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PreprintCited by 18Plant and animal studiesInsect-Plant Interactions and ControlInsect and Arachnid Ecology and Behavior

Variation in context‐dependent foraging behavior across pollinators

Open access
Briggs, Heather M., Graham, Stuart, Switzer, Callin M., Hopkins, Robin·2017·bioRxiv
DOI 10.1101/210377PMC PMC6144987

Preprint — published as 10.1002/ece3.4303 · Paper page

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Abstract

Pollinator foraging behavior has direct consequences for plant reproduction and has been implicated in driving floral trait evolution. Exploring the degree to which pollinators exhibit flexibility in foraging behavior will add to a mechanistic understanding of how pollinators can impact selection on plant traits. Although plants have evolved suites of floral traits to attract pollinators, flower color is a particularly important aspect of the floral display. Some pollinators show strong innate color preference, but many pollinators display flexibility in preference due to learning associations between rewards and color, or due to variable perception of color in different environments or plant communities. This study examines the flexibility in flower color preference of two groups of native butterfly pollinators under natural field conditions. Our study reveals that pipevine swallowtails and skippers, the predominate pollinators of the two native Texas Phlox species, display distinct patterns of color preferences across different contexts. Pipevine swallowtails exhibit highly flexible color preferences and likely utilize other floral traits to make foraging decisions. In contrast, skippers have consistent color preferences and likely use flower color as a primary cue for foraging. As a result of this variation in color preference flexibility, the two pollinator groups impose concordant selection on flower color in some contexts but discordant selection in other contexts. This variability could have profound implications for how flower traits respond to pollinator-mediated selection. Our findings suggest that studying dynamics of behavior in natural field conditions is important for understanding plant-pollinator interactions.

AI snapshot

Written from the paper’s full text.

TL;DR

In field arrays of two Texas *Phlox* species, pipevine swallowtails showed context-dependent flower color preferences, whereas skippers were largely consistent in color preference. This variation means the two pollinator groups impose concordant selection on flower color in some contexts but discordant selection in others.

Key findings

  • Over 3 years, the authors observed 2,441 visits from 312 pipevine swallowtail butterflies and 962 visits from 308 skipper butterflies.
  • Pipevine swallowtails showed significant flexibility in color preference across floral contexts; for example, in light-red arrays they preferred light-blue flowers in within-species (WS) comparisons but preferred light-red flowers in between-species (BS) comparisons, and showed no preference in community context (CC). Similar context-dependent patterns occurred for dark-red and dark-blue arrays.
  • Skippers showed little to no flexibility: they strongly preferred light-blue over dark-red flowers in all three floral contexts, and showed no preference between light-red and light-blue regardless of context.
  • The two pollinator groups were concordant in WS contrasts but significantly different in BS contrasts for all three color-type arrays, and in CC contrasts for dark-blue and dark-red arrays; thus floral context dictates whether the two groups impose concordant or discordant selection on flower color.

Methods

  • Field experiment at Brackenridge Field Laboratory, Austin, TX, in May 2010, 2012, and 2015; potted *Phlox* plants were arranged in 4 × 5 grids alternating light-blue focal species with one other *P. drummondii* color (light-red, dark-red, or dark-blue), producing nine array types across three contexts: within-species (WS), between-species (BS), and community context (CC). Due to logistical limitations, one floral context was observed per year.
  • Free-flying butterflies were observed between 10 a.m. and 4 p.m. on at least 2 days per array; visits were recorded only when the proboscis entered a corolla, and pollinators were identified as pipevine swallowtail, skipper, or “other.” Other pollinators (~5% of total visits) were excluded.
  • Data were analyzed with binomial GLMMs in the lme4 R package, modeling visits to light-blue versus other-color flowers with fixed effects of floral context, pollinator type, and other *P. drummondii* flower color, plus random effects of pollinator individual and date. The full three-way interaction model was best-fit by likelihood ratio test; post hoc pairwise comparisons used glht() in multcomp, without multiple-test correction because contrasts tested targeted a priori hypotheses.

