Hydrobiologia 271: 179-189, 1993. 0 1993 Kluwer Academic Publishers.
Primed
179
in Belgium.
The consortium of the sponge Lubomirskia baicalensis in Lake Baikal, East Siberia R. M. Kamaltynov, V. I. Chernykh, Z. V. Slugina & E. B. Karabanov Limnological Institute of the Siberian Division of the Russian Academy of Sciences, Ulan-Batorskaya str. 3, 664033, Irkutsk, Russia Received 8 April 1992; in revised form 10 March 1993; accepted 23 March 1993
Key words: sponge, benthic invertebrates,
consortium,
amphipod,
Baikal.
Abstract The spatial distribution of the fauna associated with a branched sponge, Lubomirskia baicalensis, endemic of Lake Baikal has been quantitatively studied. The biomass and numbers of three amphipod species which inhabit the sponge correlate (linearly or non-linearly) with the weight of the sponge.
Introduction Bottom biocenoses of Lake Baikal (East Siberia) involve associations supported by sponges. According to Beklemishev (1951), ‘organisms belong to biocenoses as components of a consortium, rather than as independent entities... A consortium consists of an edificator, and of a number of epibionts and endobionts’. Relations between edificators and supported species are mainly topically, as suggested by Beklemishev. Almost simultaneously, a similar approach was proposed by Ramenskiy (1952): ‘considering biocenoses, it is useful to distinguish within them combinations of organisms having the same destiny in their life cycles - consortive communities, or a consortium’. Since that time, consortia have been regarded as major structural parts, or elementary energetic systems of biocenoses (Lavrenko, 1959; Rabotnov, 1974). An important paper by Mazing (1966) proposed the name ‘consort’ for organisms depending energetically, or topically (by contact) on
a central organism. It is also believed (Arnoldi et al., 1969; Rabotnov, 1973) that consortia are groups of organisms which have originated due to concerted evolution of species and consist of autotrophic plants, and of heterotrophs dependent on such plants. The term ‘consortium’ has been proposed relatively recently, and problems related to it are actively discussed (for reviews, see Voronov, 1974; Kharchenko et al., 1989). Many authors do not use this name, although they study the phenomenon (Long, 1968; Tuzet & Paris, 1964; Wendt et al., 1986; Westinga & Hoetjes, 1981). We studied the sponge Lubomirskia baicalensis (Pallas) 1772-1773, an abundant endemic of Lake Baikal. The first data on ‘animals which are parasites’ of this sponge were published by Dybowsky (1874) - these were Spinacanthus parasiticus (Dyb.) 1874 and Eulimnogammarus violaceus (Dyb.) 1874. The close association between the sponge and these two species was also pointed out by Kozhov (1931, 1962, 1963). Observations were performed by means of
180 scuba diving. They showed that the community of inhabitants of this sponge is much richer than was hitherto believed. Earlier authors did not give any quantitative data, thus we investigated the relationships between the weight of consorts and that of the determinant species.
Study sites
Table
I.
Time
Characteristics Depth,
of samples. m
435
September November
6,4
Number of sponges
Wet weight, kg
Number of sponges
Wet weight, kg
5 3
0.05-2.59 1.45-3.01
10 2
0.06-1.36 1.04-1.47
Samples of branched sponges L. baicalensis were collected 17 km East of the outlet of the Angara River in Southern Baikal(105 o 04’ E, 5 lo 54’ N) at two sites, one at a depth of 4.5 m on the edge of a shallow-water platform, the other at a depth of 6.4 m, on the upper part of an underwater slope.
1969; Barnard, Barnard, 1983). We assume that it involves the sub-genera Eulimnogammarus Baz. 1945, Philolimnogammarus Baz. 1945, and Eurybiogammarus Baz. 1945; the reasons for this assumption will be discussed elsewhere.
