REVIEWS

Making connectionsin foodwebs

Philip H. Warren

ood webs show evidence of Patterns in food web structure have the proportion of possible links

F structural regularities, or
patterns, in their arrange-
ments of species and feed-
ing link+. One aim of food web
ecology is to establish the causes
provided an important, though
contentious, testing ground for ideas
about the population dynamics and
energetics of multispecies systems. One
of the most debated of these patterns is
that actually occur). Plots of the
sort shown in Fig. la, showing a
roughly hyperbolic decline in con-
nectance with species number, are
well known. The same data pro-
of such patterns, and thereby to the apparent decrease in food web duce an approximately linear plot
identify processes important in connectance as the number of species in of number of links against number
community organization. Since a web Increases. Several contrasting of species (Fig. lb). Analysis of
feeding and being fed upon will, in mechanisms that might determine food such data (in the form of Fig. la)
general, have energetic and popu- web connectance have been suggested. was stimulated by theoretical find-
lationdynamic consequences for These mechanisms, in combination with ings of May10and others that, in
the organisms concerned, food new, food web data, suggest that the certain general models of food
webs have been interpreted as conventional pattern, and explanations for webs, high complexity (connect-
depicting, albeit crudely, the dy- it, may well be open to dispute. The true ante) reduced the probability of
namic structure of a community. nature of the relationship between systems being locally stable (i.e.
With the addition of various connectance and species number has able to return to an equilibrium
assumptions about the nature of implications for the explanation of other following small perturbations) and
trophic interactions (and, by web patterns and for theories of food web hence, that in real webs, com-
implication, other direct and in- structure, but a general explanation plexity should decline with an
direct interactions), the extent to remains elusive. increase in species number (see
which community dynamics might below). The models used were
be responsible for observed food based on community matrices
web patterns can be theoretically Philip Warren is at the Dept of Animal and Plant (see Box 1) with randomly gen-
Sciences, University of Sheffield, Sheffield,
explored*. Among all the patterns erated elements, including zeros,
UK SlO 2UQ.
to be documented and examined excepting the ali terms which
in this way’-3, one has assumed were set to give self limitation.
particular prominence: the idea of connectance. The biological unrealities of such models, and the conse-
Connectance (see Box 1) can be broadly defined as quences of altering the assumptions and construction of the
the number of actual interactions in a food web divided models, provided a rich theoretical vein, well-worked in the
by the total possible number of interactions - essentially following years to show that inclusion of various sorts of
the density of interactions. Why has connectance been non-randomness and biological reality did alter the stability-
seen as so important? First, community ecologists have complexity relationship6J1-14.However, parallel empirical ex-
been concerned for many years with the problem of what plorations, using food web and other data, appeared to indi-
determines the stability of a community and, in particular, cate that the simple predictions of an inverse hyperbolic
whether complex communities should be more stable relationship between connectance (C) and species richness
than simple ones (see recent reviews6s7). The density of (S) were, at least roughly, supportedlJs-la. This evidence
interactions in a food web provides one measure of com- has done much to sustain the idea of excess connectance
munity complexity, and has been extensively used in the causing instability, to the extent that May’s conditions for
complexity-stability debate6, as much for its ease of deri- stability (see Hypothesis 1 below) has been used to estimate
vation from published data, as for any biological relevances. average interaction strength from data on Sand Cl7Js.
Second, it lies at the heart of recent static models of food However, the food web data on which most of these
webs which, by applying simple constraints to the numbers analyses were based were not collected for the purpose of
and arrangements of links, appear to account for some of quantitative analyses of linkage density, and arguably pro
the patterns in web structure3. Although food web studies vide a poor basis for testing such far-reaching theory~J2s.
tend to list connectance simply as one of the patterns to In contrast, recent interest in food webs has resulted in a
be explained’-3, it is increasingly evident that food web number of studies in which webs were documented specifi-
patterns are not all independentzJ,Q. Connectance may cally for the purposes of analysing food web patternsQO,*i,
be one of the key patterns from which others follow. or compiled for these purposes from detailed, often long-
Here 1reconsider the well-established patterns in food term, data-sets from specific habitatS2-24. They include
web connectance with respect to recent data which suggest webs from a pond*O,many lakes4,21,streams**, an estuary*s,
that such patterns may be misleading, review old and new a desert area24 and a Caribbean islandg.
explanations for such patterns in the light of these data, and These webs, although far from perfect (as their com-
outline some of the possible links between connectance pilers all emphasize), have the merit that they were con-
and other aspects of food web structure. structed with the well-known criticisms of the existing
web catalogues in mind. These new webs differ from the
Does connectance decrease wlth increasing species existing catalogues in various, not always consistent, ways.
richness? A recent review8 assesses many of these differences and
The established wisdom is that the more species-rich the impact the new data may have on our perception of web
the community the lower the connectance (i.e. the smaller structure. One of the most striking differences, however,

