Nordic Society Oikos

Scale-Dependence of Food Web Structures: Tropical Ponds as Paradigm

Author(s): Debal Deb
Source: Oikos, Vol. 72, No. 2 (Mar., 1995), pp. 245-262
Published by: Blackwell Publishing on behalf of Nordic Society Oikos
Stable URL: http://www.jstor.org/stable/3546227 .
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OIKOS 72: 245-262. Copenhagen1995

Scale-dependenceof food web structures:tropicalponds as

paradigm
Debal Deb

Deb, D. 1995.Scale-dependence
of food web structures:
tropicalpondsas paradigm.
Oikos 72: 245-262.

Two sets of time-specificfood webs were constructedusing all known details of

trophicbiology of the 87 componentspecies found in two tropicalfreshwaterpond
systems. Analysis of the webs showed that severalfood web statisticswere scale-
dependent.To examinethe robustnessof this finding, 11500 computerizedanalog
webs werebuiltuponthe specifiedrulesof non-randomtrophiclinks (L) betweenthe
organismsfoundin the ponds.The structuralpropertiesof the analogwebs, like the
recentlypublishedspecies-richwebs, disagreedwith severalgeneralizationsmadeby
food web theoristsoverthe lasttwo decades.Themagnitudeof L andthe frequencyof
long food chainsincreasedwith S. Proportionsof top, intermediateandbasalspecies
showedstrongnonlinearvariationwithS. Themeannumberof linksbetweendifferent
trophiclevels seemedto be scale-dependent, thoughapparentlyscale-invariant for the
pond webs. However,the relativeproportionsof link fractionsappearedto be scale-
invariant,with the intermediate-basal links most numerous.The meanpredator-prey
ratioappearedto be the most scale-invariant of all the web properties,but the scale
invarianceof the ratiomay not be directlyderivedfrom species fractions.The study
representsthelargesttropicalaquaticfood web database,andindicatesthat(a) as many
detailsof inter-specificinteractionsas possiblewithina web arenecessaryfor unravel-
ing the actualcomplexityof real ecosystems,and (b) computerizedanalogsof real
webs, incorporatingall necessarydetails of interactions,may be fruitfullyused to
ensurerobustnessof conclusionsaboutthe web features.

D. Deb, World Wide Fund for Nature - India, Eastern Region Office, Tata Centre 5th
Floor, 43 Chowringhee Road, Calcutta 700 071, India.

Over the last two decades, ecologists conjectured that
several food web features were invariant. That is, the
number of trophic levels; the number of trophic interac-
tions (L) for each species, or the linkage density (d); the
fractions of top predators (species with no predators in
the web), intermediate species, and basal species (au-
totrophs and detritus); the proportions of trophic links
amongst the total species (S); and the ratio between
predator to prey species, did not depend on S (Pimm
1982, Briand and Cohen 1984, 1987, Cohen and Briand
1984, Cohen and Newman 1985, Cohen et al. 1986,
1990). These generalizations were built on the descrip-
tions of webs catalogued by Briand and Cohen (Briand
1983, Cohen et al. 1990), which have recently been
criticized for a number of methodological difficulties
(Paine 1988, Sprules and Bowerman 1988, Lawton 1989,
Schoener 1989, Winemiller 1989, Polis 1991). As more
species-rich webs became available, it became evident
that most web properties are scale-dependent. Thus,
Pimm et al. (1991) suggest that trophic linkages typically
increase with S, describing the power function relation-
ship
L = k S+E, (1)
where E = 0.3 to 0.4, and Martinez (1992) and Havens

Accepted 12 August 1994

Copyright? OIKOS1995
ISSN 0030-1299
Printedin Denmark- all rightsreserved

OIKOS 72:2 (1995)

245
t-.)
.I\. Table 1A. Qualitativeestimateof zooplanktonin the culturepond.
CodeOrganism 1985 1986
Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb

Copepoda
1 +++ ++ I
Mesocyclops hyalinus ++ + +++ ++ ++ ++ + ++ +
2 M. leuckarti -++ ++ ++ +
3 Allodiaptomus raoi +-
4 Heliodiaptomus sp. +
6 Paradiaptomus sp. ++- -++ ++? ++
6 Rhinediaptomus sp.
7 Tropocyclops sp.
8 +-
Nauplii +++ ++ ++ ++ ++ ++ ++ ++
+
Rotifera
+
9 Asplanchna priodonta +++ ++ $++ ++
+
++
10 Asplanchnopus sp. -+- ++ ++ -
I1 Brachionus angularis +++ ++ $+ ++ ++ +
12 B. bidentata ++ ++ +?
++
13 B. budapestinensis +++ ++
14 B. calyciflorus ++- ++ ++--
++? ++ +?
15 B. caudatus ++- 4- + ++
16 B. falcatus ++
++--
17 B. quadridentatus -+ +
+
18 B. rubens
19 Conochiloides natans ++
20 -4-4 ++
Dicranophorus forcipatus ++
21 ++ -
Eosphora sp.
22 Filinia longisetta +
+-- ++ ++ ++
23 Hexarthra sp. -+- -+ +3- ++ ++
24 Horaella sp. ++
++
25 Keratella cochlearis
26 K tropica - +-+
-++ ++ -++-
++ 3++ +
27 Lecane (L.) luna 3+ + ++
28 L. (Monostylla) bulla ++ + ++ ++ +?
29 Lepadella sp. ++ ++- ++
30 Polyarthra multiappendiculata --+ ++
31 Pompholyx sp. ++ ++ ++
+
++?
Cladocera ++-
32 ++ +- +
Ceriodaphnia cornuta +++ ++t ++ -++ + 4+ +
33 Daphnia carinata ++
+ ++ +
34 D. lunholtzi -+ ++ ++ -+ +
35 Moina micrura duhia ++- ++ ++
++
0 36 Nauplii +++ ++ 3+ ++ ++ + 4+ +
0 Ostracoda
37 Cypridella sp.
38 Eucypris sp. - +-
- --

A
+ present;-absent. Data adoptedfrom Deb (1989).
o
Table 1B. Qualitative estimate of non-zooplanktonic organisms in the culture pond.
O
Code Organism 1985 1986
Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb

Phytoplankton
39 Chlaymydomonas sp. +++ ++ ++- ++ ++ ++ -+~ -+3 + ~+ ++3 +
40 Chlorella sp. +++ -- +-- -+3 + ++- +
41 Chlosterium sp. ++ -+ +
42 Diatona sp. +++ ++ ++3 +- +- ++$ +
43 Difflugia sp ----++ ++3 ++ +
44 Euglena sp. -++ -+ ++? ++-t ++ +
45 Gyrosigma sp. +++ +- + ++-
46 Scenedesmus sp. -+- -+ ++- ++ ++ ++ + +
47 Volvox sp. +++ ---- -+3 +- ++ ++?

