-1-

“Zooplankton Egg Bank Diversity in the

Sediments of two Lakes (Lake Bhalaswa and

Tajpur Pahadi Lake): A Comparative Field and

Experimental Investigation”

Submitted by

Debadityo Sinha (debadityo@gmail.com, +919210851757)

Roll No. N1003

Course: BSc(H.) Zoology

As part of ecology project Conducted under the supervision of Dr.

Ram Kumar

-2-
 CERTIFICATE

It is certified that this project is done in the Ecosystem research

laboratory; Department of Zoology, Acharya Narendra Dev college

(University of Delhi ) Govindpuri, Kalkaji, New Delhi 110019 by me,

Debadityo Sinha under supervision of Dr. Ram Kumar as part of B.Sc.

Hons. project.

Ram Kumar Debadityo Sinha

-3-
 Acknowledgement

At the outset I acknowledge the constant support and encouragement from the Principal Dr.

Savithri Singh who has been a source of inspiration to undertake any unsolved scientific

puzzle having evolutionary meaning. I am also grateful to her for providing facilities which

made the study possible.

I express my deepest gratitude Dr. Ram Kumar, Department of Zoology, Acharya Narendra

Dev College, University of Delhi, under whose guidance and supervision this work has been

completed. I am thankful to him for being extremely kind and generous in giving me the

opportunity to work with him. I am highly indebted to him for his support and constant

encouragement at all times. His in-depth knowledge, tactful criticism, dedication to work and

generosity shall continue to be a source of inspiration to me.

I also thank Victor R. Alekseev, Professor, Dr., Head of Department Taxonomy and

Systematics , Director, Zoological Institute of the Russian Academy of Sciences and

Innovative Center of Biotechnologies of St. Petersburg Association of Scholars and

Scientists.

I am thankful to all faculty members of the Zoology Department of Acharya Narendra Dev

College who has been constantly encouraging and supporting the project work regardless of

their busy schedule.

I would also thank to Mr Amit K. Awasthi, Senior Research Fellow at the ecosystem

research lab, Acharya Narendra Dev College for his help and encouragement.

I sincerely thanks to the staffs of Zoology Deperatment, Acharya Narendra Dev College, who

supported me throughout this project. I acknowledge the assistance provided by Mr. Tara

Dutt Sharma during sampling.

Debadityo Sinha

-4-
 CONTENTS

Abstract .…….……………….…… 6

Review Of Literature …………………………... 8

Materials And Method …………………………….17

Result ……………………………..28

References .….…………………………35

-5-
Abstract:

Central to the Darwinian theory of evolution through natural selection is

survival of the fittest, universally recognized as a potent evolutionary force.

The existence and survival of a species depend upon its adaptation to the

harsh and extreme environmental condition. One of the frequently occurring

harsh environmental conditions in fresh water ecosystem is desiccation.

Many aquatic invertebrates are able to overcome harsh environmental conditions

such as extreme low/high temperature, desiccation, intensive predation, low oxygen

etc. by producing special dormant stages that have evolved to overcome

catastrophic periods of their life cycle through millions of years of adaptation. Most

of the zooplankton taxa form resting stages which get accumulated in the sediment

of the water body. The accumulation of resting stages in bottom sediments forms a

reservoir of both the past and recent species composition in a particular water body

and acts as currency to be used under unfavorable condition (e.g. draught,

eutrophication and pollution etc.) hence also termed as egg bank for animals and

seed bank for plants. Therefore resting stage egg bank constitute an ecological

evolutionary reservoir that impacts population, community and ecosystem

processes. This allows researchers to reveal the past species composition in a

particular water body. Resting eggs of planktonic organisms from sediment are

extensively used to reconstruct historical information of the community composition.

As rotifers, cladocerans and copepods produce resting stages which can remain

viable for many decades, they have been now an important tool of ‘resurrection

ecology’.

