Convergent iridescence and divergent chemical signals in sympatric sister-species of Amazonian butterflies

Numerous ecological interactions can impact trait evolution in closely-related species in sympatry, leading to trait convergence or divergence (terHorst et al., 2018). Closely-related species often display similar suites of traits because of their shared evolutionary history (Blomberg et al., 2003). Local selection can also act as a filter and prevent trait divergence (Keddy, 1992), therefore enhancing trait similarity between closely-related species when they occur in sympatry (Chazot et al., 2014). Trait similarity among sympatric species within a given ecological niche can thus stem from retention of locally adapted ancestral traits or from evolutionary convergence (Muschick et al., 2012). Selective pressures promoting the retention of locally adapted traits within species and/or trait convergence among sympatric species can also be due to local ecological interactions: for instance, shared predation pressure may promote the convergence of predator-deterrent traits in sympatry, but allow the trait to differentiate in allopatry (Mallet, 1999).

However, when closely-related sympatric species share a given trait, either as a result of ancestry and/or convergence, they often diverge in other traits because (1) they may be partitioned in different ecological niches (Berlocher and Feder, 2002), or (2) as a result of character displacement due to reproductive interference (Grether et al., 2020) or reinforcement due to poor hybrid fitness (Butlin and Smadja, 2018). As a result, traits involved in sexual competition or mate choice tend to diverge significantly more often between species in sympatry compared to allopatry (Haavie et al., 2004; Marko, 2005).

In this study, we investigate how ecological interactions in sympatry can constrain natural and sexual selection shaping trait evolution. We specifically focus on traits submitted to both natural and sexual selection and compare differences in these traits in allopatric vs. sympatric ranges. Theoretical and empirical studies have shown that sexual selection may favor the evolution of preferences for locally-adapted traits within species (Servedio, 2004; Servedio and Boughman, 2017; van Doorn et al., 2009). For instance, the predator-deterrent coloration of poison frogs is also detected and used as mating cues by females (Reynolds and Fitzpatrick, 2007). Similarly, habitat-dependent coloration of sympatric cichlid fish is also used as a visual cue for mate recognition (Seehausen et al., 2008). Yet, sexual interactions are likely to occur between individuals from closely-related species when they live in sympatry, and similar preferences for adaptive traits may thus result in substantial reproductive interference (Gröning and Hochkirch, 2008; Soni et al., 2025). Hybrids, when produced, can be unfit, thus favoring the evolution of sexual preferences for species-specific cues, rather than locally-adapted traits (Maisonneuve et al., 2024). To determine to what extent ecological interactions shape trait evolution, it is thus necessary to compare patterns of trait evolution in sympatry and allopatry: allopatric populations indeed allow us to estimate background levels of divergence or similarity that arise in the absence of direct ecological interactions (Pfennig and Pfennig, 2009). Comparing variations in adaptive traits in sympatric vs. allopatric populations of recently-diverged species and testing the sexual preference for those traits can shed light on the selective processes targeting traits modulating reproductive isolation and co-existence in sympatry.

In butterflies, the evolution of wing color patterns can be influenced by both natural and sexual selection. The visual discrimination of wing color pattern can enable intraspecific recognition during courtship in many species (Costanzo and Monteiro, 2007; Li et al., 2017). However, the evolution of wing color patterns is also strongly influenced by the risk of detection and/or recognition by predators (Finkbeiner et al., 2014; Oliver et al., 2009). Whether these opposite selective pressures ultimately promote trait convergence or divergence in sympatric species might depend on their relatedness: for instance, a study in Papilionidae showed multiple color pattern convergences between distantly-related species living in sympatry, while divergent colorations are found in closely-related species (Puissant et al., 2023). Divergence in traits involved in species recognition could be favored because of higher reproductive interference in closely related species than in distantly related taxa (Pfennig and Pfennig, 2009). In sympatric species with chemical defenses, such as Heliconinii butterflies, local predation pressures tend to promote the convergence of similar conspicuous warning wing patterns compared to allopatric species (i.e. Müllerian mimicry, Joron et al., 1999; Merrill et al., 2014). But the costs associated with hybrid production, in turn, favor the evolution of alternative divergent mating cues in mimetic butterflies (Estrada and Jiggins, 2008), and divergence in male pheromone bouquets and female attraction has been found among mimetic sister species (González-Rojas et al., 2020). Similarly, the evolution of specific visual mate recognition signals, limiting reproductive interference but indistinguishable by predators, can also be promoted on the wings of mimetic butterflies (Llaurens et al., 2014).