136 o 1994,Elsevier Science Ltd TREE vol. 9, no. 4 April 1994


is that the mean number of links per species is generally
markedly higher in all the new webs than in those from
the existing catalogues8. The two established catalogues
of food webs, collected largely from the literature25,26,
have mean numbers of links per species (L/S) of 1.9 and
2.2, respectively (web sizes, 2-90 species); the values for
the newer webs range from 3.5 to 11 links per species,
with a mean of 7.6 (web sizes, lo-104 species)g. Some of
these studies, where more than one web, or version of a
web, has been generated, have also addressed the relation-
ship between species number and connectance (or links),
with the general conclusion that, in many webs, connect-
ante does not decrease with species numbers (it may even
increase), and that the number of links in a web increases
as a power function (with the power greater than one)
of the number of species*1.*7.28(Fig. lc,d). Reanalyses of
existing and new web data confirm this possibility across
the range of webs from different habitats*9130,and have
resulted in the suggestion that connectance (L/S*) is
roughly scale-invariant at an average value of about 0.1
(Ref. 30). Most explanations, however, do not make specific
predictions about expected values (except see Ref. 31)
without further, generally unknown and perhaps system-
dependent, parameters such as interaction strength.
Hypotheses for patterns in food web connectance
If our perception of such a fundamental pattern as the
number of links per species in a food web is shifting, how
does this affect ideas about what factors determine web
connectance? Ideas here can be separated into five groups:
(1) stability - overconnected webs tend to be unstable;
(2) morphology - there are biological restrictions on the
range of resource types a consumer can successfully cap-
ture; (3) artefacts - web linkage is subject to incomplete and
biased recording; (4) random effects - the observed pat-
terns are similar to those expected by chance; and (5)
compartmentation - many links are absent due to habitat
differences between organisms. These ideas do not all make
the same predictions, but neither are they mutually exclus-
ive: they may interact to produce the observed patterns.
A basic constraint to all the hypotheses is the idea of
minimum linkage. Food webs tend to be recorded as those
groups of species actually linked by feeding interactions,
omitting species or groups of species that are disconnected.
The restriction that all species should be connected to at
least one other, and no parts of a web may be disconnected,
requires a minimum linkage of L=S-1 and hence C=S-l/S2
giving SC=S-l/S which is close to constant for large S, i.e.
a roughly inverse hyperbolic relationship between S and
C(Refs 17 and 18).
Stability
Analyses of certain generalized models of randomly
assembled ecosystems show that model webs are likely
to be locally stable if i(SC)“z<l, and unstable otherwise
(where i is average interaction strength), though the
generality of this result has been questioned (Ref. 32 and
others therein). If such a constraint holds, for a web to
retain stability as S increases, either i, C, or both, must
decrease. Since i is rarely, if ever, known it is assumed to be
constant, and hence the switch from stability to instability
is defined by an inverse hyperbolic relation between S
and C (i.e. SC=constant)lo.
Morphology
Morphological arguments have been used to make
two differing predictions. These are both based on the
TREE uol. 9. no 4 April 19.94
_ REVIEWS
Box 1. Definition and measurement of connectance and
linkage density
Various definitions of connectance (C) have been used In the food web literature,
all based on dividing the actual number of links (L) by the number of possible links
(where S = number of species). These include:
(1) All actual direct trophic interactions divided by ail potential direct trophic inter-
actlons (food web, or ‘directed’, connectance41: C=LjS”
(2) As (l), but excluding cannibalism from the potential tnteractlons: C=L/[S(S-l)]
(3) All trophic interactions in the community matrix [tn which the elements rep
resent dynamic effects of species on each other, i.e. the effect (a,,) of predator (I)
on prey (j) and also the effect (a,) of prey on predator] divided by all potential inter-
actions in the community matrix:
c=2 L/[S(S-I)]
(4) As (3). but in addition assuming that each pair of species sharing one or more
resources experiences direct (interference) competltion in addition to the indirect
(exploitation) competition implied by the shared resource. Actual links are there
fore calculated as for (3) with additional links for each pair of species that don’t
feed on each other, but share one or more resources -[I.e. if I and jfeed on k then
the community matrix elements ark, ak,, a,k. a,, (feedingi and a,, and a,! (interference)
will all be non-zero].
Although most attention has concentrated on connectance, due to its explicit use
in dynamics models, more-recent food web analyses have tended to favour an
alternative statistic: the average number of lmks per species, or linkage density
(l/S).
The above definitions give quantitative credentials to what is, in reality, a very
imperfect statistic, resulting from a variety of arbitrary decisions (what species to
include in the web; how frequent an interaction must be to be included; spatial and
temporal scales of observation), and different methods for observation (gut con
tents analysis/direct observation. which favour common links and abundant
species; and laboratoty feeding trials and literature information, which ignore habl-
tat-specific effects and many effects of spatial and temporal scale)5. One caveat
that follows from these observations is that the patterns (e.g. Fig. 1) could be influ-
enced by the way the data on linkage are generated. Links assigned from feeding
trials, or by assumption about what a species can do, rather than what it is
actually doing in each separate web, provide a measure of biologically possible
connectance. This may be appropriate for some analyses, but not others. Different
biases, but similar cautions, apply to data generated by other methods.
observation that there are biological restrictions to the
range of resources an organism can be adapted to feed
on, i.e. it is difficult to be a ‘Jack of all trades’. This has
differing consequences depending on exactly how these
restrictions are manifested:
Hypothesis (1): If each species is adapted to feed on only
a fixed number (k) of other species (or other resources)
then each species contributes a fixed, maximum number
of links (kS) to the web, hence the links-species relation-
ship is linear (f.=ti)), and so C=kS/S2,which is equivalent to
SC=k (where k is a constant) and results in the hyperbolic
relationship’. Different values of k give a series of possible
inverse relationships between Sand C, all hyperbolic.
Hypothesis (2): If each species is adapted to feed on
resources in a fixed proportion of the total resource space
(i.e. to feed upon anything whose characteristics, such as
size, toughness, chemistry etc., fall within the range to
which it is adapted), then as the number of species in a
web increases, more species are likely to occur within the
part of the resource space exploited by each species; thus
the number of links per species will increase. If p is the
average proportion of resource space exploited by species
in the web, then L=pS2 and hence C=pSz/S2 (Refs 28,30).
This generates roughly constant connectance (C=p) for
any particular value of p, but a wide range of connectance
values if p varies between webs.
The two morphological hypotheses, although producing
different predictions, coincide where webs consist of very
specialist species, when, in hypothesis (2), the proportion
of resource space exploited is so small it corresponds
137
REVIEWS