-6-
Herein, I studied the zooplankton diversity in the sediment egg banks of two lakes:

Lake Bhalaswa and lake differing in trophic status, anthropogenic pressure and

draught condition. Bottom sediments from both the lakes were collected

synoptically. Zooplankton hatching experiments were done by restoring the aquatic

environment under indoor laboratory condition. Zooplankton emerged from the

sediment were identified to the lowest possible taxa. Observations were done at

regular intervals spanning a period of 3 months. A total of 20 species of

zooplankton (Rotifera- 10; Cladocera: 10) emerged from the sediment

collected from TAJPUR PAHADI LAKE whereas in the sediment collected from

Bhaleswa I recorded 13 species (Rotifera- 9; Cladocera: 4; Ostracoda: 1). Results

obtained in my study indicate that the bottom sediment is evolutionary archive of the

zooplankton community. My study also indicates that the bottom sediment may be

used as started culture of fish food in aqua culture industry.

-7-
Introduction:

Many lakes and ponds are subjected to eutrophication, draught (e.g. Lake
Bhalaswa, Lake Badkhal) and heavy anthropogenic pressure (Tajpur Pahadi lake).
Consequently most water bodies are becoming temporary habitats for fresh water
communities. Temporary waterbodies are widespread in tropical regions and in
recent years the abundance and ecological importance of these ephemeral
waterbodies have drawn attention ecologically (Williams, 1987). These habitats are
transient for the aquatic organisms, shifting from inhabitable to uninhabitable, and
then back to inhabitable. Aquatic organisms for continual existence must either
disperse to other habitats (forming spatial metapopulations) or remain dormant for
the inhabitable period. The most work done in aquatic ecology have been directed to
permanent (ie. Hydroperiod> 1 year) lentic and lotic waters. The lack of attention
may be due to the fact that temporary waters are often too small for fishing,
recreation or holding water for agricultural or other uses. Without obvious utility to
man they have been largely ignored. Without obvious utility to man they have been
largely ignored, hence having less basis for the protection of these unique and
endangered habitats (Kumar & Prasad, 2004). Concern is expressed for the state of
the art of temporary aquatic habitats, as our exploitation and destruction of such
habitats continue globally. Habitat destruction is a primary threat to invertebrates in
general, and in particular invertebrates with specific requirements (Collins,
1991).Keeping in view the conservation biology of aquatic organisms, the study of
such habitats is essential (Kumar & Prasad, 2005). In order to understand the
adaptation and survival strategies of zooplankton it is important to understand their
life history attributes.

-8-
Reproduction and Life Histories of Zooplankton:

Rotifers and Cladocerans reproduce by diploid female parthenogenesis (females

produce young without mating) throughout much of the growing season. During

periods of stress (low, food, light, temp., crowding) meiosis occurs leading to the

formation of haploid males and females.

The haploid males and females mate and the fertilized, diploid eggs are produced

which are encased in a heavy shell or ephippium. These resting eggs are shed

when the female molts and over winter in the sediments and will develop into amictic

diploid females

These resting eggs are very durable and can remain a viable “seed bank” of

organisms for decades.

After the resting or parthenogenetic eggs hatch, the Cladoceran undergoes a

number of rapid instar stages which differ mainly in size. Temperature is the main

factor that determines growth rate. A Daphnia gives birth to her first brood in about

48 hours at a water temperature of 25ºC. However, the fecundity or numbers of

young/brood are more influenced by availability and quality of food.

Copepods are bisexual and carry their fertilized eggs in external egg sacs. As with

the Cladocera, Copepod egg production is determined by temperature and clutch

size is determined by food quality. Development of a Copepod from egg to adult is

much longer than compared to Rotifers and Cladocerans. The degree to which a

copepod population enters diapause, the instar stage when diapause occurs and the

location (water, sediments) of diapause is highly variable among species.

-9-
Fig. 3. Schematic diagram showing dry and hydroperiod of a tropical temporary

aquatic habitat and survival strategies of the inhabitants. ( Kumar & Prasad, 2004).

