impact on produc- Diagnosis. Test shape varies from spherical, subspherical to elon-
tivity caused by stratification, benthic arcellacean faunas gated. It is characterized by a thick apertural lip.
may be seriously affected throughout most oligomictic
Centropyxis constricta (Ehrenberg 1843)
lakes. This probably means that the group, such an impor- strain: ‘‘constricta’’
tant limnological indicator in temperate lakes, is not useful
Arcella constricta EHRENBERG 1843, p. 410, pl. 4, fig. 35, pl. 5,
in a significant proportion of low latitude oligomictic lakes. fig. 1
Diagnosis. Test less flattened than strain ‘‘spinosa’’ with 3 or less
SYSTEMATIC PALEONTOLOGY spines on the fundus.
Although infrasubspecific variants are not considered val- Discussion. These specimens were fragile and became broken when
mounted on the scanning electron microscope plug.
id according to the International Code of Zoological No-
menclature (ICZN), their informal use is often useful in pa- Centropyxis constricta (Ehrenberg 1843)
leoecological studies. This is particularly true with regards strain: ‘‘spinosa’’
to arcellaceans as the species concept within this clonal Centropyxis spinosa CASH in CASH and HOPKINSON 1905, p. 135,
group is a highly subjective matter (see Deflandre, 1928; text figs. 26 a–c, pl. 16, fig. 15
Medioli and Scott, 1983; Medioli and Scott, 1988 for dis- Centropyxis spinosa Cash, OGDEN and HEDLEY 1980, p. 62, pl. 20,
cussion). For example, Reinhardt and others (1998) were figs. a–d
able to separate and define numerous distinct and stable ar- Diagnosis. Test more flattened than strain ‘‘constricta’’ with 3 or
cellacean strains that characterized various limnological more spines on the fundus.
Discussion. Only two specimens were found and they became bro-
conditions in northern Ontario lakes. In addition, Kumar and ken when mounted on the scanning electron microscope plug.
Dalby (1998) proposed a standardized key for lacustrine ar-
cellacean species and strains, which is used here. As this Family DIFFLUGIDAE Stein 1859
paper is not of a taxonomic nature, only abbreviated sys- Difflugia Leclerc in Lamarck 1816
Difflugia oblonga Ehrenberg 1832
tematic descriptions are included. strain ‘‘glans’’
(Plate 1; Figure 4)
Subphylum SARCODINA Schmarda 1871
Class RHIZOPODEA von Siebold 1845 Difflugia glans PENARD 1902
Subclass LOBOSA Carpenter 1861
Diagnosis. Test oval to ovoid, slightly elongated, fundus rounded,
Order ARCELLINIDA Kent 1880
neck absent, aperture circular with smooth lip, test made of fine sand
Superfamily ARCELLACEA Ehrenberg 1830
particles, small.
Family ARCELLIDAE Ehrenberg 1830
Arcella Ehrenberg 1830 Difflugia oblonga Ehrenberg 1832
Arcella vulgaris Ehrenberg 1830 strain: ‘‘tenuis’’
(Plate 1; Figure 5b)
Difflugia pyriformis var. tenuis PENARD 1890, p. 138, pl. 3, figs. 47–49
Arcella vulgaris EHRENBERG 1830, p. 40, pl. 1, fig. 6
Diagnosis. Test elongated, ovoid almost bean shaped, fundus sub-
Diagnosis. Test depressed, hyaline, and autogenous, with an invag- rounded to subacute, neck indistinct or absent, aperture narrow and
inated aperture. circular with crenulated lip, test made of generally medium to fine sand
grains.
Family CENTROPYXIDIDAE Deflandre 1953 Discussion. Only one specimen was found and it became broken.
Centropyxis Stein 1859
Centropyxis aculeata (Ehrenberg 1832) Difflugia protaeiformis Lamarck 1816
strain: ‘‘aculeata’’ strain: ‘‘claviformis’’
(Plate 1; Figure 6 a–e) (Plate 1; Figure 3)
Arcella aculeata EHRENBERG 1832, p. 91 Difflugia protaeiformis LAMARCK 1816, p. 95 (with reference to
Diagnosis. Test depressed, circular with 1–8 spines on postero-lat- material in a manuscript by Leclerc)
eral margin. Difflugia pyriformis var. claviformis PENARD 1899, p. 25, pl. 2, figs.
12–14
Centropyxis aculeata (Ehrenberg 1832) Difflugia claviformis OGDEN and HEDLEY 1980, p. 126, pl. 52, figs.
strain: ‘‘discoides’’ a–d
(Plate 1; Figure 5a) Diagnosis. Test elongated almost cylindroconical, fundus acuminate,
Arcella discoides EHRENBERG 1843, p. 139 tapering to form a blunt spine, neck absent, aperture circular, narrow
Arcella discoides Ehrenberg, EHRENBERG 1872, p. 259, pl. 3, fig. 1 without lip, test made up of medium to coarse grained sand
Arcella discoides Ehrenberg, LEIDY 1879, p. 173, pl. 28, figs. 14–38
Centropyxis aculeata var. discoides PENARD 1890, p. 150, pl. 5, figs. INCERTA
38–41
Centropyxis discoides Penard [sic], OGDEN and HEDLEY 1980, p. Incerta sp. A
54, pl. 16, figs. a–e (Plate 1; Figure 1 a–e)
Diagnosis. Test depressed, circular almost ‘‘doughnut shaped’’ with-
Diagnosis. Agglutinated sphere with no discernible aperture.
out spines.
Discussion. This could be an encystment of an arcellacean or the
aperture has been covered (F. Medioli, oral communication, 1997)
Centropyxis constricta (Ehrenberg 1843)
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