webs to be created than

(4 (cl others (more, possible com-
0.30 1 0.30 -; binatorial arrangements of
links). It is these that are
.
.. likely to be observed most
:/ -
frequently in nature. The
i ’
results from this type of null
model approach depend on
whether other restrictions
are applied (such as con-
straining the arrangement
of species into trophic levels
0 10 20 30 40 50 60 70 80 90 100 0 10 20 30 40 50 60 70 80 90 100 in the web), but do indicate
Number of species Number of species a decrease in connectance
with increasing species rich-
(b) (d) ness, though not necessarily
600 600 , a hyperbolic relatioshiplsJ1.
.
.

Compartmentation
Since two organisms can
I only interact if they actually
meet, webs divided into many
distinct microhabitats (with
each species occurring in
only some of these) will be
less connected than the same
web might be if all the con-
stituent species co-occurred.
0 20 40 60 80 100 0 20 40 60 80 100 In webs with few species, it
Number of species Number of species is unlikely that these species
will be drawn from very dif-
Fig. 1.The relationships between connectance (calculated as C=L/.F) and number of species, and between number of ferent microhabitats, but
trophic links and number of species, for two data-sets. (a) and (b) show data for food webs from many different, insect- webs that have 50400 species
dominated webs collected from the literature (mainly data not compiled explicitly for food web analysisp6. (c) and (d) show may tend to come from larger
data for pelagic food webs from 50 small lakes in North America compiled as part of a single study21 (using biologically
possible links - see Box 1). Axes for each type of plot are on the same scales for the two data-sets. The solid line indicates
habitats (e.g. a lake) where
the minimum level of connectance, or number of links, required for all parts of a web to be connected. The two data-sets it is quite likely that the
illustrate, (1) the commonly accepted pattern of connectance between webs - a roughly hyperbolic decline in C with S, or species included occupy
linear increase in L with S, shown by (a) and (b), and (2) the type of pattern being suggested by more recent data -C more many different microhabitats
or-less independent of S, or a non-linear increase in I. with S, shown by (c) and (d).
and many rarely coincide,
and so rarely interact.

