Complexity and Fragility in Ecological Networks

Ricard V. Solé, José M. Montoya · Proceedings of the Royal Society B / arXiv · 2001 · ecology · 10.1098/rspb.2001.1767 · record sole2001-ecological-networks
tag layer · index · concepts · authors · עברית
Where this record's text came from. faceted 2026-08 by reading the locally held text (pdftext); every evidence string verified as an exact substring of that text before merging. Claim type is 'mechanism': the headline result ('robust against random removal but very fragile when selective attacks are used') is attributed throughout to the scale-free degree distribution, i.e. fragility happens because of topology; a reading as plain 'existence' is defensible and the abstract sentence stating the result verbatim is 'Quotation withheld.' The Montoya and Solé quote retains the PDF's mangled accent ('Sol´e') because the rule is character-for-character copying. The paper mentions unpublished corroboration ('Preliminary analysis of model ecosystems suggest that our results are robust (Sol´e and Montoya, unpublished)'), which is not a published replication, so replication status stays 'untested'. Figure-caption and table text in this extraction is heavily garbled by column interleaving, so no quotes were taken from it. 2026-08-12 and 2026-08-15 schema-0.3 passes: every quote verified as an exact substring of the staged text and anchored at merge time; concept terms verified present; every edge endpoint an existing node.
Some words in these quotes contain the wrong character. The PDF this text was extracted from substituted 10 characters inside a word — a currency sign or a stray accent where a letter pair belongs, as in signi¢cantly for significantly, or a single ligature glyph, as in confidence. They are valid characters in the wrong place, so they survive every encoding check. The quotes keep them for the same reason as above, and a search of this page will not find the affected words.
How to read this page. The article itself is not reproduced here — this paper is not cleared for redistribution, so what follows is the tag layer alone. The evidence sentences are withheld with the article. The table at the foot lists every tag the record carries; each row names the facet it belongs to, and the whose words column says whether the tag records the paper's own wording or the tagger's sentence about the paper. Every tag links to each other paper in the corpus carrying it. The machine-readable edition of the same thing is the TEI file beside this one, where the tags live in standOff, and the article is at its publisher via the link below. When we have changed our mind. Some entries in the table begin with a word in capitals and the reason for it. EVIDENCE CORRECTED — the finding still stands, but we had quoted the wrong sentence from the paper, and the quotation has been replaced with the right one. RETRACTED — the entry itself was wrong; it stays on the page, marked, with the reason, rather than being deleted. WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below. Each mark carries its own reason, in brackets, because the reasons differ from one entry to the next. An entry we have WITHDRAWN is removed from this page and kept in the record, with its reason. A line reading Audit and a date is a later re-reading of the entry against its whole sentence: DOUBTFUL means it was kept, and the reason says what is loose about it. Quotation marks inside the tagger's sentence. Where a tag is the tagger's own sentence, a phrase in quotation marks inside it is borrowed from the paper — the tagger's wording around it is not. The paper's own sentence is the evidence quote for that row; on this page it is withheld with the article, and the article is at its publisher. Nothing inside quotation marks anywhere on this page is ours: if it is quoted, it is the paper's. Whose words a concept carries. A concept entry records a term the paper itself uses, so the term, the sense given for it, and its evidence are the paper's own words, quoted where the paper fixes them. This is the opposite of a sense entry in the tag table, which is the tagger's sentence about how the paper uses a term. Where a concept entry could not be grounded in the paper's words, the entry says so on its own row.
Other fields define these differently the same word, defined another way in another discipline
species 3 fields · robustness 3 fields · graph 2 fields · simulation 2 fields · homeostasis 2 fields · keystone species 2 fields · and 2 more →

Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 14 concept tags on this paper, 12 are grounded in a quoted sentence. 10 carry the tagger's gloss of what the term means in this paper, which is not something the article can license.
food web ecological network robustness keystone species species node link scale-free small world random networks secondary extinctions homeostasis topology percolation food webs are robust to random species loss but fragile to targeted removal of highly-connected (keystone) species - the error-tolerance / attack-vulnerability of heterogeneous networks a power-law degree distribution makes random removal almost always hit a sparsely connected species, so little follows from it removing the most connected species instead fragments the web into disconnected clusters at a percolation threshold the scale-free shape itself survives random loss but is destroyed by directed loss, so the web loses the property that protected it network robust fragile extinction nodes connections trophic power law graph power law distribution size of the largest species cluster p-value power-law poisson-distribution mean simulation