Materials
Results
and methods
Sampling was performed by scuba divers in September and October 1990 (Table 1). To obtain quantitative data, a sponge was covered by a net (mesh size 0.2 mm) and lifted to the surface together with the stone to which it was attached. Sponges and their fauna were fixed in 4% formaldehyde. For preliminary investigation of their infauna, some sponges were dissected: five sections in apical position, five sections in basal position. All organisms were collected, identified to the lowest possible taxon, counted, and weighed. Sponge consorts are difficult to classify as nectobenthos, or benthos, therefore, we use the terms proposed by Clark (198 1): ‘epon’ - organisms living on surfaces of objects above the bottom, i.e., on substrates; and ‘peron’ - organisms residing nearby the surface, i.e., nearby substrates. The genus Eulimnogammarus Bazikalova, 1945 is under discussion (Bazikalova, 1945; Stock, Fig. la. Zonal distribution of consortium supported by a sponge enthos: 2 - amphipod P. erinaceus; 3 - sculpin P. jeittelesi. C. M. demersa; 6 - amphipod B. latissima; 7 - Trichoptera. D. Epiete Nais sp.; 10 - cyclops A. spongicola. F. Under sponge base: 11 base: 13 - polychaete M. baicalensis.
The bright-green, branched sponges L. baicalensis (Fig. 1) lives in Lake Baikal on stony bottoms at depths of 4-15 m, and often forms thickets on stony debris and rocks. Sponges tightly attack to substrates by their crust-shaped base, which sometimes cover large surfaces. Branches grow out of the crust. Branches of the younger are some 15 cm long, and their number is small (1 to 4). The number of branches increases with age; secondary branches sprout from primary ones; secondary branches often merge. The largest sponges collected had a height of 50-63 cm; their number of primary branches was 10 to 20; that number of terminal branches 48-114. Sponges with a small mass (0.05 to 0.15 kg, height 11 to 15 cm) typically contained no fauna of consorts, but this became abundant on sponges with masses over 0.17 kg (20 cm, 8 primary branches); a diverse fauna was also collected on a 0.06 kg (14 cm) sponge (Fig. 1, Table 2). The L. baicalensis. A. plankton: 1 - cyclops C. kolensis. B. NectobEpifauna (epon): 4 - amphipod S. parasiticus; 5 - mollusk infauna: 8 - amphipod E. violuceus. E. Infauna: 9 - oligocha- Turbellaria; 12 - oligochaete L. nigrescens. G. Around sponge
181
EQ :..I..,:.
El0
182
first horizon
is occupied
by a pelagic copepod
Cyclops kolensis, in the form of dense schools,
Fig.
(according to Bazikalova, One score of scale - 1 mm.
lb. S. parasiticus
modifications).
Fig. lc. E. violaceus
Fig. Id. A. spongicola modifications).
(original).
1945, with
Scale - the same.
(according
to Mazepova,
1978, with
visible to scuba divers. Few Cladocera are also present here. The second horizon is the nectobentos, or peron. The dominant organism here was an amphipod, Poekilogammarus erinaceus, which jumps over the surface of the sponge and in its vicinity; another organism abundant in this horizon is Baicalogammarus pullus. When a diver approaches, all movement ceases and animals hide among the branches. Shaking a sponge results in rapid escape of amphipods to other sponges. The sculpin Procottus jeittelesi (Sideleva et al., 1990) also resided in this horizon. It hides in-between branches, and leaves the sponge only when strongly disturbed, returning to the same place when danger is over. The richest horizon is that of epi-fauna, or epon. The branches surface is occupied by brightly colored amphipods of the species Spinacanthus parasiticus. Here they are much more abundant than on the base of sponges. Normally they sit quiet, firmly attached to the sponge by their strong claws. Only occasionally do they move, slowly, over the surface. It is difficult to make them leave their ‘home’, and they still cling to the sponge even after it is lifted to the surface. The amphipod Eulimnogammarus cruentus also hides at branching sites. Few Cyclops also move over the surface. The most interesting one, Acanthocyclops spongicola, clings to the sponge by hook-like spines or its extremities, or hides in oscula holes; its ‘harpacticoid’ form helps such 1978). Mollusks occur behavior (Mazepova, more often on the base of sponges, and rarely climb its branches. The same holds for few other, less abundant, groups of organisms. A horizon intermediate between epi- and infauna is occupied by Eulimnogammarus violaceus, which lives in holes gnawed in sponge bodies. The sizes of crustaceans strictly correspond to the sizes of their refuges; they sit coiled, their head and extremities facing the outlet. No infauna was found in the upper parts of the branches. In four out of five bases, juvenile oligochaetes (Nais sp.) were found (number:
183 Table 2. Composition, maximum abundance (N, number of animals) and maximum weight (B, mg) of different horizons of the fauna associated with L. baicalensis, according to data of the present study, and to evidence published earlier by other authors. + , no quantitative data; -, taxon absent; ?, presence of taxon doubtful; *, summary data presented in other sections of this row. (1) - identified by T. V. Akinshina; (2) - according to Chekanovskaya, 1962; (3) - according to Kozhov, 1931; (4) - identified by G. F. Mazepova; (5) - according to Mazepova, 1978, and personal observations; (6) - identified as a new species by W. W. Tachteew (1992); (7) - according to Sideleva et al., 1990, and personal observations. Taxons
Nectobenthos, peron
Epon
1
-
Turbellaria Nematoda Oligochaeta including: Nais sp. (1) Propappus Lycodrilus
B
N
B
N
B
N
B
2
3
4
5
6
7
8
9
-
-
8 8 10
7 0.1 12
30 * 43
78 * 26
-
-
+
+
+
+
1905 (2)
-
-
+
+
+
+
-
-
+
+
+
+
phreodriloides
Mich. 1905 (2) Lamprodrilus
?
