e c o l o g i c a l m o d e l l i n g 2 0 8 ( 2 0 0 7 ) 286294

available at www.sciencedirect.com

journal homepage: www.elsevier.com/locate/ecolmodel

The relevance of ecological pyramids

in community assemblages

Brian D. Fath a,b, , Megan C. Killian c

a Biology Department, Towson University, Towson, MD 21252, USA
b Dynamic Systems Program, IIASA, Laxenburg A-2236, Austria
c Environmental Science Department, Towson University, Towson, MD 21252, USA

a r t i c l e i n f o a b s t r a c t

Article history: In this paper, we investigate the trophic pyramid structure of 17 commonly studied ecosys-
Accepted 4 June 2007 tems (terrestrial and varying aquatic environments). Food-web taxa from these ecosystems
Published on line 10 July 2007 are classied into six functional groups: primary producers, herbivores, carnivores, omni-
vores, detritus, and detrital feeder; we nd that the majority of the food webs exhibit an
Keywords: inverted pyramid structure with more higher trophic groups than lower ones. We nd the
Community assembly rules average pyramid and inverted pyramid trophic distributions for these webs, and along with
Ecological network analysis the uniform distribution, construct large-scale ecological networks based on these distribu-
Food webs tions. By creating and analyzing structural and ow matrices of these distributions using the
Trophic pyramids cyber-ecosystem community assembly rule model, we compare the ecological parameters
cycling, amplication, homogenization, indirect effects, and synergism. Excluding ampli-
cation, which does not occur in larger-scale models, we nd that all network parameters
show strong characteristic behaviors regardless of the trophic structure. Therefore, ecolog-
ical network parameters are not considerably sensitive to different structures, indicating
there may not be a preferred diet distribution.
2007 Elsevier B.V. All rights reserved.

1. Introduction is transferred from prey to predator, and as this transfer

The creation and analysis of food-web models has been
benecial to the development of structural and functional
relationships within both terrestrial and aquatic ecosys-
tems. Elton (1927) introduced the idea of food pyramids as
a result of observing characteristics of population dynam-
ics within multiple ecosystems. Elton noted a pattern, which
he described as the pyramid of numbers, exhibited within
any given area: a large number of producers supporting
a smaller number of herbivores, who then in turn sup-
port a smaller number of predators, which support an
even smaller number of predators of the predators. Energy
occurs, energy is lost. The process of energy transfer and
the resultant decrease in energy available to the predator
provides one explanation of population dynamics within
an environmentthere is an abundance of producers (who
receive the most energy) compared to a small amount of top
predators (who receive the least amount of energy) (Fig. 1a,
Elton, 1927).
To analyze standing crop amount of energy within the
environment at one time Elton took the pyramid of numbers
a step further creating the idea of the pyramid of biomass.
Because of varying biomass of predator and prey, the number
of consumers at each feeding level is multiplied by weight,

Corresponding author at: Biology Department, Towson University, Towson, MD 21252, USA. Tel.: +1 410 704 2535; fax: +1 410 704 2405.
E-mail address: bfath@towson.edu (B.D. Fath).
0304-3800/$ see front matter 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.ecolmodel.2007.06.001
 e c o l o g i c a l m o d e l l i n g 2 0 8 ( 2 0 0 7 ) 286294 287

Table 1 Generalized connection matrix for ows from one functional group to another (from columns to rows)
Trophic structure Primary producer Herbivore Carnivore Omnivore Detritivore Detritus

Primary producer 0 0 0 0 0 0
Herbivore 1 0 0 0 0 0
Carnivore 0 1 1 1 1 0
Omnivore 1 1 1 1 1 0
Detritivore 0 0 0 0 0 1
Detritus 1 1 1 1 1 1

The number of compartments in each functional group can be expanded while this meta-structure constrains the connection possibilities. In
this manner networks of any size and functional group conguration can be constructed.