Here, we focus on the evolution of mating cues in the neotropical butterfly genus Morpho, where multiple closely-related species co-exist in sympatry (Blandin and Purser, 2013). In the Morpho species observed in the understory, striking iridescent blue coloration is displayed on the dorsal side of the wings, due to specific wing scale structures (Giraldo et al., 2016; Siddique et al., 2013). The light signal reflected by iridescent surfaces can be very directional, as hue and brightness of iridescent objects can drastically change depending on the light environment or the observer’s position (Doucet and Meadows, 2009). While the iridescent blue color is probably ancestral to the diversification of the understory clade (Chazot et al., 2021), the precise reflectance spectra at different angles likely differ among Morpho species. Directional iridescent signals generated in animals can likely enhance recognition by mates while remaining poorly detected by predators (Endler, 1992). In birds (Simpson and McGraw, 2019) and butterflies (White et al., 2015), the specific directional signal produced by the iridescent trait can be used as a cue during courtship, suggesting that the antagonistic sexual and natural selective pressures may finely tune the evolution of iridescent effects. How much sexual selection shapes the evolution of iridescent properties in sympatric Morpho species is currently unknown, but behavioral experiments carried out in the field in Amazonian Peru highlighted strong visually-based territorial interactions among males from sympatric species and limited species discrimination based on female coloration in males (Le Roy et al., 2021b).

This raises questions on the key visual cues involved in mate choice, given that the iridescent blue coloration shared by closely-related species encountered within the understory is also likely under selection by predators. The iridescent bright blue dorsal coloration of Morpho wings contrasts with the brown and matte ventral side and generates a peculiar visual effect during flight. The combination of the blue flashes produced by the alternate exposure of the bright blue vs. brown sides of the wings during flapping flight, in addition to erratic flight trajectories, makes these Morpho very difficult to catch by bird predators (Young, 1971), potentially enhancing their evasive capabilities (Murali and Kodandaramaiah, 2020). Experimental trials with evasive prey have shown that predators learn to avoid prey they repeatedly fail to catch (Páez et al., 2021). The display of iridescent wings could thus be associated with a higher survival rate in nature because of both (i) direct effects, through successful escape of predator attacks, and (ii) indirect effects, by limiting predation attempts by birds recognizing the blue signals and refraining from attacking, as highlighted by butterfly release experiments investigating the hunting behavior of wild insectivorous birds in Brazil (Pinheiro et al., 2016). Mark recapture experiments in the field with manipulated dorsoventral contrasts in wild Morphos have suggested that dynamic flash coloration can reduce predation rate (Vieira‐Silva et al., 2024). This indirect effect could promote the evolution of convergent blue patterns in sympatric species, similar to the mimicry observed in species with chemical defenses (Joron et al., 1999; Merrill et al., 2014). In line with this hypothesis, repeated local convergence in the proportion of iridescent blue vs. black areas on the dorsal side of the wings has been documented in the sister-species Morpho helenor and Morpho achilles living in sympatry throughout the Amazonian basin (Llaurens et al., 2021). Precise quantification of variations in iridescence is now needed to assess the respective effects of selection by predators and mates that may drive convergent vs. divergent evolution of iridescence in sympatric and allopatric ranges.

First, we quantified iridescence in allopatric vs. sympatric populations of M. helenor subspecies. Since coloration is expected to be more similar within than among species under neutral evolution, we used allopatric populations of M. helenor as a baseline to assess convergence of iridescence between two sympatric species (M. helenor and M. achilles). We then conducted behavioral experiments to test the effect of variation in iridescence on mate recognition, using two subspecies of M. helenor displaying different iridescent phenotypes. This intraspecific comparison allows identifying the visual cues used in mate choice in M. helenor, teasing apart the effects of iridescence and/or wing pattern. We then tested whether those visual cues are used in species recognition between sympatric M. helenor and M. achilles. Finally, we studied variations in the volatile compounds produced by wild males and females from sympatric populations of the two species to explore the evolution of potentially alternative traits, such as chemical cues, possibly acting as a reproductive barrier.