exactly to one other species or resource, and, in hypoth-
esis (1) k=l. Schoener29 proposes an intermediate hy-
pothesis in which the number of resources a consumer
can take (‘generalization’) is fixed [hypothesis (l)], but the
number of consumers a resource suffers (vulnerability’)
increases with S (there is a limit to the number of species
a prey can be adapted against, therefore no limit to those
it cannot be adapted against); this also predicts L/S should
increase with S.
Artefact
The labour and time involved in documenting feeding
interactions is considerable, and since the number of poss-
ible interactions increases as 9, it is harder and harder
accurately to document larger webs. Diagrammatic represen-
tation of webs will also become increasingly difficult with
large numbers of links. So, particularly if the primary goal
of the work is not complete documentation of a food web,
large webs are likely to be disproportionately incomplete,
resulting in a decrease in C with increasing S (Refs
5,23,31,32). This effect will be particularly pronounced if
the distribution of link frequencies is very unever-91.
Random effects
If links are assigned purely at random amongst species
then some numbers of links allow more, different food
Evaluating the hypotheses
The stability hypothesis was the major stimulant to
analyses of the connectance-species relationship. The
generality and utility of the conclusions drawn from these
models have been questionedzJ4JsJ2 (though see Ref. 13)
but the idea that interactive complexity may generate
instability in model systems remainsrJ2 and is supported,
at least in part, by some alternative (more global) stability
measure@ and the suggestion that connectance is lower
in webs from less environmentally variable habitatsiT.
Different approaches to analysing community stability
may, however, yield different results. For example, using
the criterion of permanence (a global stability measure
that requires simply that populations of all species in the
system tend to increase when rare) suggests that more
connected systems may have a greater number of viable
reassembly pathways and so recover more easily follow-
ing disruption34. A second major uncertainty over this idea
is the absence of information about interaction strength. If
most interactions are weak or donor-controlled (consumers
having little or no effect on their resource dynamics), then
stability may impose much less restrictive conditions on
web complexity@.
One way to measure interaction strength is through
experimental manipulationss. Although such exper-
iments are not usually designed to yield distributions of