Limitations

  • Stated by the authors: only one floral context was collected per year, so year or abiotic effects are possible; the study does not represent the full breadth of plant contexts pollinators encounter; skippers were not identified to species because pollinators were not destructively sampled, so species-specific preferences could be masked by pooling; and constancy was not examined because the study focused on color preference.
  • Evident from the text: the experiment includes only two pollinator groups and two *Phlox* species, uses wild butterflies with unknown foraging experience, and excludes other pollinator species (~5% of visits) due to small sample sizes and apparent inability to access floral rewards.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1101/210377

Cites

61 of 61 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

5 of 58 cited sources checked · 3 supported · 0 flagged · accuracy not yet measured

  • Premisecitation

    Flower color is a particularly important trait driving patterns of pollinator visits to plants.

    “Flower color is a particularly important trait driving patterns of pollinator visits to plants”
    Quote in the paperNot a substantial claim1 of 3 sources checked
    • cites Dötterl et al., 2014 source
    • cites Ômura & Honda, 2005

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites von Frisch et al., 1914

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Findingown data

    Pipevine swallowtail flexibility in color preference is evident in each of the three flower color comparisons.

    “This flexibility is evident in each of the three flower color comparisons (Figure 3, Table 1).”
    Quote in the paperSubstantial claim
  • Findingown data

    The model that best predicted color preference of the two butterflies was the full model that included a three-way interaction between the fixed effects (foraging context, pollinator type, and flower color type).

    “The model that best predicted color preference of the two butterflies was the full model that included a three‐way interaction between the fixed effects (foraging context, pollinator type, and flower color type).”
    Quote in the paperSubstantial claim
  • Findingown data

    For pipevine swallowtails, the strength and direction of preference are significantly different between floral contexts.

    “For pipevine swallowtails, the strength and direction of preference are significantly different between floral contexts.”
    Quote in the paperSubstantial claim
  • Findingown data

    Pipevine swallowtails exhibit highly flexible color preferences.

    “Pipevine swallowtails exhibit highly flexible color preferences and likely utilize other floral traits to make foraging decisions.”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. 23
  • May combine two claims. 32
  • The quote only partly states this claim. 1

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Variation in the degree to which flower color contrasts with the background can drive flexibility in color preference.

    “variation in the degree to which flower color contrasts with the background can drive flexibility in color preference”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Osorio & Vorobyev, 2008 source

      Section 3, “Animal colour vision and the colours of natural objects” (page unavailable), discusses bees detecting flowers against leaves and discriminating among flower colours. It does not say that variation in flower–background contrast drives flexibility in colour preference; searches for that relationship found no supporting passage. (LLM, conf=0.82)

      Experimental — accuracy not yet measured

  • May combine two claims.

    Premisecitation

    Wild lepidopterans are important but understudied pollinators.

    “Wild lepidopterans are well suited to investigating flexibility in color preference in a natural field setting because they are important but understudied pollinators”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Rader et al., 2016 source

  • May combine two claims.

    Premisecitation

    Lepidopterans exhibit variation in visual systems across families and even species, which could translate to differential selection pressures on flower color within a given community of co-occurring pollinators and plants.

    “They exhibit variation in visual systems across families and even species, which could translate to differential selection pressures on flower color (and other traits) within a given community of co‐occurring pollinators and plants”
    Quote in the paperSubstantial claim1 of 2 sources checked
    • cites Briscoe, 2008 source
    • cites Stavenga & Arikawa, 2006 source

      The Introduction says differences in butterfly eyes suggest that color-discrimination capacity varies among species, and §3.1 describes different visual-pigment diversification in several butterfly families. The paper does not address whether those differences create differential selection pressures on flower color in communities of co-occurring plants and pollinators. (LLM, conf=0.84)

      Experimental — accuracy not yet measured

  • May combine two claims.

    Premisecitation

    The composition of the pollinator visitors can vary both spatially and temporally.

    “Furthermore, the composition of the pollinator visitors can vary both spatially (Gomez, Abdelaziz, Lorite, Jesús Muñoz‐Pajares, & Perfectti, 2010) and temporally (CaraDonna et al., 2017), further complicating our understanding of how pollinator variation drives patterns of selection on floral traits.”
    Quote in the paperNot a substantial claim1 of 2 sources checked
    • cites Gomez, Abdelaziz, Lorite, Jesús Muñoz‐Pajares, & Perfectti, 2010

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites CaraDonna et al., 2017 source

  • May combine two claims.