observer (e.g., Cohen et al., 1993; Hall et al., 2000). There

Fig. 1 (a) A simple trophic pyramid of numbers or biomass
and (b) the inverted pyramid in which a smaller number or
biomass of primary producers supports a larger amount of
herbivores and carnivores.
accounting for a more accurate representation of total energy
present within the ecosystem. In terrestrial and shallow water
ecosystems, the greatest total biomass is from the producers
(i.e., trees in a forest) because they are large organisms, with
high accumulation of organic matter, long life cycles, and low
harvesting rates. However, in marine ecosystems, phytoplank-
ton are the producers of the system having a short life cycle
with rapid reproduction, little accumulation of organic matter,
and are heavily consumed by herbivorous zooplankton and
larger consumersresulting in an inverted pyramid (Elton,
1927) (Fig. 1b).
A weakness of both the pyramid of numbers and biomass
is the misrepresentation of energy ow within the ecosys-
tem. By visually analyzing the models, one may assume
the ow of energy terminates at the top predator. However,
this is only the case when detritus, decomposers, and het-
erotrophic microbes are excluded from the system by the
is a cyclic ow of energy within all ecosystems: when an
organism dies, its remains decayreturning energy back
into the system and essentially recycling biomass back into
the food-web (see for example, Pomeroy, 1974; Patten, 1985;
Fath and Halnes, 2007). Differing sampling intensities and
techniques by investigators are critical to food-web analy-
sis, affecting the true representation of species richness and
trophic interactions (Martinez et al., 1999a,b). This is why it
is imperative to sample and record every part of the ecosys-
tem in question, and if it is essential to exclude some species
from the study, record what was not included (Cohen et al.,
1993).
2. Community assembly rules
Due to the difculty in acquiring the requisite data for fully
developed empirical food webs, researchers have developed
community assembly rules that are heuristics to construct
ecological food webs. Common rules that have been developed
are random or constant connectance (Martinez, 1992), cascade
(Cohen and Newman, 1985), niche (Williams and Martinez,
2000), and cyber-ecosystem (Fath, 2004), each with its own
assumptions and limitations. Here we follow-up on the inves-
tigations of the cyber-ecosystem model (Fath, 2004) which uses
a meta-structure of six functional groups: producer (P), herbi-
vore (H), carnivore (C), omnivore (O), detritus (D), and detrital
feeders (F), within which random connections link species

Table 2 Five network parameters, where G is the non-dimensional ow intensity matrix (gij = fij /Tj ), N the integral ow
intensity matrix accounting for direct and indirect pathways (N = (I G)1 ), and zj is the input vector
Property Denition Test
n n
(n /n 1)zj
j=1 ij ii
Cycling Percentage of ow derived from cycling 
i=1
n n
n z
i=1 j=1 ij j
Amplication Integral ow along a pathway exceeds direct input nij > 1 for i = j
CV(G)
Homogenization Action of the network makes ow distribution more uniform. CV CV(N)
>1
is the coefcient of variation
n n
(nij iij gij )
Indirect-to-direct ratio Indirect ow contribution exceeds direct ow in the network n j=1
i=1
n >1
g
 i=1 j=1 ij
+utility
Synergism Positive utility exceeds negative utility giving rise to dominant  >1
|utility|
positive relations
288 e c o l o g i c a l m o d e l l i n g 2 0 8 ( 2 0 0 7 ) 286294

Table 3 Seventeen empirical food webs from Dunne et al. (2002) representing aquatic and terrestrial ecosystems
ranging in size from 29 to 220 taxa
Ecosystem Habitat Taxa Reference

Pyramid
Canton Creek Stream 109 Townsend et al. (1998)
Stony Creek Stream 113 Townsend et al. (1998)
Bridge Brook Lake Lake/pond 220 Havens (1992)