nigrescens
-
Mich. 1903 (3) Polychaeta Manayunkia
baicalensis
(Nusbaum) 1901 Cyclopoida including: Cyclops kolensis Lill. 1901 (4) (plankton) Acanthocyclops
Baicalogammarus
?
?
+
+
+
* +
* +
*
+
+
19 +
1 +
-
-
-
-
+
+
+ + + +
+ + + +
+ + +
+ + +
-
26
14
-
80
72
-
228
8
141
-
-
1
13
spongicola
Maz. 1962 (5) Harpacticoida Ostracoda Cladocera Amphipoda
-
pullus
(Dybowsky) Brandtia
N
glandulosus
Michaelsen
Around sponge base
Infauna under sponge base
Epibenthos
1874
latissima
(Gerstfeldt)
1858
Crypturopus
rugosus
-
(Dyb.) 1874 Eulimnogammarus
cruentus
(Dorogostaisky) 1930 E. czerskii (Dyb.) 1874 E. grandimanus Bazikalova 1945 E. obsoletus Baz. 1945 E. violaceus (Dyb.) 1874 Hakonboeckia
(Dyb.) 1874
9
-
5 -
376 -
4 -
587 -
-
3 1 35
38 2 1372
5 8 -
50 6 -
-
2
3
3
18
strauchii
* 152
184 Table
(Continued)
2.
Taxons
Nectobenthos, peron
Epibenthos Epon
1 Heterogammarus
N
B
N
B
N
B
N
B
2
3
4
5
6
I
8
9
bifasciatus
(Dyb.) 1874 Pallasea
Around sponge base
Infauna under sponge base
-
15
-
-
-
50
-
-
-
-
-
-
18
*
*
59 *
1 *
1 *
cancellus
(Pall.) 1776 Poekilogammarus
erinaceus
Tachteew 1992 (6) Spinacanthus
parasiticus
238 -
Isopoda Baicalasellus
543 -
1895 22
20600 12
angarensis
(Grube) 1872 (Dyb.) 1884 Trichoptera, Baicalinini Chironomidae Mollusca
B. baicalensis
Megalovalvata
-
3 5
5 0.5
baicalensis
(Gerstf.) 1859 Baicalia sp. Pisces, Abyssocottidae Procottus
-
-
-
-
-
12 *
86 65
5210 375
*
*
-
-
356
2184
-
-
-
-
+
(3)+
-
-
,jeittelesi
(Dyb.) 1874
+
(7)+
xk lSE= 5.4-~ 3.8, weight: 3.5-t 2.6mg per 100 g of sponge weight, II = 5). The surface of sponge bases also contained A. spongicolu (number 1.3 + 0.9 per 100 g of sponge weight, y1= 5) but cyclops were only found in two sponges (weight 2.4-2.6 kg). The main endobiont of L. baicalensis is the unicellular alga Zoochlorella conductrix Brandt (Meyer, 1930) which inhabits cells of the surface layer of sponges and stains them green. The diversity of fauna living between the sponge base and the substrate depends on the presence and on the size of cavities. The smallest ones are inhabited by oligochaetes and Turbellaria, the larger cavities by oligochaetes, Turbellaria, Trichoptera, polychaetes, and small amphipods. The greatest cavities serve as refuges to gammarids (E. cruentus and EuZimnogammarus czerskii). The numbers of animals living under the base were
most variable; all these forms occurred in some, not all sponge specimens. Bases of sponges were encircled by numerous cases of Trichoptera (mostly empty, but some of them inhabited by larvae). Various animals belonging to the meio- and macrozoobenthos live in refuges between these cases. Only the numerous mollusks of the genus Baicalia do without refuges. The nectobenthic crustacean B. puZZus was one more component of this epifauna. A majority of species again did not occur in all sponge specimens; Brandtia Zatissima was in 83 %, E. cruentus in 61%, other amphipods in less than 50% of the sponges examined, but 100% occurrence was typical of S. parasiticus, E. violaceus and P. erinaceus. M. baicalensis inhabited only sponges living on the edge of the shallow-water platform. A correlation between the abundances of con-