Insects
Diptera (larvae)
48 Aedes sp. -
49 Chironomus sp.
++-- ++-
++ ++ ++ ++ ++ ++ ++ + ++~ +- ++ +
50 ?+~~~-
++ +- ++ +- -
++
Tanypus sp.
Hemiptera (adults)
51 Limnogonus sp. ---+ -?- ++ +t+ ++ - ++ ++ +
52 Anisops sp.
53 Gerris fossarum --++ +- +?- +
Heteroptera (adults)
54 Nepa sp.
55 Ranatra sp. -.-.? ..+ ++ +- ++ -- __ + --
Odonata (larvae)
56 Anax sp. late instar +
57 Anax sp. early instar ++ ++
--
+ +
58 Agria sp. late instar ++ --+ + +
59 Agria sp. early instar ++-t ++ ++ + ++ ++ + +

Oligochaeta
60 Aulophorus sp. ---++ ++ ++3 ++3 ++?3 ++ ++ + $+ ++- + ++
61 Branchiodrillus sp. ++
+E+ ++ ++ + + ++ +
+
62 Branchiura sp. ---
-- -+
++ ++ ++ ++ +4- ++t ++ ++
4.
+ ++? ++~ +
4- +4 +
4.?
63 Dero sp. ++ ++ ++- ++ ++ +t-- -+3 ++ ++ +
++ 4.
64 ?Limnodrilussp. +-? -+3 ++ +
65 Pistina sp.

Vertebrates
66 Labeo rohita adult ++ ++ +-+ ++ ++ 4.-
+ ++ ++4- + ++t 4-
85 L. rohita fry ++-+ ++ ++
67 Catla catla adult ++S +-- ++ ++ ++~ ++ f+ + ++ ++ +- ++t +-
86 C. catla fry +++ -++ ++
++ +
t) 68 Cirrhinus mrigala adult ++ ++ ++ ++ 4.? ++ ++ - ++t 4-
4 87 C. mrigala fry +++ ++ ++
Table2A. Qualitativeestimateof zooplanktonin the wild pond.
Code Organism 1985 1986 1987
Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr

Copepoda
1 Mesocyclops hyalinus ++ ++ ++ ++ ++ ++ ++ ++ ++ + ++ ++
2 M. leuckarti +- ++ ++ ++ ++ -+ ++ ++ + +-
7 +- -+ ++ + -+
Tropocyclops sp.
8 Nauplii ++ ++ ++ ++ ++ ++ ++ ++ ++ + ++ ++

Rotifera
69 Aneureopsis sp. ++
~+ --

9 Asplanchna priodonta ++
++ --

11 Brachionus angularis ++ +f ++
++t + --
++
++
13 B. budapestinensis -+ -- ++ -- --

14 B. calyciflorus
-++
15 B. caudatus ++ -- ++-
++- ++
16 B. falcatus -- --

17 B. quadridentata ++ -- ++ -$- ++ -- --

18 B. rubens -- ++ + -- --

19 Conochiloides natans -+ ++ -- ++ - + --

20 Dicranophorus forcipatus ++ ~+ -- +-- f+ -- --

22 Filinia longisetta
++
-- --

23 Hexarthra sp. -- --
++ 1--
25 Keratella cochlearis --

26 K. tropica -- - --

27 Lecane (L.) luna ?+ ++ ++ ++ ++ +- ++ -- -- --

28 L. (Monostyla) bulla ?+f ++ ++ +t+ ++ ++ ++ +- -~ +++ +

30 Polyarthra
multiappendiculata --+ ++ +? + +-- -- +
31 Pompholyx sp. ?+ ~~+++?
70 Testudinella sp. -+$ ++ +- ++? + +-- --

Cladocera
71 Alona pulchella -+ ++
32 Ceriodaphnia coruta ++ +- ++ -+ + ++ + ++
33 Daphnia carinata _+ ++ +- -+
35 Moina micrura dubia +- +- +- +-
36 Nauplii ++ ++ ++ ++ ++ ++ ++ ++ ++ ++ + ++ ++ + + + ++ ++

Ostracoda
37 Eucypris sp. --?+ -?-
72 Heterocypris sp. ?-+ ? ---- f+ ++ --

+present;-absent. Data adoptedfrom Deb (1989).

(1992) have suggested, based on their non-aggregated
webs, the value of E to lie roughly between 0.4 and 1.
While the new sets of data constitute a substantial
improvement over Briand's webs, there exists a non-
uniformity of trophic resolution in the data. Many of the
data sets, (e.g., Sugihara et al. 1989, Schoenly et al. 1991)
are isolated subgraphs rather than complete ecosystems
of which they are a part (Pimm et al. 1991). Martinez'
(1992) set of data includes two comprehensive and
evenly resolved webs, while the other webs are aggre-
gated ones catalogued by Briand and Cohen and by
Schoenly. An important lacuna in Havens' (1992) analy-
sis is that his webs have no detritus and, consequently,
hardly any links between the top and basal species, which
is severely unrealistic. When detritus is not included in
the webs, pure detritivores with no predatorsmust appear
as "dangling species" (Ibid.), which Havens has removed
- unrealistically - from his edited webs.
To overcome these shortcomings, it is necessary to
record as many as possible of the existing trophic links
between all trophic levels (including detritus) to approach
a comprehensive description of real food webs. I intend
here to describe in detail the timespecific trophic interac-
tions of two freshwater pond communities, and to study
the relationship between community size and complexity.
I subsequently explore the complexity of a large number
of simulated communities with nonrandom links. Like
the time slices represented in real pond systems, the
model assemblages could either be considered as snap-
shot censuses of the same community fluctuating with
time, or as different independent species assemblages.
Whereas the cumulative webs tend to overestimate link-

248 OIKOS 72:2 (1995)

Table2B. Qualitativeestimateof non-zooplanktonic
organismsin the wild pond.
Code Organism 1985 1986 1987
Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr

Phytoplankton
39 Chlamydomonas sp. ++ ++ ++ + ++ ++ + + +
40 Chlorella sp. ++ ++ ++ ++ ++ ++
++
41 Closterium sp. ++ ++ ++ ++ ++ + + ++
++
42 Diatoma sp. ++ ++ ++ ++ ++ + + +
43 Difflugia sp.
44 Euglena sp. ++ + ++ + + ++
++ ++
73 Gomphonema sp. ++ ++ ++ ++ ++
74 Microcystis sp. ++ ++ ++ ++ ++ + ++
++ ++
75 Navicula sp.
++
++ ++ ++ + ++
76 Oedogonium sp. ++ ++ ++ + ++ ++ + +
47 Volvox sp. --- -+ ++ ++ +- -- -- -- -- --- --- -- +-
Insects
Diptera(larvae)
48 Aedes sp. ?~~~++
++ -- -- ++ + __.. +
Hemiptera(adults)
52 Anisops sp. ?-+ ++ ++ ++ ++ --- --- ++
53 Gerris fossarum ?~~~~~++
++ +-- --