- 10 -
Resting Eggs:

Micro-organisms mostly (here zooplankton) that inhabit temporary variable

aquatic environments or ephemeral or seasonal lakes follow some strategies to

survive the unsuitable conditions like drying, temperature variations, competition,

predation etc. They bridge the periods of unsuitable conditions by production of

resting eggs or diapause stages (Cáeceres, 1997; Brendonck et al., 1998a). This

stage is a state of suppressed development (Danks, 1987). In normal dormancy

the organisms resume their metabolism and other development as soon as

suitable conditions persists again, but in diapause the organism does not

resume its development until its diapause is broken, even on the arrival of

suitable conditions. The conditions which are necessary for breaking the diapause

depend mostly on the species to which the egg belongs and often vary even among

populations of same species (Wyngaard, 1988). Generally the eggs either settle at

the bottom of lakes sediment or remain freely floating. Photoperiodism and light

intensity are the main stimulus. With time, some eggs hatch but most remain

unhatched, and they accumulate and contribute to the existing eggs and forms the

“Egg Bank”. Egg banks are a common feature in variable water systems such as

shallow lakes, ephemeral pools (de Stasio, 1989). After the ephemeral pools get

filled by rain some of them float and hatch and remaining remain as egg bank in the

sediment.

Most of the zooplankton taxa form resting stages which get accumulated in

the sediment of the water body. In a particular water body, this forms a reservoir of

both the earlier and recent species composition. As resting eggs in the sediment can

- 11 -
remain viable for decades or centuries, the longlived dormant stages of zooplankton

additionally offer a unique source of living historical information (eg. Weider et al.,

Hairston et al., 1999a,b; Kerfoot et al., 1999; Cousyn et al., 2001; Limburg & Weider,

2002), the study of which was denoted as resurrection ecology by Kerfoot et al.

(1999). Resting stage egg bank thus constitute an ecological evolutionary reservoir

that impacts population, community and ecosystem processes (Brendonck & De

Meester, 2003).

All of the zooplankton species do not produce resting eggs or dormant stages

and those produced do not essentially survive for long (Hairston, 1996). The

zooplankton known to produce resting eggs or dormant stages are the rotifers (eg.

Brachionus, Asplanchna, Rotaria) ; cladocerans ( eg. Daphnia, Moina,

Cereidaphnia).

In monogonont rotifers, the egg has yolk and the embryo get surrounded by a

thick multilayered shell. The eggs are spherical and get sink in the sediment. The

dormant copepodite or adult stages in cyclopoid and harpacticoid copepods, are

often encased in a transparent gelatinous cyst with an outer layer of mud particles

(Fryer & Smyly, 1954; Smyly, 1961, 1967; Elgmork, 1962). In calanoid copepods,

resting eggs are spherical and sink freely to the bottom after deposition (Viitasalo &

Katajisto, 1994).

In cladocerans, the daphnid eggs are well known for their structure. Here, 1 or

2 resting eggs are encapsulated by the carapace valves and forms a structure called

“ephippium” which is shed at molting. The ephippium get attached to the floating

- 12 -
debris or algae and thereafter sink to the bottom after a brief floating phase

( Carvalho & Wolf, 1989).

Large pools of dormant stages exist in the active egg bank in all groups of

freshwater zooplankton, as shown in rotifers (Nipkow, 1961; Hairston et al., 2000),

large branchiopods (Hildrew, 1985; Thiéry, 1997; Brendonck & Riddoch, 2000b),

calanoid copepods (de Stasio, 1989; Hairston et al.; 2000), cyclopoid copepods

(Hairston et al., 2000), and cladocerans (Lampert & Krause, 1976; Smyly, 1977;

Herzig, 1985; Carvalho & Wolf, 1989; review in Hairston, 1996; Cáeceres, 1998b;

Cousyn & de Meester, 1998; Hairston et al., 2000). Hatching is mainly restricted to

the resting eggs in the upper 2 cm of the sediment (Herzig, 1985) that is indicated

as the ‘active egg bank’ according to Cáeceres & Hairston, (1998). The largest

fraction of viable eggs occurs in the top 4-6 cm of the sediment (Herzig, 1985;

Carvalho & Wolf, 1989; Hairston & van Brunt, 1994; Cáceres, 1998b).

Factors affecting egg bank and its hatching:

In permanent relatively shallow aquatic habitats, photoperiodism (reviewed in

Fryer, 1996) and temperature have been shown to be important for inducing or

breaking diapause. In deep lakes, temperature and the gaseous state (Fryer, 1996)

do the work. In shallow ephemeral lakes temperature changes and changes in

osmotic and oxygen values during an inundation cycle are the most important

indicators of the suitability for growth and reproduction (Brendonck, 1996). In rotifers,

cladocerans and copepods, most hatching takes place in spring (Herzig, 1985; Wolf

& Carvalho, 1989; de Stasio, 1989, 1990; Cáceres, 1998a, b; Hairston et al., 2000).