Legend

second readingalternativesan independent re-reading of a facet, recorded whether or not it agrees
relation to other workclaim_relationshow this paper stands to another: agrees, contradicts, extends, qualifies
claim typeclaim_typethe shape of the claim: existence, mechanism, method, critique, prediction, null
concept edgeconcept_edgesan asserted relation between two concepts, with the sentence asserting it
conceptconceptsan idea the paper works with, quoted where the paper uses it
content tagcontent_tagsa finding, definition, quantity, statistic or variable named in the paper
contributioncontributionwhat the paper claims to add
controlcontrolsan experiment run to eliminate a rival explanation
limitationlimitationswhat the paper says its evidence cannot support
mechanismmechanismshow the reported effect is said to come about
methodmethodhow the paper produced its evidence
object of studyobject_of_studywhat the paper is about — the thing it investigates
phenomenonphenomenonthe effect or regularity under study
research questionresearch_questionthe question the paper sets itself
sensesensesa distinct use of a term, quoted where the paper fixes it
transfertransfersa method or concept carried in from another field

The paper

The full text is not reproduced here. This article is not cleared for redistribution, so the page carries the tag layer only. Its evidence sentences are withheld; the grounding status remains with each tag, and the article itself is at https://doi.org/10.1098/rspb.2001.1767. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 2 · claim_relations 9 · claim_type 1 · concept_edges 4 · concepts 12 · content_tags 21 · contribution 1 · controls 2 · limitations 7 · mechanisms 4 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 3 · transfers 2