Heteroptera(adultsnaiiads)
54 -++ _
Nepa sp. ?+- ++ ? __- +- + --++ --
55 Ranatra filiformis ?~~~~++
++ ++ -- + + ++ +-

Odonata(larvae)
56 Anax sp. late instar +
57 Anax sp. early instar ++ ++ + + + ++
58 Agria sp. late instar +
59 +
Agria sp. early instar ++ ++ ++ + ++

Mollusks
Bivalvia
77 Margaritifera sp. ++ ++ ++ ++ --- ++ +-- --- ++ ++ + ++ ++ +
Gastropoda
78 Campeloma sp. --+ ++ + ?--- --+ ?--- -- --++ + ++ +-- + + + ++ ++
79 Goniobasis sp. -+ ++ + -? -+ - ?- -++ + ++ +- + + + ++ ++
80 Lymnaea sp. --+ ++ + ?--- --+ ?--- -- --++ + ++ +-- + + + ++ ++

Vertebrates
81 Panchax panchax adult ++ ++ ++ ++ ++ ++ ++ ++ ++ ++ + ++ ++ + + + ++ ++
82 P. panchax larvae ++ +-
83 Rana tigrina early tadpole + +
84 R. tigrina late tadpole +

age densities (Schoenly and Cohen 1991), the separate
treatmentof each of the pond community time-slices and
the randomized analog webs here would obviate that
predilection.
Materials and methods
Study sites and field experiments
Two ponds with similar species composition were se-
lected for study at Sonarpur (22.26?N, 88.25?E) in West
Bengal, India. One of the ponds (650 m2) was managed
for pisciculture, and is referred to here as the culture
pond. The maximum depth of the culture pond (CP)
during the monsoon was 185 + 8 cm, and during the dry
seasons 97 ? 6 cm. The other pond, henceforth called the
unmanaged pond (UP), was about the same shape and
size (ca 646 m2), with the average depth of 170 + 10 cm,
in the monsoon, and 90 + 8 cm in the dry seasons. Both
the ponds were fed by ground water, in addition to being
rain-fed. The ponds are known to have existed for at least
30 yr, and were not subjected to management until 1985
when the CP was preparedfor pisciculture, and this study
was undertaken.
In the pre-stocking phase the CP was treated with the
piscicide mahua oil cake (MOC), preparedfrom the sapo-
nin-containing fruits of mahua Bassia latifolia Roxb. The

OIKOS 72:2 (1995) 249

Table3A. Dietarylinks establishedbetweenpairsof organisms.Numbersindicatethe corresponding
speciescodes fromTables1
and 2.
Predator Prey Selectedreferences

Asplanchna(9) Paramoecium, Euglena (44), Cryptomonas, rotifers, juvenile Maly (1975), Salt et al.
asplanchnids,cladocerannauplii(36), algae (fecultative). (1978), Stembergerand
Gilbert(1985), Wetzel
(1983).
Cladocera,Rotifera, dissolvedandparticulatedetritus.
Phytoplankton, Saunders(1972), Hobbie
Calanoidcopepods(3-7) (1980), Salonenand
and nauplii(8, 36) Hammar(1988), Williamson
(1986)
Daphnia (33, 34) As above, and Keratella (25). Gilbertand McIsaac(1989),
Munshiet al. (1991).
Mesocyclops (1, 2) Zooplankton,nauplii(8, 36), cyprinidfry (85, 86, 87) Sprulesand Bowerman
(1988), Dziuban(1939)
Hartiget al. (1962), Jhingran
(1982, Dash and Hots
(1980), Munshiet al. (1991).
Fish fry and fingerlings detritus.
Zooplankton,phytoplankton,
(85, 86, 87, 82), young
tadpoles(83)
Advancedtadpoles(84) Zooplankton,insect larvae. Dash and Hota (1980),
Munshiet al. (1991).
Adultrohu(66) Phytoplankton,
zooplankton. Jhingran(1982), Das and
Moitra(1963).
Adultcatla (67) Zooplankton,detritus. Jhingran(1982), Munshiet
al. (1991).
Adultmrigal(69) Adult Benthos,detritus,mud.Zooplankton,mosquitolarvae(48). Jhingran(1982). Das and
Panchax (81) Moitra(1963).
Odonatelarvae(56, 57, Zooplankton,fry, ranidtadpole,mosquitolarvae. Roy (1985), Traviset al.
58, 59), Anisops (52), (1985), Munshiet al. (1991),
Gerris (53), Nepa (54), Roy and Munshi(1984),
Ranatra (55) Buskirk(1988).
Chironomus larvae (49) detritus,brachionids.
Phytoplankton, Alfred(1974),
Chattopadhyay and Datta
(1988).
Tanypus(50) Chironomidlarvae,detritus,diatoms,unicellularalgae. Chattopadhyay and Datta
(1988), Chaudhuryet al.
(1983). Munshiet al. (1991).
Mollusks(77-80) Phytoplankton. Munshiet al. (1991).

Table3B. Absenceof dietarylinks betweenpairsof potentialpredatorsand prey organisms.

Predator Prey Selectedreferences

Asplanchnids(9, 10)
Polyarthra(30) and
conochiloids(19)
Carnivorousinsects (imago)
Mesocylops (1, 2)
Fish, tadpoles
Bacteria,protists,fungi, Filinia (22) Hexarthra(23), Bogdanet al. (1980) Gilbert
Polyarthra (30), longspined Brachionus calyciflorus (14), (1985), Gilbert and Kirk (1988).
Keratella (25, 26).
Chlorella(40). Hutchinson(1967)
Daphniacarinata(33), chironomids(49, 50). O'Brienand Vinyard(1978),
Williams(1983).
Keratella (25, 26), asplanchnids. Stemberger (1985), Wetzel
(1983).
Asplanchnids. Haberman(1983).