138 TREE vol. 9, no. 4 April 1994


interaction strengths for whole webs, recent experimental
work aimed at this problem suggests that webs may be
characterized by a few interactions being strong and
many being rather weak35. In the light of these uncer-
tainties, and the recent food web data in which connect-
ante appears to be independent of species number, empiri-
cal evidence for the stability hypothesis, at least in its
simple forms, is equivocal. It is possible to reconcile the
evidence with the predictions if the extra links result from
inclusion of very weak interactions. However, data from
tropical fish webs suggest that C and S are not negatively
correlated even if infrequent (though not necessarily
weak) interactions are excluded27.
The morphological hypotheses both predict a series of
relationships. In hypothesis (1) C always declines with S.
In hypothesis (2) for webs of generalist species, C is
more-or-less independent of S; while for specialist organ-
isms the pattern, although theoretically predicting con-
stant connectance, will be constrained by the (artificial?)
restriction that no part of a web may be disconnected
from the rest, which creates a minimum bound for con-
nectance and forces a hyperbolic decrease (Fig. la). There
is certainly some support for the idea of constant connect-
ante in the new food web dataz1J73*8J0 (Fig. lc), and perhaps
some evidence of the inverse C:S pattern among special-
ists in the older data (e.g. Ref. 15). Schoener finds some
evidence for the intermediate hypothesis (the relationship
between generalization and richness being weaker than
that between vulnerability and richness)*g. There is also
evidence of system-specific variation in connectance36Q7.
However, an hypothesis that makes such a wide range of
predictions is clearly hard to evaluate.
Arguments that the pattern is, at least in part, arte-
factual5 are convincing. Paine5 showed that webs from
the same system, documented in differing detail by differ-
ent studies, followed the negative C:S pattern. More quan-
titatively, modelling the effect of field ecologists’ ‘missing’
links under various different criteria also strongly suggests
that large webs are much more likely to be recorded as
having low connectance”*J*. The new data support the view
that the existing catalogues contain many, very incompletely
documented webs, and show that where a series of webs
is documented in equivalent detail, the pattern of de-
creasing connectance is moderated or lost*lJO. The influ-
ence of sample bias is, however, not an explanation in
itself; the underlying pattern on which the bias acts could
still take various different forms.
Taking the expected number of links in a food web to
be the number producing the modal frequency of possible
alternative link arrangements was proposed as a null model
of connectance. It predicts that C will decrease initially,
becoming rapidly independent of S, with a value higher than
that observed in the original food web data31.Interestingly,
it gets much closer to a reasonable (though still high) pre
diction for some of the new data. However, although the
statistics of the approach are clear, it is much less obvi-
ous what biological interpretation can be placed on the
proposed mechanism if it is to be used to describe real
systems.
The organization of webs into compartments has been
suggested as an explanation for the decreased connect-
ante in larger webs. Data and theory both stimulate mixed
opinions over the importance of compartmentationl-3S8,
However, the idea of microhabitat differences creating
spatial heterogeneity in the occurrence of species within
a large, or even small, habitat seems reasonable. Temporal
heterogeneity could have similar effects38. Even if such
REVIEWS
heterogeneity did not result in obvious food web com-
partments, it could still result in lower connectance.
Finally, interesting (though debated? theoretical studies
of web assembly under simple energetics ‘rules’“9suggest
that levels of connectance in some webs may be consist-
ent with an assembly process in which (1) each invading
species has to include a sufficient number of species in its
diet to satisfy its energy demands, and (2) where the
energy available from each species is energy which has
not already been preemptively allocated to preceding
invaders. This sort of approach provides an interesting link
to foraging theory and to evolutionary constraints on diet
breadth, and deserves further investigation.
The biology of connectance
Although presented separately, the above explanations
are far from independent. The proximate explanation for
the occurrence of any feeding interaction must be the
combined effects of factors such as morphology, micro-
habitat use and phenology, which place two species (one
of which is physically capable of feeding on the other) in
the same place at the same time. It follows from this that
webs do not, at a general structural level, just gain or lose
links independently of the species composition (though
more subtle changes, such as diet switching or local popu-
lation specialisms, may occur). For a linkage to change,
the identity of one or more of the component species
of the web needs to change. Other hypotheses (artefacts
excepted) need to be interpreted in these terms. For
example, overconnected (unstable) webs must lose the
species causing the high connectance, and these must be
replaced by others with a more-restricted diet or subject
to fewer natural enemies, if lower connectance, and hence
stability, is to be retained (which echoes the more general
finding that the community stability and structure may
depend critically on the processes of assembly and species
turnover7JJ4). If an ultimate explanation for connectance
is to be derived, then perhaps the first question should
be whether webs are more, or less, connected than we
would expect based solely on the morphological and eco-
logical characteristics of the pool of available species?
Connectance, although generally used simply as a
whole-web statistic, does have a biological meaning in
terms of individual species: it is a measure of the average
degree of trophic generalism or specialism amongst a set
of organisms27J8. Simplistically viewed, highly connected
webs have a high proportion of generalists, webs with
low connectance have more specialists. Presuming that
adaptations to varying degrees of specialism and general-
ism (with all the complications of such definitions40) are a
function of natural selection, this provides a possible link
between individual-level selection and the emergent charac-
teristics of whole food webs. If we knew why particular
groups of species (e.g. those from stable or those from
variable habitats) were more, or less, generalist, then we
might be in a position to predict the relative connectance
of webs from each of those habitats. Consideration of gen-
eralism and specialism also suggests a further reason why
connectance might decline with increasing richness as a
community assembles; when few resources are present,
invasion by generalists will be more likely, with specialist
species more likely to invade when richness is greater,
later in the successional sequence41.
Connectance and the cascade model
Although interest in connectance was originally stimu-
lated by investigations of stability, its importance has
139
REVIEWS
more recently been emphasized as the key parameter in a
static model of food webs - the cascade modeW. This
essentially comprises two assumptions that: (1) species