185 sort and the mass of sponges was found for only were linear three species; some correlations (Fig. 2, 4), others were nonlinear (Fig. 3, 5-6). The main weight of S. parasiticus specimens taken in September at a depth of 6.4 m depended (Fig. 7.1) on the mass of the sponges in the range 700-2600 g; no such dependence (x? 1 SE = 10.8 + 0.5, n = 4) was found for sponges living on the shallow-water platform. When the bulk of data for September are united, they yield a weak correlation (Fig. 7.2). All data for November could be described by one straight line (Fig. 7.3). A regression calculated from the data of all observations is presented in Fig. 7.4. The mean mass of E. violaceus and P. erinaceus does not reliably depend on the mass of their sponges. Only few egg- bearing females of S. parasiticus have been found in September; they became dominant in November. Reproduction of E. violaceus finished before our studies as suggested by the absence of egg-bearing females and
-4.0
- d.5
0
LOG OF SPONGE WEIGHT,
0.5
kg
3. Dependence of the numbers of S. parasiticus on logarithm of sponge weight in September (depth 6.4 m) and November (depth 4.5 - 6.4 m): yk 139 = 385 + (421 k 81.9)* x, r= 0.82, P
a high abundance 1 mg.
of juveniles weighing less than
Discussion
WEIGHT Fig. 2. Dependence of S. parasiticus on sponge
OF SPONGE,
kg
the quantitative characteristics of weight: 1 - weight, September, depth 4.5m, yt 1.46=(7.61*0.68)* x - 0.13, r=0.99, P~0.01, n=5; 2 weight, September, depth 6.4 m, ykO.35=(2.12+0.31)* x - 0.28, r=0.93, P
Our data do not contradict those of earlier studies, as these (Kozhov, 193 1; 1962; 1963) gave a description of the fauna of the stony, shallowwater bottom of Lake Baikal, rather than of a sponge consortium. Bulk description also resulted in mixing the inhabitants of two sponges; the branched L. b. baicalensis, and the encrusting L. baicalensis littoralis Resvoj, 1936. This latter sponge occurs at depths below 1.0-1.5 m. It is rarely inhabited by S. parasiticus, but serves as a refuge for animals which are less abundant on the branched form; see Table 2, and the data of Kozhov, 193 1; Table 9. Unlike Kozhov (193 I), we failed to find ‘dense populations of polychaetes in oscula’ of L. baicalensis. As for the lower crustaceans, the most abundant were bottom Cyclopids, rather than ‘mass schools of harpacticids’. Moreover, the number of such crustaceans in our samples were small. The fauna residing on sponges is much less diverse than that residing around their base, although the sponge surface area is much larger than that of the substrate around their base.
186
0
0.5
WEIGHT Fig.
l-
of the quantitative 4. Dependence weight, yk 119=5.04+(91.6&40.3)*
characteristics x, r=0.53,
.
- 4.0
- 0.5
LOG OF SPONGE 5. Dependence rithm of sponge r= 0.85, P
Fig.
of the number weight: yk6.47 n = 19.