250 OIKOS 72:2 (1995)

Fig. 1. Relationshipof 600 600
trophiclinkageL (top) and
linkagedensityd (bottom)
with system size S for the
culturepond CP (left) and
the unmanagedpond UP
(right).The powerfunction L 300-
relationshipof L with S is L
= 0.15 S'93 for the CP and L
= 0.09 S2.19for the UP.
Straightlines show
regressionson S (for values
see Table4).
0
10 20 30 40 50 10 20 30 40 50
S S
20- 20-

d i0- 10-

.n.4
_? ...~r~
p.?
-
Li n
I
-
I
-
I r
.
I
.
I i
.
i i 0I I I I I .II I
I I
I
I I

40 20 30 40 50 10 20 30 40 50
S S

known lethal dose of 250 ppm (Homechowdhury et al.

1986) was applied on 7 October, 1985 in order to erad-
icate "weed" fish species Ambassis ranga Hamilton, Ox-
ygaster bacaila Ham, and Panchax panchax Ham. The
pond was later treated with lime on 18 October, 1985 for
alkalization, and with mustard cake for manuring (Jhin-
gran 1982) on 25 October, before the fry (< 20 mm) of
Indian major carps (IMC), namely, Labeo rohita Ham,
Catla catla Ham and Cirrhinus mrigala Ham were
stocked on 30 October, 1985 for rearing. This whole
process of preparing the pond for the culture of IMC in
the CP can be called a "pulse" perturbationexperiment
(Bender et al. 1984), on account of the instantaneous
changes in the composition and densities of the species
that it affected. The fish were fed thereafter,according to
the standard feeding schedule (Sinha 1979), to supple-
ment the nutritional demands for fish growth.
On 27 March, 1987 all the cultured fish were netted out
and a mild dose (100 ppm) of MOC was applied to the CP
on 5 April, 1987 to eliminate the zooplankton. This sec-
ond pulse experiment was conducted to ascertain the new
complexity that the CP community would attain follow-
ing the perturbation(Deb 1989).
Collection of samples
Static representations of biological interactions in food
webs are misleading (Sprules and Bowerman 1988, Polis
1991). Not only do link types change, but also signs and
entry points shift over seasonal cycles (Lane 1985). A
series of time slices is therefore needed to portray the
seasonal dynamics of the community (Winemiller 1990,
Schoenly and Cohen 1991). To divide the pond commu-
nities into several such time slices, all the organisms
except nannoplanktonand microbes were sampled almost
every fortnight at around 08.30 over a period of 24
months in the case of CP (Table 1), and 18 months in the
case of the UP (Table 2).
Plankton were fortnightly collected by filtering a total
of 100 1 of random samples of surface water, using a
plankton net with a mesh of 40/cm. The samples were
preserved in 4% formalin and counted in triplicates on a
Sedgewick-Rafter chamber. Insect larvae collected along
with the plankton in the plankton net were preserved in
4% formalin. Collection of macro-zoobenthos was also
made fortnightly, by dredging the upper layer of the pond
bottom at four random sites with a metal bucket (25 1
capacity), mixing the samples in a large wooden box, and
then sieving the collection by a wire mesh of 10/cm. The
specimens of each collection were fixed in 4% formalin.
Samples of the reared fish, all being the same age, were

OIKOS 72:2 (1995) 251

 1 - 1- Fig. 2. Relationshipof
fractionsof top (TF),
intermediate(IF) an basal
species (BF) with total
species number(S) for the
CP (left) and UP (right)
systems.Straightlines
5- indicateregressionson S.
0 . a
0

0 0,

0
00 v
00

0 I I I I I I I I I

10 20 30 40 50 10 20 30 40 50
S S
1 - 1-

IF .5- .5-
:0

0 I I I I I~ I I I I 0 I I I I I I I I I

10 20 30 40 50 10 20 30 40 50
S S
1 - 4 -

BF .5 - .5 -
* I0 0
0
0 .. ?? ?

0 f I I I I I I I I I 0 I I I 1 I (1 I I I

10 20 30 40 50 10 20 30 40 50
S S

netted out from the CP for ascertaining their size and
maturationalstages. Each of such collections was treated
as a time-slice of the system, and represented a discrete
assemblage of species. Each could then be compared to
other such time-slices of either of the pond systems.
The pond species are described here as nonaggregated,
individual species, and many of the species in which diet
shifts occur have been resolved into relevant life history
stages. Detrital matter is also described as a member of
the food webs. Thus, all the web elements may be de-
scribed here as ontospecies, defined as ontologically in-
dependent and discrete web elements, including the onto-
genic stages of the biological species. The term ontospe-
cies is used here in opposition to Briand and Cohen's
concept of "trophospecies", defined as an aggregate of
functionally or taxonomically related species on the same
trophic level. The numbered entries in Tables 1 and 2 of
the kinds of organisms denote the ontospecies of concern.