    Findingown data

    Skippers in the dark-blue arrays exhibit preference for light-blue flowers in two of the three contexts and weak to no preference when light-blue P. cuspidata is paired with dark-blue P. drummondii.

    “Skippers in the dark‐blue arrays exhibit preference for light‐blue flowers in two of the three contexts and weak to no preference when light‐blue P. cuspidata is paired with dark‐blue P. drummondii (SI).”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    In the light-red arrays, pipevine swallowtails have a preference for light-blue flowers when the two flower colors are the same species (WS) but preference for light-red color when the two flower colors are different species (BS).

    “For example, in the light‐red arrays, pipevine swallowtails have a preference for light‐blue flowers when the two flower colors are the same species (WS) but preference for light‐red color when the two flower colors are different species (BS).”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    These butterflies show preference for light-blue over dark-red under one floral context, no preference in another context, and preference for dark-red over light-blue in the third context.

    “For example, we found that these butterflies show preference for light‐blue over dark‐red under one floral context, no preference in another context, and preference for dark‐red over light‐blue in the third context.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    In our study, pipevine swallowtails preferred blue flowers of one species (P. drummondii), were strongly deterred by the blue flowers of another co-occurring species (P. cuspidata), and ultimately displayed preference for blue flowers based on species identity.

    “In our study, we found that while pipevine swallowtails preferred blue flowers of one species (P. drummondii), they were strongly deterred by the blue flowers of another co‐occurring species (P. cuspidata) and ultimately displayed preference for blue flowers based on species identity.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    Our study reveals that two butterfly groups that provide the majority of pollination visitation to two native wildflowers display different flexibility in color preference and, in the case of the pipevine swallowtail, behave in ways that might be difficult to predict from laboratory studies.

    “Our study reveals that two butterfly groups that provide the majority of pollination visitation to two native wildflowers display different flexibility in color preference and, in the case of the pipevine swallowtail, behave in ways that might be difficult to predict from laboratory studies.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The two Phlox wildflower species used in the experiment depend on these pollinators for as much as 95% of their pollination visits.

  • May combine two claims.

    Findingown data

    In this study, in some foraging contexts, pipevine swallowtails and skippers exhibit the same color preferences, and in other contexts, the two pollinators display disparate color preferences.

    “In this study, we found that in some foraging contexts, pipevine swallowtails and skippers exhibit the same color preferences, and in other contexts, we found that the two pollinators display disparate color preferences.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductreasoning

    Due to logistical limitations, foraging data were collected on one floral context per year.

    “Due to logistical limitations, we collected foraging data on one floral context per year”
    Quote in the paperNot a substantial claim

  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    Because arrays contained equal numbers of each compared color, pollinator preference could be measured as the proportion of total floral visits to the light-blue focal species.

    “Because arrays contained equal numbers of each compared color, pollinator preference could be measured as the proportion of total floral visits to the light‐blue focal species.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    Butterfly foraging behavior generates selection on flower color and maintains the flower color polymorphisms across the P. drummondii range.

    “Butterfly foraging behavior generates selection on flower color and maintains the flower color polymorphisms across the P. drummondii range.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Hopkins & Rausher, 2011, 2012, 2014

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Classified as probably not a substantial claim.

    Study design and conductcitation

    We did not adjust for multiple tests because our contrasts were targeted to test specific a priori hypotheses.

    “We did not adjust for multiple tests because our contrasts were targeted to test specific a priori hypotheses”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Benjamini, 2010 source
    • The authors state this as an assumption: Implementing a correction for multiple tests does not change the interpretation of our results.

  • May combine two claims.

    Premisecitation

    Lepidopterans have been shown to display innate color preferences and yet can alter preference through learning.

    “lepidopterans have been shown to display innate color preferences and yet can alter preference through learning”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Blackiston et al., 2011 source
    • cites Kandori et al., 2009

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Interpretationreasoning

    The two pollinator groups impose concordant selection on flower color in some contexts but discordant selection in other contexts.

    “As a result of this variation in color preference flexibility, the two pollinator groups impose concordant selection on flower color in some contexts but discordant selection in other contexts.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Pollinator foraging behavior has direct consequences for plant reproduction and has been implicated in driving floral trait evolution.

  • Classified as probably not a substantial claim. May combine two claims.

    Study design and conductreasoning

    Contexts (1) and (3) differ only in the presence/absence of P. cuspidata, allowing assessment of the impact of a morphologically similar coflowering species on pollinator preference.