0
WEIGHT,
1:5
OF SPONGE,
2.0
25
kg
of P. erir~aceus on sponge weight in September P~0.05; 2 -number, y+51.2=7.53+(39.9*
Aquatic animals generally tend to inhabit any substrates on the bottom. Baikalian sponges, like other sponges (Gastic, 1988; Kohmoto et al., 1988) may excrete toxic substances, or repellents. Indeed, when sponges with their fauna were placed in small aquaria, all animals except S. parasiticus and E. violaceus died in a short time. Most workers (for reviews see Kharchenko & Protasov, 1981; Kharchenko et al., 1989) believe that all organisms found on determinant species are consorts. However, our studies reveal that the weight of a few species associated with sponges correlates with the weight of the sponge itself. Hence, one of us (R. M. Kamaltynov) suggests that true consorts are organisms which live exclusively on the edificator species, whereas non-
m d
4:o
233
at depth 4.5 - 6.4 m (n = 15): 17.3)* x, r=0.54, PiO.05.
consort species occur here only occasionally. Death of the edificator will result in death of its consort. For example, Newman (1989) reports that extinction of a single species of a tropical tree may result in the extinction of some 30 species of plants and animals supported by this edificator. L. baicalensis is not the only edificator among sponge consortia. The major consort - S. para-
./
0.5
kg
of E. violaceus on the loga=(21.7 k 3.31)* x -20.2,
LOG OF SPONGE
WEIGHT,
kg
Fig. 6. Dependence of the weight of E. violaceus on logarithm of sponge weight: 1 - September, y+ 0.15 = 0.39 + (0.43+0.09)* x, r=0.80, PcO.001, n=14; 2-November, y+O.26=0.79+(1.01~0.27)* x, r=0.94, P
187
WEIGHT
OF SPONGE , kg
Fig. 7. Dependence of the mean weight of S. parasiticus (September: a - depth 6.4 m, b - depth 4.5 m; c - November, both sites) n = 11; 2 - September, both on sponge weight: 1 - September, depth 6.4 m, y k 1.95 = (9.45 f 1.61)* x - 0.70, r= 0.89. P
siticus - may also inhabit L. b. littoralis, Baicalospongia bacillifera, Baicalospongia intermedia, and unidentified bark-shaped sponges growing on subvertical rocks down to 140 m depths. E. violaceus has been found on a bark-shaped sponge collected in Frolikha Bay (Northern Baikal) at 410 m. Co-evolution of the members of sponge consortia probably began before the separation of the edificator sponge species. This case is similar to others (Mazing, 1966), in which a group of species which identical composition of consorts was supposed to be the nucleus of a consortium. Our (R.M.K.) viewpoint is close to that suggested by Arnoldi & Lavrenko (1960; reference taken from Arnoldi et al., 1969) who classify relations within a consortium as primary when a phytophage uses given edificator plant species as a food source almost exclusively; as secondary, when there exists a network of food webs within a biocenosis; and, finally, as a satellite relation when the link between plants and animals is transient. Seasonal changes in the number and mean size of S. parasiticus specimens indicate that this species peaks around September, thus in summer, on the shallow-water platform. The bigger animals are found in September not only on the large
sponges but also on a smaller ones. This is the reason for the lack of the dependence of the mean weight of S. parasiticus on the mass of the sponges living on the shallow-water platform. In November part of the population of S. parasiticus migrates to deeper water, presumably to escape winter storms. During this period the correlation between mean weight of amphipods and the weight of sponges restores. This may indicate that predominantly large animals migrate. Mean size of E. violaceus does not correlate with sponge weight, evidently due to the high number variation of amphipods offspring. A better correlation could probably be found immediately prior to the reproduction, like it is the case for S. parasiticus. The mean size of P. erinaceus also does not correlate with sponge weight, but this resulted from a too small variance in size, because the animals studied belonged to one cohort. L. baicalensis grows approximately 1 cm per year (Gombreich, 1988). Hence, a sponge consorts remain unstable during the 20 years, and only thereafter stationary populations of consorts become settled; the weight of these populations becomes dependent on sponge weight. The sponges are at first occupied by younger amphi-
188 pods. Increase in the age leads to an increase of the number of larger animals.
Conclusion The Baikalian sponge L. baicalensis acts as determinant of a peculiar, typically baikalian consortium during part of the life cycle of the sponge. This consortium is here characterized quantitatively for the first time. New data have been obtained on the species diversity of consorts, and on seasonal dependence of the relations between sponges and their consorts. These consortia may play an important role in the cycling of nutrients in shallow waters and on steep rocky slopes of Lake Baikal.
Acknowledgements The authors thank Dr M. A. Grachev for help in manuscript preparation; Mr V. V. Romanov, scuba diver, for delivery of samples; Dr G. F. Mazepova for identification of Cyclopids and for information on ecology of meiobenthos; to M. S. Ivanova for technical assistance.
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