Relations on this page carry a quotation for the claim. Where a relation was separately evidenced — and so tested against the vocabulary for its type — the row says so; none of 4 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studyThree of the best-described food webs in the ecological literature - Ythan estuary, Silwood Park and Little Rock Lake - treated as graphs and subjected to simulated species loss.
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object_of_studythe paper's words
research_questionHow much of an ecosystem's fragility follows from the shape of its food web rather than from the ecology of the species in it?
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research_questionthe paper's words
contributionShows that food webs inherit the robust-yet-fragile signature of scale-free networks, and turns 'keystone species' from a qualitative label into a topological one that can be computed from the web.
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contributionthe paper's words
claim_typemechanism
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claim_typethe paper's words
phenomenonfood web robustness and fragility under species removal
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phenomenonthe paper's words
methodsimulation
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also in: simulation
methodthe paper's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[0]the tagger's words
mechanismsmechanisms
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mechanisms[1]1.0the tagger's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[2]1.0the tagger's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[3]1.0the tagger's words
limitationsThe approach lacks population dynamics and is constrained by the taxonomic resolution of the webs.
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what this limits: every fragility estimate in the paper: THIS APPROXIMATION TO COMMUNITY FRAGILITY HAS OBVIOUS CAVEATS DERIVED FROM A LACK OF DYNAMICS AND TAXONOMIC LIMITATIONS
limitations[0]the tagger's words
limitationsThe analysis treats food webs as undirected, unweighted graphs.
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what this limits: any claim about who eats whom or how much: the analysis treats food webs as undirected, unweighted graphs - direction and magnitude are discarded before it begins
limitations[1]the tagger's words
limitationsOnly three food webs could be analysed, because of the scarcity of adequately resolved data.
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what this limits: the generality of the food-web results: OUR STUDY IS LIMITED TO THE THREE RICHEST AND BEST-DESCR[ibed webs], BECAUSE OF THE LIMITATIONS OF THE AVAILABLE DATA IN TERMS OF BOTH TAXONOMIC RESOLUTION AND SIZE
limitations[2]the tagger's words
limitationsThe analysis has no population dynamics in it at all - only the graph.
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what this limits: every fragility result: the analysis has no population dynamics in it, only the graph
limitations[3]1.0the tagger's words
limitationsInteractions are treated as undirected and unweighted, so who eats whom and how strongly are both discarded.
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what this limits: any claim about interaction direction or strength: OUR FINDINGS ARE BASED ON A NON-DIRECTED, AND NON-WEIGHTED GRAPH
limitations[4]1.0the tagger's words
limitationsThree food webs, chosen because they are the only ones well enough described - the sample is the whole available population, not a random draw from it.
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what this limits: the generality of the conclusion: Quotation withheld - three because three is all there was
limitations[5]1.0the tagger's words
limitationsOne of the three webs resolves only 31% of its nodes to species, so its degree distribution is partly an artefact of taxonomy.
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what this limits: the degree distribution of that web: ONLY 31% OF NODES ARE REAL SPECIES, BEING MOST OF THE NODES GENERA-LEVEL (63%) AND THE REST CORRESPONDING TO HIGHER TAXA - connectivity depends partly on how finely each group was resolved
limitations[6]0.9the tagger's words
transfersthe robustness/fragility (error- vs attack-tolerance) framework from complex networks applied to ecological food webs
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transfers[0]the tagger's words
transfersthe error-and-attack-tolerance analysis of scale-free graphs, applied to species loss in ecological communities
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transfers[1]1.0the tagger's words
controlsthe ecological graphs subjected to RANDOM node removal as well as to selective removal aimed at the most connected species
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eliminates: the fragility being a property of removing nodes at all rather than of removing the RIGHT nodes - random removal is the same operation without the targeting · outcome (stated by the tagger, no quotation): held
controls[0]0.85the tagger's words
controlsthe behaviour compared against purely random networks, where the degree distribution is Poissonian
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eliminates: a robustness property that any network would show - the random graph is the object the claim must beat, and it has to be constructed rather than observed · outcome (stated by the tagger, no quotation): held
controls[1]0.85the tagger's words
conceptsfood web
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also in: food web
concepts[0]the paper's words
conceptskeystone species
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concepts[3]the paper's words
conceptsspecies
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: species
concepts[4]1.0the paper's words
conceptsnode
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: node
concepts[5]1.0the paper's words
conceptslink
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: link
concepts[6]1.0the paper's words
conceptsscale-free
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: scale-free
concepts[7]1.0the paper's words
conceptssmall world
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: small world
concepts[8]1.0the paper's words
conceptsrandom networks
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
concepts[9]0.85the paper's words
conceptssecondary extinctions
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
concepts[10]1.0the paper's words
conceptshomeostasis
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: homeostasis
concepts[11]1.0the paper's words
conceptstopology
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: topology
concepts[12]1.0the paper's words
conceptspercolation
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: percolation
concepts[13]1.0the paper's words
sensesnetwork - a food web, whose nodes are species and whose links are trophic relations
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senses[0]1.0the tagger's words
sensesnetwork - biological and technological networks generally, sharing one graph of interaction and one response to node removal
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senses[1]1.0the tagger's words
sensesfragility - in ecology, how a community responds to losing species, which the paper reads as the graph's response to losing nodes
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senses[2]0.9the tagger's words
alternativesclaim_type: mechanism
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a second reading reaching the same value. Recorded: mechanism — note: a second reading reaching the same value from a different sentence. Not applied - a person decides.
alternatives.claim_typethe tagger's words
alternativesphenomenon: robustness of food webs to random species loss and fragility to loss of the most connected species