    “Contexts (1) and (3) differ only in the presence/absence of P. cuspidata. Foraging visits to P. cuspidata were not recorded in this context. This comparison allows us to assess the impact that the presence of a morphologically similar coflowering species can have on pollinator preference.”
    Quote in the paperSubstantial claim

  • Classified as probably not a substantial claim. May combine two claims.

    Study design and conductreasoning

    Contexts (1) and (2) differ only in which Phlox species has light-blue flowers, allowing a test of the effect of species identity on light-blue phenotype preference.

    “Contexts (1) and (2) differ only in which Phlox species has light‐blue flowers. This allows us to test the effect of species identity on light‐blue phenotype preference.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    The presence of other morphologically similar flowering species or a dominant pollinator competitor can contribute to context-dependent flower color preferences.

    “the presence of other morphologically similar flowering species or presence of a dominant pollinator competitor can contribute to context‐dependent flower color preferences”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Brosi & Briggs, 2013 source
    • cites Fornoff et al., 2016

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    P. drummondii has the same light-blue flower color across much of its range, but in some eastern and central Texas populations, P. drummondii has evolved dark-red, light-red, and dark-blue flower colors.

    “P. drummondii also has the same light‐blue flower color across much of its range, but in some eastern and central Texas populations, P. drummondii has evolved dark‐red, light‐red, and dark‐blue flower colors”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Hopkins & Rausher, 2011, 2012

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    Pollinator preference leads to increased floral visitation and thus selection for the preferred flower type.

    “Pollinator preference leads to increased floral visitation and thus selection for the preferred flower type”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Aldridge & Campbell, 2007 source
    • cites Schemske & Bradshaw, 1999 source

  • Classified as probably not a substantial claim.

    Study design and conductnone given

    In the experimental arrays, each color had the same number of open flowers, ranging from 518 to 1,254 across days.

    “In total, each color had the same number of open flowers (ranging from 518 to 1,254 across days).”
    Quote in the paperSubstantial claim

  • Classified as probably not a substantial claim.

    Study design and conductreasoning

    Date of data collection was included as a random effect because samples taken on the same array type on different days could not be considered independent.

    “The date of data collection was included because samples taken on the same array type on different days could not be considered independent.”
    Quote in the paperSubstantial claim

  • Classified as probably not a substantial claim. May combine two claims. The quote only partly states this claim.

    Study design and conductown data

    Only plant visits from pipevine swallowtails and skippers were included in analyses; other pollinator species (~5% of total visits) were excluded because of small sample sizes and because their behavior suggested they could not access pollen or nectar rewards.

    “Other pollinator species (~5% of total visits) were excluded because of their small sample sizes and because their behavior on flowers suggested that they could not access the pollen or nectar rewards of the flowers.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The excluded other pollinator species could not access the pollen or nectar rewards of the flowers.

  • May combine two claims.

    Premisecitation

    In western populations, P. drummondii individuals have light-blue flowers; however, in populations sympatric with the light-blue-flowered P. cuspidata, P. drummondii has evolved dark-red flowers.

    “In western populations, P. drummondii individuals have light‐blue flowers; however, in populations sympatric with the light‐blue‐flowered P. cuspidata, P. drummondii has evolved dark‐red flowers”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Hopkins & Rausher, 2011, 2012, 2014

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    Bumblebees often display an innate preference for blue flowers but can readily learn to associate a reward with a novel flower color.

    “while bumblebees often display an innate preference for blue flowers, they can readily learn to associate a reward with a novel flower color”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Gumbert, 2000 source
    • cites Raine & Chittka, 2007

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Classified as probably not a substantial claim.

    Interpretationreasoning

    Variation in traits other than flower color could lead to context-dependent preferences in pipevine swallowtails.

    “Variation in traits other than flower color could lead to context‐dependent preferences in pipevine swallowtails.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Pollinator behavior in allopatry favors light-blue flower color and thus maintains the ancestral phenotype in western P. drummondii populations.

    “In addition, pollinator behavior in allopatry favors light‐blue flower color and thus maintains the ancestral phenotype in western P. drummondii populations (Hopkins & Rausher, 2014).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Hopkins & Rausher, 2014

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Interpretationreasoning

    Butterfly individuals of different species can both collect and perceive spectral information in different ways.