Inverted
Benguela Marine 29 Yodzis (1998)
Caribbean Reef small Marine 50 Opitz (1996)
NE US Shelf Marine 81 Link (2002)
Chesapeake Bay Estuary 32 Baird and Ulanowicz (1989)
St. Marks Estuary Estuary 48 Christian and Luczkovich (1999)
Ythan Estuary, 1991 Estuary 90 Hall and Raffaelli (1991)
Ythan Estuary, 1996 Estuary 134 Huxham et al. (1996)
Little Rock Lake Lake/pond 181 Martinez (1992)
Skipwith Pond Lake/pond 35 Warren (1989)
Scotch Broom Terrestrial 154 Memmott et al. (2000)
Coachella Valley Terrestrial 30 Polis (1991)
El Verde Rainforest Terrestrial 154 Waide and Reagan (1996)
Grassland Terrestrial 75 Martinez et al. (1999a,b)
St. Martin Island Terrestrial 44 Goldwasser and Roughgarden (1993)

Table 4 Categorization of taxa into food groups

Ecosystem Producer Herbivore Carnivore Omnivore Detritivore Detritus Compartments Taxa

Pyramid
Canton Creek 55 32 19 6 29 1 142 109
Stony Creek 62 34 17 10 35 1 159 113
Bridge Brook 112 69 29 10 1 1 222 220

Inverted
Benguela 1 3 22 3 2 0 31 29
Caribbean Reef 2 13 29 6 11 2 63 50
NE US Shelf 1 7 50 18 17 1 94 81
Chesapeake Bay 2 5 13 7 11 0 38 32
St. Marks Estuary 5 12 24 7 12 1 61 48
Ythan Estuary, 1991 2 11 47 27 31 0 118 90
Ythan Estuary, 1996 3 11 89 27 31 1 162 134
Little Rock Lake 59 92 64 7 47 1 270 181
Skipwith Pond 0 24 29 2 10 1 66 35
Scotch Broom 4 84 126 5 3 0 222 154
Coachella Valley 1 6 18 4 3 2 34 30
El Verde Rainforest 13 56 79 12 23 4 187 154
Grassland 8 15 52 0 0 0 75 75
St. Martin Island 5 15 20 6 9 3 58 44

Categorized taxa from 17 ecosystems into six categorical food groups: producer, herbivore, carnivore, omnivore, detritivore, and detritus with
habitat information. Taxa numbers are taken from the original data; compartment numbers are the total amount of food categories represented
within the ecosystem. Compartment numbers are greater for the majority of ecosystems due to taxa that are present in multiple food groups.
Pyramid ecosystems exhibit the traditional pattern of more producers than any other food group. Inverted ecosystems exhibit more carnivores
than any other food group.

based on these denitional constraints (Table 1). The original
intent of the cyber-ecosystem model was to create large eco-
logical networks up to 600 compartments to distinguish
the impact of size on a suite of commonly used ecologi-
cal network parameters (Table 2).1 From that analysis, some
interesting results followed; namely, with increasing system
size, network amplication failed to appear and synergism,
although occurring, became weaker. For simplicity, in those

of network parameters such as indirect effects ratio, homog-

1
Ecological network analysis considers the role of each entity
embedded in a larger system and has been a fruitful way of investi-
gating system level properties of ecosystems. In particular, a series
enization, synergism, and mutualism have evolved around this
analysis. See Patten, in preparation, Fath and Patten (1999), and
Fath (2004) for further details regarding these properties and the
history of Ecological network analysis.
 e c o l o g i c a l m o d e l l i n g 2 0 8 ( 2 0 0 7 ) 286294 289

simulations a uniform distribution of all functional groups

was used. In other words, the 60-compartment network had
10 each of the six functional groups [P, H, C, O, D, F]; a 120-
compartment network, 20 each; and the 600-compartment
network, 100 each. It was suggested that other distributions
should be considered to see if there is any impact on the
results.
3. Methods
The best choice to determine the trophic distribution for use
in the cyber-ecosystem model would be based on real ecologi-
cal data. The set of 17 ecological networks (Dunne et al., 2002,
2004), ranging from n = 29 to 220, provided a good source of data