are arranged in a trophic hierarchy such that species can
be fed upon by any above them, and can feed on any below
them, in the hierarchy; and (2) that links are assigned,
within this restriction, independently with a fixed density
(connectance). It provides reasonable descriptions of many
web properties (e.g. chain length, species proportions)
evident in the established catalogueszJ. However, its cen-
tral parameter, linkage density, is derived from the observed
values in these catalogues, and what evidence there is
suggests that some predictions of the model are sensitive
to the value of this parameter and depend on C declining
with S. For example, with high connectance, the model
predicts food chains that are too long20.If many webs are
actually highly connected and C is independent of S, such
failures imply imperfections in the model, but could also
indicate that other web patterns deserve more detailed
re-evaluation. For example, in some systems, maximum
chain lengths may indeed be greater than previously sup
posedsJO. Such questions notwithstanding, the cascade
model does show that some aspects of web structure may
be predictable from a knowledge of connectance and an
assumption about feeding hierarchies. This implies that
connectance may be a key parameter upon which other
web characteristics depend.
Connectance and other food web patterns
This being the case, it may be pertinent to ask what
the relationships are between connectance and other web
patterns. This question has received little explicit study,
but a number of suggestions, in relation to a selection of
documented food web pattern&s, are given in Box 2. The
most systematic analysis of the effect of constraints on
connectance on other food web patterns comes from
work by Schoener29, which uses a hybrid of the two mor-
phological hypotheses (above) in which the number of re-
sources a consumer can take is fixed, but the number of
consumers it can suffer increases with S (which generates
an increase in linkage density with S) and predicts the
effect of this on species proportions and predator:prey
ratios (Box 2).
Conclusions
It is difficult to say to what extent the connections out-
lined in Box 2 are causal, in one direction or another. But
the purpose of making such connections is to highlight the
possible non-independence of the widely cited food web
patterns. Identifying which are the independent patterns
and which are the dependent will be the critical step
and, given that many web patterns seem to be affected by
variation in connectance, exploring the consequences of
treating connectance as the independent variable may be
worthwhile. Some of the ideas reviewed here (perhaps in
combination) do have the potential for explaining con-
nectance without dependence on other web patterns.
Furthermore, interest in what determines wholecommunity
stability, and in the role of connectance in this debate, is
far from dead or unimportant7; and connectance is also
implicated in other ecologically important questions. For
example, theoretical evidence points to more-connected
communities being less-invasible3v7. Some of these ideas
may be amenable to experimental probing (e.g. Ref. 43)
but it is the abilities of each to account for the patterns
evident in new and more-detailed data that will probably
serve as the critical testing ground.
140
Box 2. Pdble links between connectance and
other food web patterns
?? Feeding loops are rare. High connectance, in the absence of any
other restrictions on link arrangement, increases the probability of gen
erating loopsii. Recently documented, highly connected webs have
more loops than found in earlier data*.
??Proportions of top species (those fed upon by no others in the web),
basal species (those feeding on no others in the web) and intermedi-
ate species (those which feed on and are fed upon by others) are inde
pendent of the number of species in the web. Webs with high
connectance seem likely to have smaller proportions of the former two
types and larger of the latter. The more links there are in a web, the
more likely a species is to have links both to it and from it (i.e. to be
intermediate). This suggestion is borne out by data from newer
webs4.9.20,24. Reanalyses of food web data also dispute the indepen-
dence of species proportions from S (Refs 29,42); such an effect (and
variations in the predator:prey ratio) could be generated if the L/S ratio
increased with S (Ref. 29).
??Omnivory (feeding on more than one trophic level) is rare. Increasing
connectance, all else beingequal, seems likely to result in an increase
In omnivory. Recent, highly connected webs have extensive omnivory8.
?? Food chains are short. High connectance seems, in theory3,20 and
from new web datas, to be associated with higher maximum and mean
chain lengths than previously thought typical. In addition, the number
of food chains in a web increases very markedly with connectance
[Goldwasser and Roughgarden give the example of a five-trophic-level
web, with five species at each level - if each species feeds on three
species on the level below (L/S=2.4) there will be 405 different food
chains; if each feeds on five species on the level below (L/S=4) there
will be 3135 different chains].
?? Webs are not compartmented within habitats. High linkage density
would reduce the probability of compartmentation, and for those new
webs in which it has been investigated, compartmentation has not
been found*.
?? Webs have predator-overlap graphs (diagrams linking every pair of
predators that have any prey in common) that are more likely to be
‘interval’ and ‘rigid circuit’ graphs than would be expected by chance in
small webs, but the effect decreases as webs become large. (For full
explanations of these properties see, for example, Ref. 7; in essence
both properties describe the extent and pattern of resource-use over-
lap amongst consumers, in particular the degree to which resource-use
overlap can be represented on a single-niche dimension3,7.) Since
more-connected webs will have greater numbers of predators sharing
prey, and high overlap seems likely to increase the chance of rigid cir-
cuitry (and hence intervality) the reported decline of these in larger
webs3 may be a reflection of the apparent decline of connectance in
those data.
Is the pursuit worthwhile? Criticisms of the over-
simplistic use of a highly imperfect food web statistic as a
basis for community theory5 are undoubtedly important,
and their relevance is borne out by the evidence from
recent webs. However, it may be premature to dismiss
entirely the importance of understanding what determines
the numbers and arrangements of trophic interactions in
natural communities, even at the crude level of presence
and absence. Although not a sufficient basis for predicting
wholecommunity dynamics, such trophic ‘scaffolding’does
define the bounds within which those dynamic effects
operate and links the evolution of species’ trophic niches
with emergent patterns in community structure. The ex-
tent to which understanding trophic organization at this
level will have significant consequences for community
ecology is not at all clear. Perhaps only by achieving such
an understanding can that judgement be made.
Acknowledgements
I am very grateful to Tim Birkhead, Kevin Caston, Steve Hall,
Richard Law, Kate Lessells, Neo Martinez, Peter Morin,
Matthew Spencer, John Spicer and two anonymous
referees for helpful comments on the manuscript.
TREE vol. 9, no. 4 April 1994
 REVIEWS