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a second reading DIFFERING from the recorded value. Recorded: food web robustness and fragility under species removal — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.phenomenonthe tagger's words
claim_relationsbuilds-on → scale-free network theory
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claim_relations[0]the tagger's words
claim_relationsbuilds-on → network error-and-attack tolerance results
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claim_relations[1]the tagger's words
claim_relationsbuilds-on → Montoya and Solé (2000), small-world and scale-free patterns in food webs
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claim_relations[2]the tagger's words
claim_relationsextends → Albert et al. (2000) and Jeong et al. (2000), error and attack tolerance of complex networks
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claim_relations[3]the tagger's words
claim_relationsbuilds-on → Bond (1993) and Davic (2000) on keystone species beyond top predators
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claim_relations[4]the tagger's words
claim_relationsbuilds-on → Albert, Jeong and Barabasi (2000), error and attack tolerance of complex networks
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claim_relations[5]1.0the tagger's words
claim_relationsbuilds-on → Jeong et al. 2000, the metabolic-network analysis in this same corpus
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claim_relations[6]1.0the tagger's words
claim_relationsWITHDRAWN AND REPLACED (RETRACTED AND RE-ADDED WITH THE FINDING CORRECTED) · extends → Montoya and Sole (2000), which showed food webs are scale-free
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WITHDRAWN AND REPLACED: RETRACTED AND RE-ADDED WITH THE FINDING CORRECTED. The target said the cited paper 'SHOWED FOOD WEBS ARE SCALE-FREE'. The span says '…DISPLAY SMALL-WORLD PROPERTIES (Montoya and Sole, 2000)…' SMALL-WORLD IS NOT SCALE-FREE - they are distinct properties and the difference is exactly what this literature argues about. The quote was exact, the anchor resolved and the endpoints were attested throughout: 'target' is the one field of the pair that nothing checks, and it is the fold key.
claim_relations[7]1.0the tagger's words
claim_relationsextends → Montoya and Sole (2000), which showed these networks display small-world properties
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claim_relations[8]1.0the tagger's words
concept_edgeskeystone species —operationalised-as→ topology
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strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says topology rather than trophic position determines which species are keystone; operationalised-as is loose.
concept_edges[0]the tagger's words
concept_edgesfood web —is-a→ scale-free
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strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span writes 'scalefree distributions of connections'; endpoint variant, is-a loose.
also in: food web · scale-free
concept_edges[2]the tagger's words
concept_edgesfood web —is-a→ small world
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strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but endpoint by anaphora: 'these networks' = the three food webs (previous sentences); is-a is loose.
also in: food web · small world
concept_edges[3]the tagger's words
concept_edgesEVIDENCE CORRECTED (RE-GROUNDED) · keystone species —causes→ secondary extinctions
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EVIDENCE CORRECTED: RE-GROUNDED. The old span named neither endpoint. This sentence names both and states the causal link the edge asserts - removal of keystone species producing secondary extinctions. — strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span reports extinction rates up to 95 times higher when keystone species are removed; 'secondary extinctions' is a variant.
concept_edges[5]the tagger's words
content_tagsthree — number of species-rich ecosystem food webs analysed
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content_tags.quantities[0]the tagger's words
content_tagsP(k) ≈ k^-γ — scale-free power-law degree distribution of the food webs
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about this quote: QUOTE REPAIRED: the extractor split a hyphen-space and the quote was recorded faithfully with the damage in it, which made it unanchorable. The repaired string is an EXACT SUBSTRING of the staged text - that is why it was accepted, not because a rule produced it. The text as originally recorded is kept in evidence as extracted.
content_tags.quantities[1]the tagger's words
content_tagsNs = 134 species — size of the Ythan estuary food web
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content_tags.quantities[2]1.0the tagger's words
content_tags<k> = 4.75 — average links per species in the Silwood Park web
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content_tags.quantities[3]1.0the tagger's words
content_tags<k> = 26.05 — average links per species in the Little Rock Lake web
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[4]1.0the tagger's words
content_tags95 times higher — ratio of extinction rate under directed removal to random removal
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[5]1.0the tagger's words
content_tags88% real species — taxonomic resolution of the Ythan estuary web
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content_tags.quantities[6]1.0the tagger's words
content_tagsEcological networks are extremely robust against random removal but very fragile under selective attacks.
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content_tags.findings[0]the tagger's words
content_tagsThis robust-yet-fragile structure bears on biodiversity dynamics, extinction-rate estimates, and the definition of keystone species.
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content_tags.findings[1]the tagger's words
content_tagsThe food webs show high homeostasis under random species removal, with a slow, linear decrease in the fraction of species in the largest cluster.
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content_tags.findings[2]1.0the tagger's words
content_tagsDirected removal of the most connected species is catastrophic - extinction rates run up to 95 times higher than under random removal.
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content_tags.findings[3]1.0the tagger's words
content_tagsThe scale-free distribution itself is stable under random loss of more than half the species, and collapses after only a few directed removals.
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content_tags.findings[4]1.0the tagger's words
content_tagsThe topology of the food web, rather than the trophic position of species, determines which species are keystone.
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content_tags.findings[5]0.9the tagger's words
content_tagssize of the largest species cluster
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content_tags.variables[3]1.0the tagger's words
content_tagssecondary extinctions
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content_tags.definitions[2]1.0the tagger's words
content_tagskeystone species
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content_tags.definitions[3]1.0the tagger's words
content_tagspower law distribution
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content_tags.definitions[5]1.0the tagger's words
content_tagsp-value
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: p-value
content_tags.statistics[0]the tagger's words
content_tagspower-law
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also in: power-law
content_tags.statistics[1]1.0the tagger's words
content_tagspoisson-distribution
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content_tags.statistics[2]1.0the tagger's words
content_tagsmean
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also in: mean
content_tags.statistics[3]1.0the tagger's words

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