Fig. 2 Network parameters for 120-compartment models with varying trophic distributions. Each data point represents the
average of 50 different simulations for a given structure except (b) where each data point represents the total number of
amplication occurrences per 50 simulations of a given structure: (a) cycling index, (b) amplication occurrence, (c)
homogenization index, (d) indirect-to-direct ratio, and (e) synergism.
290 e c o l o g i c a l m o d e l l i n g 2 0 8 ( 2 0 0 7 ) 286294

Table 5 Distribution of each trophic structure normalized to 60, 120, and 300 compartments, respectively
Trophic structure Producer Herbivore Carnivore Omnivore Detritivore Detritus Compartments

Uniform 10 10 10 10 10 10 60
Pyramid 25 15 7 3 9 1 60
Inverted 3 12 28 6 9 1 60

Uniform 20 20 20 20 20 20 120
Pyramid 51 30 15 6 17 1 120
Inverted 7 25 57 12 18 2 120

Uniform 50 50 50 50 50 50 300
Pyramid 128 75 37 15 43 2 300
Inverted 17 62 142 31 44 5 300

for this work (Table 3). To assess possible connections between
varying taxa, we categorized taxa from these 17 ecosystems
into the six categorical food groups. Research was completed
on each taxa represented in each ecosystem to determine diet.
For example, within the NE US Shelf ecosystem there are 81
taxa represented; one taxa being that of polychaetes, which
consists of many species with differing diets; and therefore,
may be classied as both detrital feeder and omnivore. If the
taxa fell into multiple food categories, like the polychaetes
in the NE US Shelf ecosystem, then they were represented
in each compartment. Another example, zooplankton within
the Chesapeake Bay ecosystem, which may be herbivorous
or carnivorous, was presented in both the herbivore and car-
nivore food categories. Therefore, the total number of taxa
in all food groups may be greater than the actual number
of taxa accounted for within the ecosystem. Those ecosys-
tems that exhibited more producers than any other food group
were considered to be a traditional pyramid system; those
ecosystems that exhibited more carnivores than any other
food group were considered to be an inverted system (Table 4).
Surprisingly, only 3 of the 17 empirical food-web data sets fell
in the former category, while 14 were in the latter. The per-

Fig. 3 Amplication occurrence for increasing network size. Each data point represents the total number of amplication
occurrences per 50 simulations of a given structure: (a) uniform distribution, (b) pyramid distribution, and (c) inverted
pyramid.
 e c o l o g i c a l m o d e l l i n g 2 0 8 ( 2 0 0 7 ) 286294 291

Table 6 Average network properties of the 2500 simulations, 50 structures for each class, in which each structure was
used to generate 50 different ow patterns
Trophic structure Connections Cycling index Network Network Network Indirectdirect
amplication homogenization synergism ratio

N = 60
Uniform 659.26 0.1799 0.1468 2.2068 2.3448 9.1029
Pyramid 495.3 0.1459 3.6524 1.4701 2.3972 3.9612
Inverted 848.92 0.1381 3.2656 1.6315 2.4390 6.0989

N = 120
Uniform 2584.2 0.1937 0.0004 2.4800 2.2258 18.1194
Pyramid 1895.3 0.1470 1.8496 1.4420 2.3047 6.0675
Inverted 3397.6 0.1412 1.5452 1.6975 2.3099 10.2123

N = 300
Uniform 15,803 0.2013 0 2.7432 2.1328 44.6546
Pyramid 11,644 0.1687 0.3896 1.4999 2.2652 13.6830
Inverted 20,977 0.1486 0.2216 1.8252 2.2319 22.3924

centages of the number of taxa within each food group were
averaged within pyramid and inverted systems. The ratio of
each group to one another was then used to construct net-
works of 60, 120 and 300 compartments (Table 5) for each
generalized distributionpyramid and inverted pyramid (i.e.,
120-compartment pyramid structures had an average func-
tional group distribution of [P, H, C, O, D, F] = [51, 30, 15, 6, 1,
17], while inverted had a distribution of [P, H, C, O, D, F] = [7,
25, 57, 12, 2, 18]). These were then compared to the uniform
distribution, a hypothetical system where 20 taxa are in each
food group (functional group distribution matrix [P, H, C, O, D,
F] = [20, 20, 20, 20, 20, 20]).
In the rst analysis, we started with 120-compartment
models, in which a particular trophic distribution was used to