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Transmissionpatternsof eukaryotic
tra able elements:arg for
and against horizontaltransfer
Michael P. Cummings

ransposable elements - Recent studies have demonstrated that Several steps are required for

T the very name conveys

movement. This theme of
motion is reiterated by
the names given to specific groups
of transposable elements, such as
several classes of transposable elements
are widely distributed within eukaryotes.
Horizontal transmission of these
transposable elements has often been
invoked In order to explain the observed
a transposable element to be-
come successfully transferred
between species: transportation,
transcription, translation, reverse
transcription and increase in
gypsy, hobo, jockey, mariner, dong variation and relationships within and copy number. Transportation -
(Chinese for moving), roo (as in between species. These same patterns of the movement of the transpos-
kangaroo) and HMS Beagle. But variatlon and relationships, however, may able element, as DNA or RNA,
what is the extent of this move- origlnate from processes that do not from one species to another - is
ment? While it is clear that trans- involve the lateral transfer of genetlc the inaugural manoeuvre for hori-
posable elements move within a material across species. zontal transmission. How these
genome and are transmitted verti- transfers might be accomplished
cally (i.e. sexually), recent studies is subject to speculation, but there
propose that they move horizon- Michael Cummings is at the Dept of Integrative is some research supporting the
Biology, University of California, Berkeley,
tally (i.e. nonsexually) between potentiality of viruses5,6 or mites;
CA 94720 USA
species, as welli-4. At the outset as vectors. Subsequent to the in-
it should be understood that these itial transfer event, the transpos-
two basic transmission patterns are not mutually exclusive. able element has to recognize, and be recognized by, the
While a heterogeneous assemblage of genetic entities appropriate components of the new genetic environment.
is classified as consisting of transposable elements on The barriers posed by transcription, translation, and where
account of crude functional similarity (i.e. they move required, reverse transcription, are conditional on the par-
about within a genome), the evolutionary origins of these ticular type of element and the cellular milieu. These
entities and the specific mechanisms associated with their interactions are requisite to the increase in copy number,
transposition are distinct. Transposable elements can be and hence, establishment of the elements. Arguments sur-
very generally divided into two categories: the Class I rounding the issue of horizontal transmission have
elements, or retrotransposons (see Box I), and the Class involved circumstantial observations, and these obser-
II elements (see Box 2). vations are often the subject of conflicting interpretations.

TREE ~1. 9, no. 4 April 1994 G 1994. Elsevier Science Ltd 141