252 OIKOS 72:2 (1995)

Table 4. Regressionof predator/preyratio, species and link1985) is unlikely to be revealed by gut analysis (Havens
fractionson total species number.
Variable R2
System Slope S.E. of Constant
slope
CP P/P 0.060 -0.014 0.008 1.35
S E (20, 39) TF 0.208 -1.26 0.379 54.40
IF 0.421 2.15 0.390 -4.54
BF 0.421 -0.89 0.162 50.14
TB 0.035* -0.635 0.512 29.68
TI 0.031 0.266 0.228 11.17
IB 0.016 -0.352 0.417 57.22
II 0.069 0.722 0.408 1.92
UP P/P 0.193 0.015 0.005 0.57
S E (14, 29) TF 0.013 0.218 0.342 23.22
IF 0.111 0.729 0.377 21.33
BF -0.381 -0.947 0.220 55.44
TB 0.094 -0.567 0.322 20.12
TI 0.000 0.070 0.586 19.96
IB 0.004 0.170 0.474 50.24
II 0.017 0.327 0.448 9.67
Analogs P/P 0.116 0.005 0.000 0.749
S E (5, 50) TF 0.197 -0.349 0.015 31.51
IF 0.606 1.166 0.019 15.63
BF 0.576 -0.817 0.015 52.85
TB -0.258 -0.967 0.038 42.57
TI -0.001 -0.025 0.016 16.68
IB 0.028 0.219 0.029 41.14
II 0.476 0.773 0.019 -0.386
S E (5, 10) TF 0.078 2.013 0.402 13.09
IF 0.292 4.466 0.403 -21.59
BF 0.724 -6.479 0.232 108.49
S E (10, 50) TF -0.204 -0.385 0.017 32.76
IF 0.480 0.886 0.020 25.69
BF 0.495 -0.501 0.011 41.54
* Not significantwhena web with 100%TB linkageis removed. would increase the number of dietary linkages in the web.
Estimation of trophic links
Dietary interaction between each pair of ontospecies was
determined from an extensive survey of the literature
(e.g. Jhingran 1982, Wetzel 1983, Williams 1983, Ker-
foot and Sih 1987, Sprules and Bowerman 1988, Munshi
et al. 1991), experiences of the colleagues in the Dept of
Zoology, Calcutta Univ., and personal observations. In
accord with the rationale followed by Martinez (1992)
and Havens (1992), gut content analyses for the actual
trophic links between the species were not performed, on
the basic assumption that links between species reported
in the literature, particularly for the species occurring in
and around Bengal, would hold true for the same species
in the pond communities under study. Gut analyses may
not always reveal the true nature of interaction dynamics
(Stoner and Zimmerman 1988). The predatormay not eat
one prey species in favour of another, depending on their
relative densities, environmental vagaries, or stochastic
physiological reasons at a given moment. The fact that
even the established links may be in a state of flux (Lane
OIKOS 72:2 (1995)
1991). Thus, currently available data on the presence or
absence of dietary links between ontospecies seemed
sufficient for the purpose of this study, which is not to
measure either strength or frequency of trophic interac-
tions, but to describe the communities - the primary
objective of constructing food webs (Polis 1991: 147).
The details of dietary relationships considered here for
linkage estimation are shown in Table 3.
It may be argued that since the species are here as-
sumed to adopt all the feeding biologies they are known
to use, the food links among them would only represent
the upper bound of the linkage possibilities. However,
analyses of any real webs so far published would appear
fictitious in light of this argument, because while the
linkages in real assemblages are likely to appear and
disappear stochastically, food webs are described on the
implicit or explicit assumption of the fixity of trophic
relationships between pairs of ontospecies (e.g., Lane and
Blouin 1985, Sprules and Bowerman 1988, Polis 1991,
Schoenly and Cohen 1991, Havens 1991, 1992). The
possibility of estimating the upper bound of linkages
(unpubl.) has been precluded in this study by (a) counting
zero linkage between pairs of species unless interactions
are reported in the literature, and (b) incorporating the
data (Table 3B). The follow-
predator-avoidance/escape
ing trophic interactions were further considered for
achieving a high-resolution analyis of the webs.
Passive predation. The herbivore cladoceran Daphnia
spp. (nos 33 and 34 in Tables 1 and 2) are known to filter
algae and in the process consume small rotifers such as
Keratella spp., apparentlyregardless of the rotifer's den-
sity (Gilbert and MacIsaac 1989). Such passive predation
Facultative herbivory. The predatory rotifer Asplanchna
spp. preferentially feeds on metazoan prey and usually
avoids phytoplankton and protozoa in the presence of
rotifer prey (Table 3B). Nevertheless, asplanchnids can
thrive on a diet of unicellular algae (Salt et al. 1978) and
protists like Euglena (no. 44) and Paramoecium (Maly
1975), and therefore these linkages have been included in
this study. Laboratorycultures of A. priodonta (no. 9) are
known to sustain a high growth rate on Cryptomonas
(Stemberger and Gilbert 1985). It is not known which
other phytoplankters it may facultatively feed on under
natural conditions.
Omnivory. Trophic level omnivores, defined as consum-
ers feeding on more than one trophic level, are rare in
catalogued webs (Pimm 1982, Cohen et al. 1990), but are
a common and persistent feature in aquatic communities
(Menge and Sutherland 1987, Sprules and Bowerman
1988).
The catla (no. 67) and ranid tadpoles (nos 83 and 84)
are known to be generalised feeders (Dash and Hota
1980, Jhingran 1982, Munshi et al. 1991). Mesocyclops
253
 1- 6
I Fig. 3. Relationshipof the
fractionsof top-basal(TB),
top-intermediate (TI),
intermediate-basal (IB) and
intermediate-intermediate
(II) links with S for the CP
IB .5' .5 (left) and the UP webs
(right).The circledpoint has
been excludedfrom
..... . .~,II' t" calculationof mean.Long
lines are drawnthroughthe
0 1 I fI 1 1 1I meansof the fractions.Short
lines indicatethe mean
10 20 30 4050 10 2030 40 50 values given by Briandand
Cohen(see Table5).
1- 1-

TI .5' .5 -
*
. ?
*111;1 I-

i I I I- I I l 1- i'1I I 1 1 'I I I
10 28 30 4850 10 2038 40 50
1-

* ..?I Ji e

IB .5' ."..
i.
I?
,
.5
e

a 1 II 1 II I -I- iI l I
10 20 30 4050 10 203 40 50

I
. t~f

II .5 .5 ,

1 IlI 11
1 1
10 20 38 4050 18 2830 40 50
S S

(nos 1 and 2) prey on herbivorous rotifers, the predatory
asplanchnid (nos 9 and 10) rotifers, and crustacean nau-
plii (nos 8 and 36), including their own (Sprules and
Bowerman 1988). Freshwatercalanoids, even some diap-
tomids previously thought to be purely herbivorous might
prey on small, weak-swimming small rotifers (William-
son and Butler 1986). Since it is not known if the partic-
ular diaptomid species from the ponds in this study prey
on any rotifers, links between them are not included here.
Diet switching and life cycle omnivory. An organism
feeding on a specific level of the food chain at one stage

254 OIKOS 72:2 (1995)

Fig. 4. Relationshipof
predator-prey ratiowith S
4 4
for the CP (left) and UP
webs (right).The mean 3 3
values are indicatedby long P/P
lines, comparedwith the
meansgiven by Briandand 2 2
Cohen (shortline).
t~
I
1
1 I 1i I I I 1 1 'i- 1 I 11 i1
18 28 38 48 58 10 203 48 50