Fig. 4 Indirect-to-direct ratio for increasing network size. Each data point represents the average I/D ratio for 50
simulations of a given structure: (a) uniform distribution, (b) pyramid distribution, and (c) inverted pyramid.
292 e c o l o g i c a l m o d e l l i n g 2 0 8 ( 2 0 0 7 ) 286294

create the network structure. Average matrix values of pyra-
mid and inverted systems, as well as a uniform system, were
assessed. To obtain average values for each system, each of
the normalized functional group diet distributions were used
to construct a corresponding network structure (adjacency
matrix). This structure was then used to generate constrained
random ow values, meaning they were random, but con-
strained by physical limitations such as mass conservation
and dissipation (see Fath, 2004). The ow information was
used to calculate the cycling index (Finn, 1976), presence of
network amplication, network homogenization parameter,
network synergism parameter, and indirect-to-direct effects
ratio. This process was repeated 50 times for each network
structure, giving 50 different weighted ow networks for each
structure. Averages of the network parameters of all 50 simu-
lations were recorded. Next, for the same trophic distribution a
new structure was constructed, which again was used to cal-
culate the average network parameters for 50 different ow
values. In this manner, for the 120-compartment pyramid
trophic distribution, 2500 simulations represent the averages
of 50 ow values for each of 50 structures. This same proce-
dure was then repeated for the pyramid, inverted pyramid,
and uniform trophic distributions with 120 compartments.
In the second analysis, we repeated the same process to
generate 50 structures and 50 ow values for each struc-
ture, but here considered the cases in which the network
size was 60 or 300 compartments for each of the trophic diet
distributionspyramid, inverted, and uniform. The average
ows for each of the 50 structural matrices resulted in the
matrix data used to compare the three systems. All matrices
were calculated in MATLAB version 7.1 (Table 6).
4. Results
First, we consider the case when comparing the 120-
compartment networks. Since each of the 50 structures is an
independent random construction, the results are sorted and
plotted from lowest to greatest for each parameter value. This
does not indicate a trend in the parameter, but rather provides
a cleaner depiction of the results and their range. The cycling
index was greatest for the uniform distribution, and showed
very similar results for the pyramid and inverted pyramid con-
gurations (Fig. 2a). Network amplication did not occur in the
uniformly distributed network, but did occur occasionally in
the 120-compartment pyramid and inverted pyramid models.
Fig. 2b shows the number of occurrences of amplication for
each of the 50 runs for each trophic distribution. Homogeniza-
tion and indirectness were both highest in the networks with
uniform trophic distribution, while synergism was lowest in