- 13 -
 The stimulus that triggers the production of diapausing eggs in the above

organisms usually acts prior to the onset of deteriorating environmental conditions

and includes token stimuli such as photoperiodism, population density, predator

density, or food (Hutchinson, 1967; Stross, 1969; Kleiven et al., 1992; Gilbert, 1995;

Hairston & Kearns, 1995; Piercey & Maly, 2000; Alekseev & Lampert, 2001).

Eutrophication also plays an important role in increased resting egg production as,

increased Daphnia ephippium densities since 1940s in the sediments of the

Keweenaw Waterway were most probably related to increased eutrophication

(Kerfoot et al., 1999). Also loss of drying may also result in loss of biodiversity within

a system (Nielsen et al., 2000).

Advantage Resting Eggs: A Process Must Happen

• The stored egg bank acts as buffer against years when reproduction is low or

even prevented due to harsh conditions, and allows repopulation of the

habitat in the future (De Statio, 1990).

• In addition to provide storage of individuals, dormant propagule pools also

can maintain genetic variability of the population and affect the rate of

evolution. (Templeton and Levin 1979; Coleman 1983, Hairston and De Statio

1988).

- 14 -
 • In spite of their differences in phylogeny and organism size, the zooplankton

egg bank and aquatic plant seed bank exhibit similar characteristics that allow

resilience (Brock et al. 2003).

Overall, the egg bank plays a critical role in allowing species living in variable

habitats to adapt to the changing environment in which they live. However except

two published papers (Kumar and Prasad 2004 a, b) I did not get any information on

egg bank or seed bank diversity in the sediment. This prompted to me to study the

egg bank diversity in the sediments of two lakes of Delhi differing in trophic status,

anthropogenic pressure and hydroperiod.

The primary aim of my project was to carry out a short duration study to elucidate

species richness and time taken to hatch from the sediment of the two lakes and to

establish the role of eutrophication on egg banks.

- 15 -
Materials and Methods:

Study lakes:

I selected two lakes located in the outskirts of Delhi, having contrasting

anthropogenic pressure, trophic status and hydroperiod.

The Lakes chosen were Tajpur Pahadi Lake, Badarpur and Bhalaswa Lake,

Bhalaswa.

- 16 -
DESCRIPTION OF THE TAJPUR PAHADI LAKE

Common Name: “Tajpur Pahari Khaan” (Fig. 2)

Co-ordinates: 28º29’56”N 77º18’29”E
Location: Tajpur Pahari, Badarpur. New Delhi
Landmark: Badarpur Thermal Power Station, Delhi.
Pin: 110 044

The area is under “New Delhi Municipal Corporation”. The place was actually a
stone quarry from which it derived its name ‘Khaan’. The area situated just beside
the “Badarpur Thermal Power Station” was used extensively for extracting stones
until the underground water came out. The area was then left as it was since 70’s.
Gradually the place turned into a very deep lake with accumulation of more rain
water underground and some local sewage. The total area of the place is
approximately 30 acres. With time the lake became a habitat of many aquatic
animals like waterfowls, damselflies, fishes etc. Till now no-one had studied the
waterbody and there is not single written information about its biodiversity. The lake
is now at verge of vanishing because of developmental projects undertaken by as
the Municipal corporation. The water is being extracted land filling is being done for
the last 2 years. The lake is now surviving through several small shallow ponds and
puddles which are the only niche to the aquatic animals which once used to occur in
the lake. It is suspected that the lake will be permanently extinct within few months.
Water criteria:
Water color- straw color
Highly turbid (secchi disc measuremt 30 cm)
Phosphate level: > 600 μgL-1
Nitrate level: >800 μgL-1

- 17 -
Fig.2. Road map of Tajpur Lake ( source:http// www. mapmyindia.com)

- 18 -
DESCRIPTION OF THE BHALASWA LAKE

Common Name: “Bhalaswa Jheel /Bhalaswa Recreational & Sports Complex” (Fig.

3)

Co-ordinates: 28º44’32”N 77º10’17”E

Location: Bhalaswa, New Delhi

Landmark: Bhalaswa Golf Course; Yamuna Biodiversity Park, Delhi.