S S

of its life may shift to another diet level at another stage
(Pimm and Rice 1987). For example, the catla fry (no.
86) feeds on algae as well as primary consumers, i.e.
rotifers and cladocerans, and occupies the secondary con-
sumer level. But as the fish grow larger, they occupy the
tertiary or top consumer level, and also eat detritus. The
fry of Cirrhinus mrigala (no. 87) prey on zooplankton
whereas the adults (no. 68) thrive on zoobenthos, detritus
and mud (Jhingran 1982). The predatory Mesocyclops
spp. feed on unicellular phyto-plankton at earlier instars
(no. 8) (Williamson 1986, Havens 1991, pers. obs.).
Despite the food web theorists' description as "biolog-
ically impossible" (e.g. Pimm 1982: 70-71), predation
loops are common in zooplankton (Williams 1980,
Sprules and Bowerman 1988, Havens 1991) and insect
communities (Southwood 1985), and are caused by de-
velopmental changes in the sizes of coexisting predators
(Warren and Lawton 1987), as well as by intra-guild
cannibalism (Sprules and Bowerman 1988). Reciprocal
predation involving species of different taxonomical
groups are also common. Mesocyclops spp. for example,
100 - *,-*
*
0'*?
?. * .,,
0.*
.
. -*4.*At
.**7'
.-
0
.qe, :44
- c: ,q...
%60
a IeI t chironomids. The detrital subweb is therefore not com-
20-
.
0 iI
10 20 30
S The alkalization and nutrient-enrichmentof water estab-
Fig. 5. Relationshipbetween the frequencyof systems with
differentchain lengths(ChL)and S for ranomizedanalogsof
pond webs; 250 iterationsfor each S are shown.
attack and kill cyprinid fry and fingerlings smaller than
20 mm (nos 85, 86 and 87) (Dziuban 1939, Hartig et al.
1962). However, they fall prey to the fish when the fish
grow larger. Here, the growth stage of the fish alone
affects the directionality of the trophic link. Similar onto-
genic predation reversal is observed between the carnivo-
rous insects (nos 51, 52, 53, 54, 55, 56, 57, 58 and 59) on
the one hand, and cyprinid fry (Jhingran 1982, Roy
1985), and anurantadpoles (no. 83) (Caldwell et al. 1980,
Travis et al. 1985, pers. obs.), on the other.
Detritivory. Although not included in most food webs,
detritivory is a major pathway of energy transfer in aq-
uatic systems. Rotifers and crustaceans have been re-
ported to consume particulate detritus (Saunders 1972,
Hobbie 1980). Dissolved organic matter (DOM) is not
only consumed by different species of algae by hetero-
trophy (Neilson and Lewin 1974), but also by zooplank-
ters (Salonen and Hammar 1986). Adult cyprinid fish are
known to consume particulateorganic matter (POM) and
mud (Jhingran 1982), which are also consumed by the
purely detritivorous oligochetes. Thus, POM, DOM and
mud are all included here as elements of the basal stratum
"
".t .*:::;t of the pond webs.
The consumers of live and dead matter appeared to
_ form what Pimm (1982) called "ill-defined compartments
linked by common predators".The overlap between the
detrital and live food chains exists not only between
predators, but also in the lower strata of the benthic and
pelagic food chains, especially between zooplankton and
partmentalised.
40 50
Field observations
lished a high water quality in the CP; its eutrophic nature
was indicated by the relative dominance of rotifers and
small cladocerans in the surface water (Terek 1983), the

OIKOS 72:2 (1995) 255

608 28
d
18
0
18 20 3840 50
S S
presence of Chironomus, Tanypus and Limnodrillus
among the benthic fauna (Bernstein 1981, Terek 1983),
and year-round abundance of diatoms and euglenophytes
(Hutchinson 1967). The total species numbers in the pond
varied from 20 to 39.
Long-spined morphs of Brachionus calyciflorus Pallas,
which are known to be invulnerable to asplanchnid preda-
tion (Wetzel 1983), were recorded in the presence of
Asplanchna priodonta Gosse in the CP (Deb and Baner-
jee 1989), which indicated the plausible absence of link-
age between the two species. Long-spined morphs of the
Brachionus were not found after the disappearanceof the
predator in March-April 1986 (Table 1A).
Despite similar zooplanktonic species composition, at
least at the beginning of the study (Table 2A), the UP
community structurewas different from the CP, with the
total species number varying between 14 and 29. The
only benthic animals collected from the UP were the
littoral mollusks (Table 2B). The fish Panchax panchax
existed in both ponds, but was eradicated from the CP
prior to pisciculture. Pelagic insects were comparatively
more frequently found in the UP (Table 2B).
Analysis of pond communities
All the time-slices consist of large webs with S > 10. The
variances of all the web parametersin both the ponds are
large. Homogeneity chi-square tests reveal that the CP
and UP systems are different at the 0.1 level of signif-
icance in link and species proportions. However, since it
is uncertain whether the webs from either of the pond
systems constituted a truly random collection, use of
standard statistical tests of difference between them is
perhaps inconclusive. The extent of scale-invariance is
therefore measured here principally by the regressional
estimates of the parameters on S.
When the number of links and the linkage density
(d=L/S) are plotted against S for each time slice (Fig. 1),
the regression is found to be positive and significantly
different from zero (P <0.001). The log-log regression of
L against S (Havens 1992, Martinez 1992, 1993,) gives
256
Fig. 6. Relationshipof L
(left) and d (right)with S
for the simulatedwebs. L =
0.12 S211. Straight line
shows regressionon S.
'"!iii
1 i
llll1
10 2038 40 50
an estimation of the exponent (for eq. 1), E = 0.93 for the
CP and 1.19 for the UP. When the intercept log(k) is
constrained to be zero, the exponent becomes 0.48 for the
CP and 0.42 for the UP webs.
The fraction of top species (TF), calculated as the
number of top species divided by the total number of
ontospecies (S) tends to decline with S for CP webs,
while the relationship is weakly positive for the UP (Ta-
ble 4 and Fig. 2). In both the ponds, the fraction of the
intermediate species (IF) is directly related, and the basal
species fraction (BF) is inversely related with S (Table 3),
each with a large coefficient of variation.
The correlation of the top-basal (TB) links with S is
negative, and that of the intermediate-intermediate (II)
links is positive, for both the CP and UP systems. The
link fractions show no specific trend as S increases (Table
4). The relationships are weak, both for Pearson's r and
Spearman's rank correlation estimates (P > 0.1), although
the latter indicates slight (two-tailed test, P = 0.08) scale-
dependence of the II linkage for the CP. The mean values
of the link fractions (Fig. 3) seem to support the link
scaling law. The variances of these link fractions are
0.020, 0.004, 0.013 and 0.014 for CP webs and 0.004,
0.012, 0.008 and 0.007 for UP webs, respectively. The
pond systems in respect of the mean TB and TI are
statistically no different, while the difference is signif-
icant (P <0.05) for the mean IB and II fractions.
The chain length (ChL) does not exceed 5 for webs in
either of the ponds. The mean ChL is 3.84 for the CP and
2.59 for the UP webs. The higher value of the mean in the
CP seems to correspond with larger species numbers in
CP webs than the UP.
The mean of the ratio of the total predator taxa to the
total prey taxa (PIP) for the ponds is roughly constant, at
0.95 for the CP, and 0.92 for the UP webs. The values are
not statistically different, though the difference between
variances of the pond systems is significant (F=3.15,
P<0.05).
The different ranges of system size and the different
degrees of scatter in the data from the pond systems seem
to indicate they might comprise subsets of a larger range
of webs composed of a pool of similar species associ-
OIKOS 72:2 (1995)
 i
4
3
I?
,?r
: I
TF ( I I I rl
P/P
0.5 I
t t
(
) ) ( 1( ( ( I t