We investigated differences in wing iridescence, color perception, and color pattern-based mate choice of two sister species of Morpho, M. helenor and M. achilles, which diverged ~3.6 Mya (Chazot et al., 2021). We relied on geographic variations of wing color patterns within M. helenor (Figure 1) to compare the iridescence of three different populations of M. helenor (subspecies M. helenor bristowi, M. helenor theodorus, and M. helenor helenor) and test the perception of these patterns by predators and mates using visual models.

Figure 1

Download asset Open asset

The convergent iridescent signals of sympatric Morpho species are not differentiable

We measured the reflectance of Morpho wings at different angles of observation and illumination, in two different planes (referred to as proximo-distal and antero-posterior planes) following two complementary protocols (referred to as ‘specular’ and ‘tilt’; see Figure 2A and Appendix 1 for extended methods). We then extracted for each spectrum three commonly used colorimetric variables: Hue, Brightness, and Chroma (analyzed in Figure 2—figure supplement 1).

Figure 2 with 2 supplements see all

Download asset Open asset

We found that hue and brightness always vary depending on the angle of incidence of the light, as confirmed by PERMANOVAs detecting a significant effect of the ‘Angle’ variable for all methods of measurements (Figure 2; statistical analyses in Appendix 2—table 1 and Figure 2—figure supplement 2) confirming that in both species, both sexes are iridescent. Second, the brightness of Morpho wings is highly sexually dimorphic, as the effect of sex was always significant. The brightness measured with the ‘tilt’ protocol (Figure 2F and G) showed that males tend to be brighter than females in all populations.

To finely appraise the complete spectrum of variation in iridescence within and among Morpho species, we then used a multivariate approach allowing us to analyze the full reflectance spectra obtained from both proximo-distal and anteroposterior measurements. At the intraspecific level, significant divergence of iridescence was detected between allopatric subspecies of M. helenor (Figure 3A). In contrast, iridescence did not differ between species living in sympatry (PERMANOVA, F=2.09, df = 1, p-value = 0.084), suggesting that iridescence converges in sympatry (Figure 3B). In both cases, the interaction term taxon*sex was non-significant, suggesting that the sexual dimorphism of iridescence was similar for all taxa.

Figure 3 with 2 supplements see all

Download asset Open asset

We then modeled the perception of color contrast in the visual systems of M. helenor and UV-sensitive birds respectively, and tested whether the differences of reflectance observed between sexes and species at different angles of illumination could potentially be perceived by a Morpho observer or a putative predator. As sexual dimorphism of wing reflectance was detected, we measured the achromatic distances and chromatic distances for males and females separately. Achromatic distance results (Figure 3—figure supplement 1) suggest that neither birds nor Morpho butterflies could theoretically differentiate the brightness of the wings of sympatric or allopatric individuals, based on visual modeling. Concerning chromatic distances, as the results were similar along the two directions of measurements (antero-posterior and proximo-distal), only proximo-distal results are shown in Figure 3C (antero-posterior results are shown in Figure 3—figure supplement 2). Results suggest that Morpho butterflies and predators can theoretically visually perceive the difference in the blue coloration between the different subspecies of M. helenor: mean chromatic contrasts were always higher than 1 JND, for both sexes and for all angles, using both bird and Morpho visual models. However, this effect was stronger for females. In males, confidence intervals were only strictly significant for the 0° illumination using the Morpho visual model, suggesting a weak discrimination at most angles of observation.

In contrast, the mean chromatic contrasts measured between sympatric species were always below 1 JND: the minute differences observed between sympatric species are thus unlikely to be discriminated either by Morpho or bird predators.

Evidence of mate preferences based on visual cues within species

Experimental evidence has shown that M. helenor males are attracted to blue coloration when patrolling in the wild (Le Roy et al., 2021b). We thus performed a series of mate choice experiments to determine the nature of the visual cues used in mate choice in M. helenor. We focused on the two previously studied allopatric subspecies of M. helenor (M. h. bristowi and M. h. theodorus), which significantly differ in both wing pattern and iridescence.