Pin: 110043

The lake is used for water sports activity. The lake unlike the Tajpur Pahadi

Lake is a permanent and very old lake. The water is not as eutrophic as in the Tajpur

Pahadi Lake. The lake is under Delhi Development Authority and recently it was

declared Recreational Complex by the Delhi Govt. in 2003. The lake is to be used

for sports in the 2010 commonwealth game.

Water criteria:
Water color- Clean / light green
Turbidity (secchi disc measuremt 80 cm)
Phosphate level: > 300 μgL-1
Nitrate level: >500 μgL-1

- 19 -
Fig.3. Road map of Bhalaswa Lake ( source:http// www. mapmyindia.com)

- 20 -
A

.Fig. 4. Image of the site, “Tajpur Pahadi Lake”. ( http//www.wikimapia.com)

Fig.5. Satellite image of the Bhalaswa lake. ( source::http// www. wikimapia.com)

Table : Comparison of the two lakes with respect to their current status

TAJPUR PAHADI LAKE BHALASWA LAKE

Less than 50 years old More than 100 years old
AGE
TROPHIC Water highly eutrophic Water not much eutrophic
STATUS
HUMAN Sewage and drainage in Water source natural, used
IMPACT &
large extent for sports and recreation.

- 21 -
WATER Lake is being landfilled Water level in the lake is
SOURCE
by the MCD being maintained by the

DDA
A stone quarry turned to

HISTORY the lake Natural lake

Sample collection:

We followed same methods for collecting the soil samples. The sediments were collected

from the site with the help of a PVC sediment corer of diameter 3.08 cm or 2 inches and

length 27 cm. the corer was pushed vertically upto 4 cm in the lake bottom and then taken

out with blocking its mouth. The soil samples were taken from 4 randomly selected places

within the lake. Soil samples were packed in plastic bags and covered with aluminium foils

to avoid any effect of light.

Zooplankton in the water column was filtered through a plankton net with 53

micrometer mesh size. Photographs of the sediment and water sample collection have

been shown in Fig. 9-13. Sediment samples so collected were brought to the laboratory sun

dried and stored at 4 degree centigrade temperature till the hatching experiment was

conducted.

Site photos of the Tajpur Pahadi Lake

- 22 -
- 23 -
Fig. (4-8). Site pictures of the Tajpur Pahadi Lake

- 24 -
Fig. 9 . Sediment collection from the Tajpur Pahari Khaan.

Fig. 10 . Collection of zooplankton from the water column in the Tajpur Pahari

Khaan.

- 25 -
Processing of the samples:

The wet soil samples were then allowed to sundry for 4 consecutive days. After the

soil samples were dried completely, ground them into powder form with hand so as

to protect the eggs from any mechanical pressure. The samples were then mixed

thoroughly into a single mixture. From this sample, 4 subsamples were taken

weighing 50g each. Then each subsamples were incubated with autoclaved tap-

water in a 250 ml beaker in ambient temperature, the water depth in the beaker was

- 26 -
maintained 5 cm. The samples were then observed everyday until asymptote was

reached. The water was decanted in a petriplate thropugh a 30µm mesh and

observed under stereoscopic microscope ( Magnüs MSZ-Bi). The animal hatched

were then taken out and allowed to grow in a separate beaker containing Chlorella

vulgaris till it become adult enough to be identified. The animal was then preserved

with 5% formalin. Then it was identified with the help of identifion keys.

Identification of the animals:

The identification keys consulted for this study were “Koste, (1978);

Edmondson (1959)” for the rotifers and “Michael and Sharma, 1988; Edmondson

(1959)” for the cladocerans.