( ( )(
I I ()
((I(
2 I
.
!
..
r I ) It I I 'I
I I ) ( I I, 1(I ,(
III,Ii,
(
r ,( I , ! I IlIllII I lIIIllI :IIIII,,.
(
(I
It (((
(I ((
I)
((
))) ()) ()( II
)I
r. Il,
,, m I II I n
=t
- .rL[.
.
,
I1 1I! .l_.1 I I 1,llI,HLlI
1 I1 l HH 11 qrl,,]l 11
n

I(( I (I(,( )(( 1(1

II, ," I' I,,1 III I
-- I -
I I I I I II I I
-----

'
--

I
I f

I I
F I

I II
I 1

II
I I
0
t1 28 30 40 50
i0 20 30 40 50 S
Fig. 8. Relationshipof predator-prey
ratiowith S for the rando-
mized analogwebs. Lines as in Fig. 4.
i
ations. Generalisations are likely to be valid if a larger
data base with a range of system size encompassing those
of the pond webs is available.
IF
0.5'
Expanding the data base
With the objective of examining the quantitative web
properties for a large number of systems with species as
T I I T - l I- _I
well as non-random linkage rules identical to those ob-
0 I I I I I I I I I I served in the ponds, a set of 11 500 randomized commu-
nities, analogous to the pond systems, was generated on a
B1 20 30 40 50 computer. Since the pond webs contain ontospecies com-
monly found in most tropical freshwater systems, random
assemblages of those species pooled from the two ponds
would represent a large proportion of feasible species
I
associations likely to be found in tropical waters. These
analog webs would therefore provide a considerably
wider database.
.6S , W While creating the model communities, the following
o
BF
.,
rules, in addition to the specified dietary interactions
discussed above, were obeyed (summarized in Appen-
,,,', : ". ,,
dix).
.,,",:,,
'"* . ,-. ' !( i I 1) The species composition of each community was
,'"";"
;" "''1 , i[,I-
I:'~i
-,,9,9 ,,,,,9
, 99
,, , ,:,,,,,,,,!tl
,,1., 11 randomly chosen from among the 87 kinds of organisms
(Tables 1 and 2) altogether encountered in the ponds. In
addition to the organisms, DOM, POM and mud were
__ .
I I I II I I I _-- -

I I ubiquitously present in all the random assemblages to

0 simulate the real pond systems.
i0
Fig. 7. Relationshipof TF,IF andBF withS fortheanalogwebs.
Straightlines showthe regressions.The curveson the ceilingof
the scattersindicatethe upperlimits, given in eqn. (2). 5 000
iterationsshown.
2) In order to eliminate the possibility of generating
2L 30 40 50 systems with zero connectance (see also condition 3), the
size for each system was chosen randomly such that S >4,
including the three detrital elements. The upper limit of
S the system size was set at S = 50, to conform to the pond
webs. For each S, 250 random assemblages (iterations)
were created.
3) Each community comprised at least one algal spe-
cies, and at least one primary consumer species.

17 OIKOS 72:2 (1995) 257

Table5. Propertiesof food webs.
Source Cohenand Sugiharaet al. Martinez Havens CP UP Computer
Briandtl) (1989), (1991, 1992) (1992) analogs
Schoenlyet
al. (1991)
No. of webs studied 113# 40 10 50 44 32 11500

Meanchain length 2.65, 2.71 2.89 NA NA 3.84 2.59 3.64#, 3.98

Maximumlength 5.19 7 NA NA 5 5 5
Minimumlength 2 1 NA NA 1 1 1
Totalpredator/total
prey taxa 1.11 1.56 NA NA 0.95 0.92 0.90
TF 0.28 0.46 0.01 0.06 0.184 0.280 0.22
IF 0.53 0.38 0.86 0.44 0.571 0.357 0.477
BF 0.19 0.16 0.13 0.50 0.246 0.344 0.303
TB (%) 8.0 NA NA 5.0 9.26 7.53 15.43#,9.86
TI (%) 36.0 NA NA 10.0 17.26 21.52 15.78#, 16.47
IB (%) 27.0 NA NA 53.0 50.93 54.02 48.09#, 50.66
1 (%) 29.0 NA NA 32.0 22.55 16.92 20.70#, 23.01
1E2) 0, 0.1(3 0.424 0.78#, 1.18 0.4 0.93 1.19 1.11#
k?2) 1 0.258(4) 0.16 1.0, 0.16(5 0.148 0.09 0.118
1 Fromvariouspublications,chiefly, (a) BriandandCohen 1984;(b) CohenandBriand1984;(c) Cohenet al. 1986;(d) Briand
and Cohen 1987. The two values of meanchain lengthare from (d) and (c) respectively.
2 For the relationship L = k.St+i.
3 Briand1983.
4 Valuenot reported.Calculationmine.
5 Martinez(1993).
NA Valuesnot available.
# For webs includingS < 10. All othervalues obtainedfrom webs with 10 or more species.

Program listing 1.3 < BF S2/3 <3.7 (2b)

The program for generating the randomized analogs of
the pond systems and for ascertaining their linkage prop-
erties were written in BASIC. It took about 14 min for
each 1000 iterations to run on an IBM compatible 80386
personal computer. Program listing is available free on
request.
Analysis of the analog webs
The mean ChL of the 11 500 random webs is 3.64, with
the specified upper limit of 5 in the paradigm of the pond
communities. The ChL tends to increase monotonically
with S (Fig. 5), so that webs with S >5 tend to have an
increasing frequency of ChL >2, and the largest webs (S
>45) all have ChL >4.
Both L and d increase with S. The range of k (eq. 1) for
the entire data set lies between 0.04 and 0.14 (Fig. 6), and
the mean value of E (1.10) matches with the value ob-
tained by Martinez (1992, 1993). When the intercept is
constrained to be zero (log(k) =0), the exponent E esti-
mates at 0.56, but provides a poor fit to the data.
The species fractions are strongly related with S -
positively for the IF, and negatively for the TF and BF
(Table 3). The scatter of the points shows an obviously
curvilinear relationship of each of the fractions with S
(Fig. 7). The relationships may be described by the func-
tions
2<TF. S<15, (2a)
and
4 < (0.9- IF)-S <19 (2c)
The means of the fractions of TB, TI, IB and II links are
0.154, 0.158, 0.481 and 0.207, respectively. The var-
iances are large, and the regressions of the fractions differ
only slightly from zero (Table 4). When systems with
only two trophic levels (100% TB links) are all removed,
the mean of TB fractions reduces to 0.095. This reduction
is compensated by about an 8% increase in the mean
values of all the other fractions.
The mean P/P ratio is constant (0.90), conforming to
that for the CP and UP webs. The ratio seems to be the
most scale-invariant of all the properties. The scatter of
the points are roughly within the confidence limits of the
mean (Fig. 8).
Discussion and conclusions
Problems of defining trophic levels and true "top preda-
tors" (Paine 1988, Lawton 1989, Polis 1991) are obviated
in this study through time-specific descriptions of the
communities, composed of trophically related ontospe-
cies (cf. Havens 1992). These time slices, and their ran-
domized analogs, also expand the data base. The large
number of simulations are likely to ensure the robustness
of conclusions.