We first tested whether mating tended to be preferentially assortative by conducting a tetrad experiment involving one male and one female of both subspecies. The highest number of mating events occurred between the two assortative pairs (see Table 1). Indeed, a Fisher’s exact test performed on the contingency table showed a significant departure from random mating (p-value = 0.04). The time until an assortative mating (mean = 48 min, sd = 38 min) or until a disassortative mating (mean = 48 min, sd = 38 min), however, did not significantly differ (T-test; stat = –0.27, p-value = 0.79). Note that the effects were clearly driven by the bolder behavior of M. h. bristowi individuals, as half of the mating events occurred between a female and a male of this subspecies.

Table 1

	M. h. theodorus female	M. h. bristowi female
M. h. theodorus male	6	3
M. h. bristowi male	5	16

To test whether this assortative mating was driven by visual cues and to identify these cues, we performed a series of experiments using dummy butterflies treated with hexane to remove any olfactory cue. We specifically focused on studying allopatric M. h. theodorus and M. h. bristowi because they both differ in their iridescent signal and in their iridescent pattern. Different pairs of female dummies were presented to M. helenor males to investigate assortative preference between subspecies (experiment 1), discrimination of the wing pattern (experiment 2), and discrimination of the iridescent coloration (experiment 3; see the experimental details in the Materials and methods section).

Experiment 1 tested male assortative visual preference using two dummies each displaying the wings of a female of a different M. helenor subspecies. M. h. theodorus males did not show any preference for any female dummy, as shown by the probability of approaches and touches that were not different from the random 0.5 expectation (Binomial GLMM approaches: Estimate = −0.121 ± 0.091 SE, z=−1.332, p-value = 0.183, Figure 4A; touches: Estimate = 1.600 ± 1.094 SE, z=1.462, p-value = 0.144, Figure 4B). M. h. bristowi males, in contrast, approached and touched a con-subspecific M. h. bristowi dummy female significantly more often than expected by chance alone (Binomial GLMM approaches: Estimate = 0.430 ± 0.105 SE, z=4.11, p-value <0.0001, Figure 4A; touches: Estimate = 1.646 ± 0.273 SE, z=6.029, p-value <0.0001, Figure 4B), suggesting a preference toward the M. h. bristowi females based on the recognition of visual cues alone.

Figure 4 with 2 supplements see all

Download asset Open asset

Experiment 2 tested male pattern-based discrimination, using two dummies made with the wings of M. h. bristowi females (i.e. same iridescence), but one of which was modified so as to present the thin blue bands found in M. h. theodorus (i.e. a different pattern). M. h. bristowi males tended to approach and touch significantly more often the wild-type M. h. bristowi female model as compared to the modified M. h. bristowi female model exhibiting a theodorus pattern (Binomial GLMMs approaches; Estimate = 0.599 ± 0.089 SE, z=6.751, p-value <0.0001, touches; Estimate = 2.805 ± 0.944 SE, z=2.972, p-value = 0.003, see Figure 4—figure supplement 1). As both female dummies shared the same blue iridescence, the choice of M. h. bristowi males was likely influenced by the difference in the pattern displayed by the dummy females, suggesting that the color pattern is used by males during mate choice.

Experiment 3 tested male color-based mate discrimination using two dummies made with the wings of females from the two subspecies (i.e. different iridescence), but those of M. h. bristowi were modified so as to resemble M. h. theodorus (i.e. same pattern). M. h. theodorus males tended to approach and touch significantly more often the M. h. theodorus female wild-type model compared to the artificially modified, theodorus-like M. h. bristowi female dummy (Binomial GLMMs on approaches; Estimate = 0.348 ± 0.121 SE, z=2.867, p-value = 0.004, on touches; Estimate = 1.172 ± 0.465 SE, z=2.521, p-value = 0.012, see Figure 4—figure supplement 2). As both dummy females shared the same color pattern, the preference expressed by M. h. theodorus males likely stemmed from the differences in the iridescent blue color, suggesting color-based discrimination in males.

In experiments 2 and 3, modified dummies were each time strongly rejected by males. Although all dummies contained traces of black marker to control for manipulation and treatment, we cannot entirely rule out that the artificial modifications performed on dummies could generate a specific visual rejection by males, because we cannot be sure whether butterflies perceive these modifications as equivalent to natural coloration. These experiments nevertheless suggest that visual cues – color pattern and iridescence – jointly contribute to mate choice in M. helenor.