RESULTS:

As the project aims at comparing only the number of species that had emerged from

the sediment egg bank of the zooplankton, we had ignored any measurement of

density or embryo number. As the lakes had different history- Tajpur lake was a

newer one (less than 50 year) and formed due to ground and sewage water, the

water was eutrophic as compared to the Bhalaswa lake which is a natural and and

very old lake. The number of species emerged from the sediment of the Tajpur lake

- 27 -
was sooner ( 3rd day of incubation) than that of the Bhalaswa lake (6 th day of

incubation); and also the number of species of cladocerans which emerged from the

egg banks were more in the case of Tajpur pahadi lake as compared to Bhalaswa

lake. The reason may be due to the highly eutrophic conditions which induce

diapause or resting stages in the cladocerans (as in daphnia ephippium, Kerfoot et

al., 1999) in the case of the Tajpur pahadi lake. In either of the lakes the diversity of

zooplankton was higher in the sediment egg bank than that in the water column. A

total of 20 species of zooplankton (Rotifera- 10; Cladocera: 10) emerged from

the sediment collected from the Tajpur Pahadi lake whereas in the sediment

collected from the Bhalawa lake I recorded 13 species (rotifera- 9; Cladocera:

4; Ostracoda: 1)

Table : Zooplankton species emerged from the sediment collected from Tajpur

Pahadi Lake

DAYS

1
2
3

- 28 -
6

10

11

12

15

90

- 29 -
Table : Zooplankton species emerged from the sediment collected from Bhalaswa

Lake

DAY

1
2
3

7
8
9
10
11
12

13

14

15

20

45
59

- 30 -
 Resting Egg Of Daphnia An Ostracod hatched

SOME MICROSCOPIC PICTURES TAKEN

Resting Egg Of Daphnia An Ostracod hatched

- 31 -
Pleuroxus aduncus hatched

My short term study also proves that the lake sediments act as an evolutionary

archive of the diversity of the zooplankton. Zooplankton in the past and their history

may be constructed with the help of resting stages analyses. Resting stages do not

only help inh survival under unfavorable condition but also act in reducing

competition and over crowding in lakes and ponds by dispersal of egg bank. Resting

stages has applied aspects also which can be explained under following sub

headings:

Applied aspects of Resting Eggs analyses:

AQUACULTURE:

Better knowledge of these processes may ultimately result in the manipulation

of these physiological phases and in more predictable hatching results in the

application of resting eggs in aquaculture and aquatic toxicology.

Today rotifers serve as the only food offered for larvae of many species of marine

and fresh water fish. Rotifers act as first food during the early developmental stages

of fish larvae. Availability of live food is very unpredictable because of difficulty in

culture however resting stages may act as buffer food which may be used during

- 32 -
developmental stages of fish larvae. It has been reported and also verified that

40,000-1,00,000 rotifers are required for raising one fish larva to age of 30-34 days

( Kafuko and Ikenoue, 1983).

But the main problem with rotifers are their storage and preservation.

Therefore, the ability of producing resting eggs can be used to preserve them

and then these may be used in aquaculture (Lubzens, 1989).

RESSURECTION ECOLOGY:

Many of the resting stages are very resistant and may survive extreme

temperatures, high pressure, or drought and remain viable for several decades. This

allows researchers to explore the earlier species composition of a particular water

body.

Identifying the resting stages can assess species diversity, where possible on the

basis of their morphology, or by hatching sediment under a range of laboratory

conditions. The reconstruction of micro-evolution using resting egg banks as

archives of past genetic changes involves the integration of different techniques and

approaches, such as radio-dating, historical reconstruction, the use of molecular

tools and experimental quantitative genetics (Hairston et al., 1999; Cousyn et al.,

2001).

RESEARCH:

It is still unclear what specific factors control the duration of dormancy for these

species what are triggering factors of production of resting stages and what are the

- 33 -
optimum condition of emergence of animals from restring stages. Therefore question

yet to be solved is “How Long To Rest????”

References

Alekseev, V. & W. Lampert, 2001. Maternal control of resting egg production in

Daphnia. Nature 414: 899–901.

Brendonck.L & De Meester.L. 2003. Egg banks in freshwater zooplankton:

Evolutionary and ecological archives in the sediment. Hydrobiologia 491: 63-

84

Brendonck, L. & B. J. Riddoch, 2000b. Egg bank dynamics in anostracan desert rock

pool populations (Crustacea: Branchiopoda). Arch. Hydrobiol. 148: 71–84.

Cáceres, C. E., 1997. Dormancy in invertebrates. Inv. Biol. 116:

371–383.

Cáceres, C. E. & N. G. Hairston, Jr., 1998. Benthic-pelagic coupling

in planktonic crustaceans: the role of the benthos. Arch.

Hydrobiol. 52: 163–174.