258 OIKOS72:2 (1995)

 Table5 summarizesthe findingsof this studyvis-a-vis
the results obtainedby previous workers.Results ob-
tainedby Polis (1991), thoughsupportiveof my results,
arenot includedin the table,becausehis datapertainto a
non-aquaticcommunity.
The resultof chain-lengthanalysis(Fig. 5) is compara-
ble with the mean ChL of 3.19 (Jeffries and Lawton
1985) and 3.77 (Warren1989) for freshwaterwebs, and
conforms to Winemiller'sand Martinez'findings and
simultaneouslyunderscoresthe incompletenessof Cohen
and Briand's as well as Havens' data. The larger the
system,the more likely is it to be longer,but the maxi-
mum limit of chain length stipulatedhere is reached
ratherquickly.It is uncertainwhetherthe constraintof
the maximumChL of 5 imposed on the randomized
model systems has any significanteffect on the trend
observed here. However, an interestingsimilitude is
found in Drake's (1990) study where web assemblies
with a maximumChL= 5 tendedto attainsimilarmean
ChL afterstabilisation.
The value of the exponentE for the pond webs and
theiranalogsis comparablewiththosesuggestedby Mar-
tinez (1992, 1993). The values indicate that the links
increase approximatelyas the squareof S. The value
rangeof k= averageL/S2 + 1 SD for thepond-andanalog
webs corroboratesMartinez'(1992, 1993) analysis.The
valueof k= 0.1 for the UP webs, thoughcorrectlymatch-
ing Martinez'estimates,is slightly less than the range,
which presumablypurportsthatthe samplesize for this
pond is insufficient.The over-allconformityto the con-
stantconnectancehypothesis(Martinez1992) shows that
it is a propertyof naturalwebs,particularlywherea large
proportionof thecomponentspeciesaregeneralisedfeed-
ers, and/orare life historyomnivoresdue to ontogenic
diet shifts.Thepresenceof filterfeederswill also contrib-
ute to an increasein thelinkagedensity,for theyconsume
all organismsof suitablesize. If one considersthe intake
of largenumbersof bacteriaby largerorganisms(evenby
algae),the linkagedensitywill necessarilybe higher,and
corroboratethe hypothesiseven more strongly.
Havens'data(Havens1992, Martinez1993), together
with this study,seem to reveala generalisationregarding
the relativemagnitudesof the link fractions:IB > II > TI
> TB. This patternpersistsfor both smaller(S <10) and
largersystems. The scale-invarianceof link fractionsis
moreprominenthere thanin Havens(1992): The II and
IB fractionsin Havens'datahavestrongcorrelationswith
S (P< 0.001), whereasthose in the pondwebs hereindo
not (Table4). Rankcorrelationanalysis(Martinez1993)
also seems to supportthe link scaling law for the pond
webs. The variancesof the link fractionsfor both the
pondsas well as theiranalogsareno widerthanthosefor
Havens' webs. However, the scale-invarianceis lost
when large samplesof randomizeddata are considered,
which again supportsMartinez'(1993) results.
The scale-dependenceof the species fractionsis obvi-
17* OIKOS 72:2 (1995)
ous. Althoughthe small sample size of the UP system
showsa (statisticallyinsignificant)rise of TF with S (Fig.
2), the overallpatternof scatterfor the fractions(Fig. 7)
agreeswith Martinez'(1993) analysisof Havens'data.
Schoener(1989) surmisedthattheP/P ratioshouldrise
with S. Thatrise is prominentonly for the smallestwebs
herein(Fig. 8). Themeanof theP/P fractionsis relatively
invariant,and could be derivedfrom the mean species
fractions(Cohenand Briand1984):
(TF + IF)/(IF + BF) = (1- BF)/(1-TF) (3)
However,thecalculationof the meanP/Pratiofromeq. 3
is indeedtautologous(despiteCohenandBriand'sclaim
to the contrary),since the predatortaxaandpreytaxaare
definedin termsof the species fractionsthemselves,and
thereforethe derivationis perhapsfortuitous.The P/P
ratio may also be derivedfrom the mean values of the
fractions,
(TF Sn+ IF S")/(IF Sn+ BF . S), (4)
n beinganyrealnumber.Thenon-linearrelationbetween
the species fractionsand the P/P ratiomay indicatethat
the calculationof the second(or even third)derivativeof
TF, IF and BF from the solutions of the functionsas
depicted in Fig. 7 and eq. 2 could explain both the
scale-invariance of P/P andthe proportionality
of species
fractionswith S.
Full descriptionsof ecological interactionsof all the
speciescomprisinga systemin a real-lifesituationwill be
completelyunwieldy,andvirtuallyimpossiblein the cur-
rentstateof knowledge(Schaffer1981, Slobodkin1987,
Schoener1989). The presentstudy is no exceptionbut
includesas muchdataas areavailableandmoredatathan
have previouslybeen used in studiesof food web com-
plexity.Additionof micro-organisms, andfurtherdetails
of dietaryinteractions,suchas thephagotrophic uptakeof
bacteriaby phytoflagellates(Tranviket al. 1989), would
have enhancedthe complexityof the webs. However,it
seemsunlikelythataddeddetailswouldseriouslychange
the conclusionspresentedhere.
The close proximityof the findings of this study to
other studies of similardetail (Havens 1992, Martinez
1992, 1993) indicatesthatconstructionof computerized
analogs of real webs, incorporatingall known trophic
biology of the componentspecies, may prove to be a
useful, if shortcut,methodof generatingsufficientlyre-
liable databasefor analyses.
- A partof this work drawson my Ph. D.
Acknowledgements
dissertation
submittedto Calcutta
Univ.I amextremely
grateful
to T.Burton,
M.GadgilandN. V.Joshiforhelpfuladvice,and
G. Polisforkindlysupplyingme withhis unpublished
manu-
script.
259

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261
 Appendix
Flowchart showing
C'START the algorithm for
I~~~~~~~~generating
randomized webs on
GENE RATE computer using
RANDOM# ontospecies from the
2 < S4 30 real ponds and non-
random linkage rules
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262
UIKOUS/2:2 (1995)