Divergent chemical profiles observed between males of the sympatric species M. helenor and M. achilles

We finally investigated the volatile chemical compounds extracted from the genitalia of males and females from the mimetic sympatric species M. helenor and M. achilles. The nMDS analyses (Figure 5) suggested a strong divergence in the chemical profiles between males of the two species, especially for C16 to C30 compounds. This result contrasts with the strong similarity of chemical profiles found between the females of these two species in these longer-chain compounds (Figure 5—figure supplement 1).

Figure 5 with 1 supplement see all

Download asset Open asset

The PERMANOVA of the chemical spectra of M. h. helenor and M. a. achilles genitalia revealed a significant effect of sex, species (p-value <0.001 on both the C8 to C16 and the C16 to C30 datasets, see Figure 5) and a significant interaction between the two variables. When analyzing the results for each sex separately (post-hoc), we found a significant effect of the species variable (PERMANOVA, df = 1, F=10.013, p-value = 0.001 for males and df = 1, F=1.928, p-value = 0.049 for females in the C8 to C16 compounds, PERMANOVA, df = 1, F=26.031, p-value = 0.001 for males and df = 1, F=2.776, p-value = 0.048 for females in the C16 to C30 compounds), suggesting a substantial divergence between species in chemical bouquets, especially marked in males.

Next, we performed an Indicator Value Analysis to identify the different compounds significantly associated with each species, separating males and females (Appendix 3—table 1). Among all the annotated compounds, we found traces of beta-ocimene, previously identified as a butterfly pheromone. Interestingly, different proportions of this compound were found on the genitalia of M. helenor and M. achilles males (Appendix 3—table 1).

All raw measurements of wing reflectance and R scripts required to generate the results are publicly available under the DOI 10.5281/zenodo.14389631, or on GitHub.

1. 1. Ledamoisel J

   (2025) Zenodo

   Wing reflectance of 2 populations of sympatric and allopatric Morpho butterfly sister-species.

   https://doi.org/10.5281/zenodo.14388104

1. 1. Allen CE
   2. Zwaan BJ
   3. Brakefield PM

   (2011) Evolution of sexual dimorphism in the Lepidoptera

   Annual Review of Entomology 56:445–464.

   https://doi.org/10.1146/annurev-ento-120709-144828

   • PubMed
   • Google Scholar
2. 1. Bacquet PMB
   2. Brattström O
   3. Wang HL
   4. Allen CE
   5. Löfstedt C
   6. Brakefield PM
   7. Nieberding CM

   (2015) Selection on male sex pheromone composition contributes to butterfly reproductive isolation

   Proceedings. Biological Sciences 282:20142734.

   https://doi.org/10.1098/rspb.2014.2734

   • PubMed
   • Google Scholar
3. 1. Baldwin J
   2. Johnsen S

   (2009) The importance of color in mate choice of the blue crab Callinectes sapidus

   The Journal of Experimental Biology 212:3762–3768.

   https://doi.org/10.1242/jeb.028027

   • PubMed
   • Google Scholar
4. 1. Bennett ATD
   2. Théry M

   (2007) Avian color vision and coloration: Multidisciplinary evolutionary biology

   The American Naturalist 169:S1–S6.

   https://doi.org/10.1086/510163

   • Google Scholar
5. 1. Berlocher SH
   2. Feder JL

   (2002) Sympatric speciation in phytophagous insects: moving beyond controversy?

   Annual Review of Entomology 47:773–815.

   https://doi.org/10.1146/annurev.ento.47.091201.145312

   • PubMed
   • Google Scholar
6. 1. Blandin P
   2. Purser B

   (2013)

   Evolution and diversification of Neotropical butterflies: Insights from the biogeography and phylogeny of the genus Morpho Fabricius, 1807 (Nymphalidae: Morphinae), with a review of the geodynamics of South America

   Tropical Lepidoptera Research 23:62–85.