    “This means that butterfly individuals of different species can both collect and perceive spectral information in different ways.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductreasoning

    We are not able to rule out the possibility that there could be species-specific preferences that were masked by our data pooling.

    “As such, we are not able to rule out the possibility that there could be species‐specific preferences that were masked by our data pooling.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationown data

    Pipevine swallowtails could use other traits and environmental signals in addition to color to make foraging decisions and are therefore flexible in their color preference under naturally variable conditions.

    “These results suggest that pipevine swallowtails could use other traits and environmental signals in addition to color to make foraging decisions and are therefore flexible in their color preference under naturally variable conditions.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S4 (based).

    Interpretationreasoning

    This would suggest an additional mechanism through which pollinator-mediated selection acts on flower color.

    “This would suggest an additional mechanism through which pollinator‐mediated selection acts on flower color.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationown data

    Skippers use color as an important foraging cue and are relatively inflexible in their preference for particular colors.

    “This strong preference consistency suggests that skippers use color as an important foraging cue and are relatively inflexible in their preference for particular colors.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisereasoning

    Despite the wealth of information about butterfly color preference, we know little about how these behaviors translate to natural systems.

    “Therefore, despite the wealth of information we have about butterfly color preference, we know little about how these behaviors translate to natural systems.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    Skippers may be more likely than swallowtails to move between plant species with the same flower color, transferring heterospecific pollen in the process.

    “Skippers may be more likely than swallowtails to move between plant species with the same flower color, transferring heterospecific pollen in the process.”
    Quote in the paperSubstantial claim

  • Classified as probably not a substantial claim. May combine two claims.

    Premisenone given

    Many laboratory studies suggest that pollinators display innate and learned color preferences and that these preferences can be flexible.

    “Many laboratory studies suggest that pollinators display innate and learned color preferences and that these preferences can be flexible, but our study is one of few that explores the flexibility of color preference in the field.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    This variation in color preference can be due to a number of factors including differences in visual systems and/or differential learning abilities.

    “This variation in color preference can be due to a number of factors including differences in visual systems and/or differential learning abilities.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    P. drummondii and P. cuspidata have similar light-blue flowers, but differ in size (P. cuspidata has smaller flowers) and nectar amount (P. cuspidata has lower nectar volume and sugar concentration compared to P. drummondii).

    “While P. drummondii and P. cuspidata have similar light‐blue flowers, the flowers differ both in size (P. cuspidata has smaller flowers) and nectar amount (P. cuspidata has lower nectar volume and sugar concentration compared to P. drummondii) (R. Hopkins, unpublished data).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites R. Hopkins, unpublished data

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Premisecitation

    Some butterflies have red visual receptors, but skippers do not, likely leading to passive discrimination against red colored flowers.

    “While some butterflies have red visual receptors, it appears that skippers do not, likely leading to passive discrimination against red colored flowers (Briscoe & Chittka, 2001).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Briscoe & Chittka, 2001

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S2 (checkable), S4 (based).

    Study design and conductnone given

    For simplicity we decided to focus on color preference for this study.

    “While we acknowledge that constancy is very important, for simplicity we decided to focus on color preference for this study.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Dark-red flower color is favored in populations sympatric with P. cuspidata because pollinator behavior decreases costly hybridization between the two species when they have different flower colors.

    “Dark‐red flower color is favored in populations sympatric with P. cuspidata because pollinator behavior decreases costly hybridization between the two species when they have different flower colors (Hopkins & Rausher, 2012).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Hopkins & Rausher, 2012 source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationreasoning

    The indiscriminate color-based preference that skippers exhibit could have important implications for co-occurring plants in the community.

    “The indiscriminate color‐based preference that skippers exhibit could have important implications for co‐occurring plants in the community.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    At present, there are no studies examining the pipevine swallowtail visual system, but closely related species exhibit exceptional long-wavelength visual abilities.

    “At present, there are no studies examining the pipevine swallowtail visual system, but closely related species exhibit exceptional long‐wavelength visual abilities (Arikawa, 2003; Takemura, Kinoshita, & Arikawa, 2005).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Arikawa, 2003 source
    • cites Takemura, Kinoshita, & Arikawa, 2005 source

  • Classified as probably not a substantial claim.