Cáceres, C. E., 1998a. Seasonal dynamics and interspecific competition in Oneida

Lake Daphnia. Oecologia 115: 233–244.

- 34 -
Cáceres, C. E., 1998b. Interspecific variation in the abundance, production, and

emergence of Daphnia diapausing eggs. Ecology 79: 1699–1710.

Carvalho, G. R. & H. G.Wolf, 1989: Resting eggs of lake-Daphnia.I. Distribution,

abundance and hatching of eggs collected from various depths in lake

sediments. Freshwat. Biol. 22: 459–470.

Coleman, A.W. 1983. The roles of resting spores and akinetes in chlorophyte

survival, p. 1-22. In G.A. Fryxell (ed.), Survival strategies of the algae.

Cambridge.

Collins.N.M. 1991. Insect conservation-priorities for the future. Antenna. 15: 73-78

Cousyn, C. & L. de Meester, 1998. The vertical profile of resting egg banks in natural

populations of the pond-dwelling cladocerans Daphnia magna. In Brendonck,

L., L. de Meester, & N. G. Hairston, Jr. (eds), Evolutionary and Ecological

Aspects of Crustacean Diapause. Arch. Hydrobiol. 52: 127–139.

Cousyn, C., L. de Meester, J. K. Colbourne, L. Brendonck, D. Verschuren & F.

Volckaert, 2001. Rapid, local adaptation of zooplankton behavior to changes

in predation pressure in the absence of neutral genetic changes. PNAS 98:

6256–6260.

Danks, H. V., 1987. Insect Dormancy: An Ecological Perspective. Tyrell Press,

Gloucester, 439 pp.

de Stasio, B. T., Jr., 1989. The seed bank of a freshwater crustacean: copepodology

for the plant ecologist. Ecology 70: 1377–1389.

- 35 -
de Stasio, B. T., Jr., 1990. The role of dormancy and emergence patterns in the

dynamics of a freshwater zooplankton community. Limnol. Oceanogr. 35:

1377–1389.

Edmondson. W. T. 1959 Freshwater Biology. 2nd ed. Wiley & Sons. New York. Pp.

1248.

Elgmork, K., 1962. A bottom resting stage in the planktonic frewhater copepod

Cyclops scutifer Sars. Oikos 13: 306–310.

Fryer, G., 1996. Diapause, a potent force in the evolution of freshwater crustaceans.

Hydrobiologia 320: 1–14.

Fryer, G. & W. J. P. Smyly, 1954. Some remarks on the resting stages of some

freshwater cyclopoid and harpacticoid copepods. Ann. mag. Nat. Hist. Ser.

12: 65–72.

Gilbert, J. J., 1995. Structure, development and induction of a new diapause stage in

rotifers. Freshwat. Biol. 34: 263–270.

Hairston, N. G., Jr., 1996. Zooplankton egg banks as biotic reservoirs in changing

environments. Limnol. Oceanogr. 41: 1087–1092.

Hairston, N. G. & C. M. Kearns, 1995. The interaction of photoperiod and

temperature in diapause timing: A copepod example.Biol. Bull. 189: 42–48.

Hairston, N. G., Jr., L. J. Perry, A. J. Bohonak, M. Q. Fellows, C.M. Kearns & D. R.

Engstrom, 1999b. Population biology of a failed invasion: Paleolimnology of

Daphnia exilis in upstate New York. Limnol. Oceanogr. 44: 477–486.

- 36 -
Hairston, N. G., Jr., W. Lampert, C. E. Cáceres, C. L. Holtmeier, L. J. Weider, U.

Gaedke, J. M. Fischer, J. A. Fox & D. M. Prost, 1999a. Rapid evolution

revealed by dormant eggs. Nature 401: 446.

Hairston, N. G., Jr., A-M Hansen & W. R. Schaffner, 2000. The effect of diapause

emergence on the seasonal dynamics of a zooplankton assemblage.

Freshwat. Biol. 45: 133–145.

Hairston, N. G., Jr. & R. A. Van Brunt, 1994. Diapause dynamics of two diaptomid

copepod species in a large lake. Hydrobiologia 292/293: 209–218.

Herzig, A., 1985. Resting eggs – a significant stage in the life cycle of crustaceans

Leptodora kindti and Bythotrephes longimanus. Verh. int. Ver. Limnol. 22:

3088–3098.