Fig. 5 Synergism index for increasing network size. Each data point represents the average I/D ratio for 50 simulations of a
given structure: (a) uniform distribution, (b) pyramid distribution, and (c) inverted pyramid.
 e c o l o g i c a l m o d e l l i n g 2 0 8 ( 2 0 0 7 ) 286294
the uniform distribution (Fig. 2ce). These gures also show
that, the inverted pyramid models had higher homogeniza-
tion and indirectness than the pyramid models, yet the two
had almost identical synergism values.
In the cross-scale analysis, results for the network param-
eters are presented separately for each trophic distribution, to
clearly show the variation with changing scale. Overall, the
results were consistent with Fath (2004) in that increasing
the network size had the effect to increase homogenization
and indirectness, decrease synergism, and effectively elimi-
nate occurrences of amplication. For example, the uniform
distribution shows some occurrences of amplication for the
60-compartment network, but none when there are 120 or
300 compartments (Fig. 3a). Occurrences of amplication also
decreased in the pyramid and inverted pyramid distributions
with increasing size (Fig. 3b and c). Fig. 4 shows the increas-
ing response of indirectness to increasing network size for all
three trophic distributions while Fig. 5 shows the opposite
trend for synergism. Regardless of the trophic distribution,
larger networks have greater indirectness but lower syner-
gism.
5. Conclusions
Since Elton, the traditional view of ecological trophic levels
has been that of pyramids of energy or numbers with a larger
amount of primary producers at the bottom of the food-web
progressively decreasing until the top predator number and
biomass supported at the top of this web. The result is the
classic trophic pyramid.
The analysis presented here expands on the earlier work of
Fath (2004), which rst introduced the cyber-ecosystem model
as a community assembly rule to construct and investigate
large-scale networks. The model, which used basic trophic
constraints for constructing the network structure, was devel-
oped to allow for any number of compartments in each of
the six functional groups. Furthermore, unlike other com-
munity assembly rules, this rule generates ow values for
each of the structures thereby making it possible to calcu-
late functional network properties in addition to structural
ones. In Fath (2004), for simplicity, only a uniform trophic dis-
tribution was used to construct and investigate networks of
various sizes. The objective here was to compare this ecologi-
cally unrealistic baseline distribution with other distributions
better representing actual ecosystem congurations.
Standard ecologically theory informs us that one would
expect the greatest number within an ecosystem to be that
of the primary producers with a continual reduction in num-
bers as one proceeds up trophic levels. We used a data set of
network interactions for 17 food webs ranging in size from
n = 29 to 220. The majority of ecosystems (16 of 17) exhibited
more food group compartments than taxa represented within
the systemmultiple taxa were functioning in multiple food
groups. Contrary to standard theory, only 3 ecosystems were
traditional pyramid systems while the remaining ecosystems
(14) had a disproportionate number of higher-level trophic
groups identied resulting in inverted pyramid trophic struc-
tures. The inverted structures may be a result of minimal
biomass stored within the producers of the ecosystem as com-
293
pared to the large biomass of the consumers; however, our
current research is only relevant to the Eltonian pyramid of
numbers, not biomass. Another explanation may be the type
of ecosystems studied, since many were aquatic, which can
exhibit inverted trophic pyramids. More likely, the reason of
a large number of inverted systems (in both terrestrial and
aquatic ecosystems) may be because of the investigators focus
while sampling and constructing the websit is possible that
some groups may have been under sampled. Detritus and
detritivores are not typically accounted for in community ecol-
ogy food webs although they play an important role in the
ecosystem functioning. For example, detritus was excluded
from 5 of the 17 ecosystems sampled. The inverted pyramids
are likely a consequence of the fact that often the larger,
higher trophic level species are of greater interest to the
researchers; thereby, resulting in greater disaggregation of the
data at this level. However, by having data for both the reg-
ular and inverted pyramid structures provided us with two
different categories of ecological systems to investigate in the
cyber-ecosystem model in addition to the uniform distribution
analyzed earlier.
In conclusion, network analysis results indicate that the
trophic distribution does not have a large effect on the overall
network parameters, in that all parameters behaved quali-
tatively like the uniform distribution. All distributions also
showed the same tendencies for increasing network size.
While additional data could further rene the average distri-
bution numbers, we believe this would only marginally affect
the overall results. Lastly, we hope this research brings to
light the discrepancies between ecological theory and data
sampling in order to convey to those constructing empirical
food-web data sets the importance of partitioning the ecosys-
tem thereby creating a more accurate representation of its
constituents.
Acknowledgement
MCK was supported by the U.S. Environmental Protection
Agency in the Undergraduate Greater Research Opportunities
(GRO) Fellow while working on this research. The results do
not necessarily reect the views of the US EPA.
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