    Study design and conductnone given

    The results are based on observing wild butterflies, which have unknown foraging experience, foraging on arrays of native plants in their natural habitat.

    “Our results are based on observing wild butterflies, which have unknown foraging experience, foraging on arrays of native plants in their natural habitat.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisereasoning

    Many butterflies display innate color preferences as well as learned associations between colors and nectar rewards.

    “From these laboratory studies, it is evident that many butterflies display innate color preferences as well as learned associations between colors and nectar rewards.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    In specialized plant-pollinator interactions, conclusions as to how a pollinator acts as an agent of selection on specific floral traits can be relatively straightforward.

    “In specialized plant–pollinator interactions, conclusions as to how a pollinator acts as an agent of selection on specific floral traits can be relatively straightforward (i.e., Muchhala & Thomson, 2009).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Muchhala & Thomson, 2009 source

  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

    Study design and conductreasoning

    This study clearly does not represent the breadth of possible plant contexts that a pollinator might encounter in the wild.

    “This study clearly does not represent the breadth of possible plant contexts that a pollinator might encounter in the wild.”
    Quote in the paperNot a substantial claim

  • Classified as probably not a substantial claim.

    Premisereasoning

    It remains unclear how results from laboratory studies translate to behavior in natural systems.

    “Therefore, it remains unclear how results from laboratory studies translate to behavior in natural systems.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationcitation

    This variation in color perception can lead to differences in innate color preferences, the ability to learn new colors associated with rewards, and the degree to which color preference will be context-dependent and influenced by the environment.

    “It is not surprising that this variation in color perception can lead to differences in innate color preferences, the ability to learn new colors associated with rewards, and the degree to which color preference will be context‐dependent and influenced by the environment (Blackiston et al., 2011; Briscoe, 2008).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Blackiston et al., 2011 source
    • cites Briscoe, 2008 source

  • Classified as probably not a substantial claim.

    Premisecitation

    Flower color variation across the range of P. drummondii is maintained by pollinator-mediated selection.

    “Previous studies in this system demonstrated that flower color variation across the range of P. drummondii is maintained by pollinator‐mediated selection (Hopkins & Rausher, 2012, 2014).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Hopkins & Rausher, 2012 source
    • cites Hopkins & Rausher, 2014

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

50 of 50

  • The paper relies on this without support.

    The extent of flexibility in pollinator floral color preference in nature is largely unknown.

    “yet, the extent of this flexibility is largely unknown”

  • The paper relies on this without support.

    Studies examining the consistency of pollinator behaviors under field conditions are lacking.

    “we are lacking studies that examine the consistency of these pollinator behaviors under field conditions”

  • The paper relies on this without support.

    Little attention has been given to how observed preference in one context might vary in other contexts.

    “little attention has been given to how observed preference in one context might vary in other contexts”

  • The paper relies on this without support.

    Very few studies have explored the extent to which butterflies are flexible in their color preference in natural systems.

    “very few studies have explored the extent to which they are flexible in their color preference in natural systems”

  • The authors state this as an assumption.

    Only five skipper species (Thorybes pylades, Erynnis horatius, Copaeodes aurantiaca, Atalopedes campestris, and Hylephila phyleas) likely visited the experimental arrays.

    “we believe, based on visual recognition, that only five of those species (Thorybes pylades, Erynnis horatius, Copaeodes aurantiaca, Atalopedes campestris, and Hylephila phyleas) likely visited our arrays”

  • The authors state this as an assumption.

    Skipper butterflies are difficult to identify on the wing, and all five species are of similar shape and size.

    “Skipper butterflies are difficult to identify on the wing, and all five species are of similar shape and size.”

  • The paper relies on this without support.

    Pollinator foraging behavior has direct consequences for plant reproduction and has been implicated in driving floral trait evolution.

    “Pollinator foraging behavior has direct consequences for plant reproduction and has been implicated in driving floral trait evolution.”

  • The authors state this as an assumption.

    Implementing a correction for multiple tests does not change the interpretation of our results.

    “Implementing a correction does not change the interpretation of our results.”

  • The paper relies on this without support.

    The excluded other pollinator species could not access the pollen or nectar rewards of the flowers.

    “their behavior on flowers suggested that they could not access the pollen or nectar rewards of the flowers”

  • The paper relies on this without support.