Hutchinson, G. E., 1967. A Treatise on Limnology. Volume II. Introduction to Lake

Biology and the Limnoplankton. John Wiley and Sons, New York.

Lampert, W. & I. Krause, 1976. Zür Biologie der Cladocere Holopedium gibberum

Zaddach imWindgefallweiher (Schwarzwald). Arch. Hydrobiol. 48: 262–286.

Lubzens E., B. perry, S. Euteneuer, and R. Berghahn. 1989. Preservation of rotifers

for use in aquaculture. Spec. Pub. Europe. Aquacult. Soc., Bredene, Belgium.

(in press)

Nielson D.L., Hillman T.J., Smith F.J. & Shiel R.J., 2002. The influence of seasonality

and duration of flooding on zooplankton in exerimantal billabongs, River

Research and Applications, 18, 227-237.

Nipkow, F., 1961. Die Rädertiere im Plankton des Zürichsees undihre

Entwicklungsphasen. Schweiz. Z. Hydrol. 23: 398–461.

- 37 -
Kafuko T. and H. Ikenoue. 1983. Modern methods of aquaculture in Japan.

Developments in aquaculture and fisheries science. 11. Kodansha Ltd.,

Tokyo, and Elsevier Scientific Publ. Company. Amsterdam. 216 p.

Kerfoot, W. C., J. A. Robbins & L. J.Weider, 1999. A new approach to historical

reconstruction: combining descriptive and experimental paleolimnology.

Limnol. Oceanogr. 44: 1232–1247.

Kleiven, O. T., P. Larsson & A. Hobæk, 1992. Sexual reproduction in Daphnia magna

requires three stimuli. Oikos 65: 165–206.

Koste. W. 1978. Rotatoria. Gebrûder Borntraeger. Berlin. Stuggart. Pp 234.

Kumar.R. and Prasad.P.2004a. Ephemeral aquatic habitats and zooplankton

diversity with reference to wetlnds of Sonipat District. Haryana. Proceedings

of the national Symposium on Perspective of Biodiversity Ecophysiology and

Conservation of Freshwater animals. Narendra Publishers. New Delhi. India

Kumar.R. and Prasad.P.2004b. Resting egg diversity and zooplankton emergence

patterns from soil egg bank of recently dried waterbodies. The Ekol. 2, 1-2:

19-30

Michael. R. G. and Sharma. B. K. 1988. Fauna of India: Indian Cladocera

( Crustacea: Branchiopoda: Cladocera). Zool. Surv. India. Govt. of India. Pp

262.

Piercey, D.W. & E. J. Maly, 2000. Factors influencing the induction of diapausing egg

production in the calanoid copepod Diaptomus leptopus. Aquat. Ecol. 34: 9–

17.

- 38 -
Smyly, W. J. P., 1961. The life-cycle of the freshwater copepod Cyclops leuckarti

Claus in Esthwaite Water. J. anim. Ecol. 30: 153–171.

Smyly, W. J. P., 1967. A resting stage in Cyclops dybowskii Lande (Crustacea:

Copepoda). Naturalist 92: 125–126.

Smyly, W. J. P., 1977. A note on the resting egg of Holopedium gibberum Zaddach

(Crustacea: Cladocera). Microscopy (Lond.) 33: 170–171.

Stross, R. G., 1969. Photoperiod control of diapause in Daphnia. II. Induction of

winter diapause in the arctic. Biol. Bull. 136:264–273.

Templeton A.R. & Levin D.A. (1979) They evolutionary consequences of seed pools.

The American Naturalist, 114, 232-249.

Viitasalo, M. & T. Katajisto, 1994. Mesozooplankton resting eggs in the Baltic Sea:

identification and vertical distribution in laminated and mixed sediments. Mar.

Biol. 120: 455–465.

Watanabe T.C. Kitajima, and S. Fujita. 1983. Nutritional values of live organisms

used in Japan for mass propagation of fish. A review. Aquaculture 34:115-143

Williams D.D. 1987. The ecology of temporary waters. Timber Press. Portland. Pp.

205

Wyngaard, G. A., 1988. Geographical variation in dormancy in a copepod: evidence

from population crosses. Hydrobiologia 167/168: 367–374.

- 39 -