    The two Phlox wildflower species used in the experiment depend on these pollinators for as much as 95% of their pollination visits.

    “This experiment was performed using two Phlox wildflower species that depend on these pollinators for as much as 95% of their pollination visits, suggesting this behavioral variability has important implications for plant evolution.”

  • The paper relies on this without support.

    Most plants are visited by multiple pollinator species and the strength and direction of selection that multiple pollinators impose on floral traits are rarely assessed.

    “However, most plants are visited by multiple pollinator species and the strength and direction of selection on multiple pollinators impose on floral traits are rarely assessed.”

  • The paper relies on this without support.

    Other pollinator behaviors such as constancy are important for plant trait evolution.

    “Finally, other pollinator behaviors such as constancy are important for plant trait evolution.”

  • The paper relies on this without support.

    The frequency and pattern of pollinator foraging, as well as the composition of both pollinator and plant communities, can have a direct impact on plant reproductive success and the evolution of plant traits.

    “The frequency and pattern of pollinator foraging, as well as the composition of both pollinator and plant communities, can have a direct impact on plant reproductive success and the evolution of plant traits.”

  • The paper relies on this without support.

    Pollinator preference for particular floral traits exerts selective pressures on plants.

    “Pollinator preference for particular floral traits exerts selective pressures on plants.”
14 of 14

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

Each contradiction between two passages listed here was confirmed by an independent check.

  • Within-species skipper preference conflicts with the light-red result

    Minor

    The pollinator-contrast section says skippers strongly prefer light-blue flowers to the other P. drummondii colors in the within-species context. The context-dependent preference section instead says skippers have no preference between light-blue and light-red flowers regardless of context, which includes the within-species context.

  • Previously published observation years do not match the study years

    Minor

    The paper says observations were collected in 2010, 2012, and 2015, but then says observations from 2011 and 2012 were included in previous publications. No 2011 observations are otherwise described.

  • Skippers' dark-blue preference changes significantly across contexts

    Minor

    The claim that skippers show no significant flexibility conflicts with Table 1: their dark-blue preference differs between BS and WS (p = 0.003) and between CC and BS (p < 0.001).

3 of 3

Cites

Flagged first, then by citations

1 of the 62 references couldn’t be matched to a known paper.

Research works printed but not linked: 1

  1. 10.2307/2407034 · DOI

Cited by

Sorted by citations

Related

Authors
Briggs, H. M., Graham, S., Switzer, C. M., & Hopkins, R.
Affiliations
  • Arnold Arboretum
  • Centre d'Écologie Fonctionnelle et Évolutive (FR)
Corresponding author
  • Briggs, Heather M.
  • Hopkins, Robin
ORCID
Venue
bioRxiv
Publisher
openRxiv
Published
2017-10-27
Volume
8
Issue
16
Pages
7964-7973
ISSN
2045-7758 (ISSN-L)
DOI
10.1101/210377
PMC
PMC6144987
OpenAlex
W2952521105
Cited by
18
References
62
Citations by year
2026: 1 · 2025: 2 · 2024: 2 · 2023: 2 · 2021: 4 · 2020: 5 · 2019: 1 · 2018: 1
FWCI
2.28
Type
journal article
Language
en
Open access
Licence
cc-by
Locations
  • Ecology and Evolution (publishedVersion, cc-by)
  • bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion)
  • PubMed (publishedVersion)
  • Digital Access to Scholarship at Harvard (DASH) (Harvard University) (acceptedVersion)
  • Digital Access to Scholarship at Harvard (DASH) (Harvard University) (submittedVersion, cc-by)
  • Europe PMC (PubMed Central) (submittedVersion, other-oa)
  • PubMed Central (submittedVersion, cc-by)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
Funders
Harvard University
Topics
  • Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Plant and animal studies
  • Life Sciences › Agricultural and Biological Sciences › Insect Science › Insect-Plant Interactions and Control
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Insect and Arachnid Ecology and Behavior
Keywords
pollinator foraging behavior, flower color preference, pipevine swallowtail, Battus philenor, Hesperiidae, Phlox drummondii, pollinator-mediated selection, plant-pollinator interactions, floral trait evolution, reward learning

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-12.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-12.

Cited sources are checked by gpt-6-sol.

Statistics and arithmetic are recomputed, not judged.

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