   • Google Scholar
7. 1. Blomberg SP
   2. Garland T
   3. Ives AR

   (2003) Testing for phylogenetic signal in comparative data: behavioral traits are more labile

   Evolution 57:717.

   https://doi.org/10.1554/0014-3820(2003)057[0717:TFPSIC]2.0.CO;2

   • Google Scholar
8. 1. Bosi SG
   2. Hayes J
   3. Large MCJ
   4. Poladian L

   (2008) Color, iridescence, and thermoregulation in Lepidoptera

   Applied Optics 47:5235–5241.

   https://doi.org/10.1364/ao.47.005235

   • PubMed
   • Google Scholar
9. 1. Bouinier T
   2. Smadi C
   3. Llaurens V

   (2025) Vertical and temporal niche partitioning in Amazonian butterflies: implications for the evolution of thermal tolerance

   Oikos 01:e11567.

   https://doi.org/10.1002/oik.11567

   • Google Scholar
10. 1. Briscoe AD

    (2008) Reconstructing the ancestral butterfly eye: focus on the opsins

    The Journal of Experimental Biology 211:1805–1813.

    https://doi.org/10.1242/jeb.013045

    • PubMed
    • Google Scholar
11. 1. Brooks ME
    2. Kristensen K
    3. Benthem KJ
    4. Magnusson A
    5. Berg CW
    6. Nielsen A
    7. Skaug HJ
    8. Mächler M
    9. Bolker BM

    (2017) glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling

    The R Journal 9:378.

    https://doi.org/10.32614/RJ-2017-066

    • Google Scholar
12. 1. Butlin RK
    2. Smadja CM

    (2018) Coupling, reinforcement, and speciation

    The American Naturalist 191:155–172.

    https://doi.org/10.1086/695136

    • PubMed
    • Google Scholar
13. 1. Caro SP
    2. Pierre L
    3. Bergès M
    4. Bakker R
    5. Doutrelant C
    6. Bonadonna F

    (2021) Mutual mate preferences and assortative mating in relation to a carotenoid-based color trait in blue tits

    Behavioral Ecology 32:1171–1182.

    https://doi.org/10.1093/beheco/arab080

    • Google Scholar
14. 1. Chazot N
    2. Willmott KR
    3. Santacruz Endara PG
    4. Toporov A
    5. Hill RI
    6. Jiggins CD
    7. Elias M

    (2014) Mutualistic mimicry and filtering by altitude shape the structure of Andean butterfly communities

    The American Naturalist 183:26–39.

    https://doi.org/10.1086/674100

    • PubMed
    • Google Scholar
15. 1. Chazot N
    2. Blandin P
    3. Debat V
    4. Elias M
    5. Condamine FL

    (2021) Punctuational ecological changes rather than global factors drive species diversification and the evolution of wing phenotypes in Morpho butterflies

    Journal of Evolutionary Biology 34:1592–1607.

    https://doi.org/10.1111/jeb.13921

    • PubMed
    • Google Scholar
16. 1. Chouteau M
    2. Arias M
    3. Joron M

    (2016) Warning signals are under positive frequency-dependent selection in nature

    PNAS 113:2164–2169.

    https://doi.org/10.1073/pnas.1519216113

    • Google Scholar
17. 1. Costanzo K
    2. Monteiro A

    (2007) The use of chemical and visual cues in female choice in the butterfly Bicyclus anynana

    Proceedings. Biological Sciences 274:845–851.

    https://doi.org/10.1098/rspb.2006.3729

    • PubMed
    • Google Scholar
18. 1. Doucet SM
    2. Meadows MG

    (2009) Iridescence: a functional perspective

    Journal of the Royal Society, Interface 6 Suppl 2:S115–S132.

    https://doi.org/10.1098/rsif.2008.0395.focus

    • PubMed
    • Google Scholar
19. 1. Dufrêne M
    2. Legendre P

    (1997) Species assemblages and indicator species:the need for a flexible asymmetrical approach

    Ecological Monographs 67:345–366.

    https://doi.org/10.1890/0012-9615(1997)067[0345:SAAIST]2.0.CO;2

    • Google Scholar
20. 1. Dyer KA
    2. White BE
    3. Sztepanacz JL
    4. Bewick ER
    5. Rundle HD

    (2014) Reproductive character displacement of epicuticular compounds and their contribution to mate choice in drosophila subquinaria and drosophila recens

    Evolution; International Journal of Organic Evolution 68:1163–1175.

    https://doi.org/10.1111/evo.12335

    • PubMed
    • Google Scholar
21. 1. Endler JA

    (1992) Signals, signal conditions, and the direction of evolution

    The American Naturalist 139:S125–S153.

    https://doi.org/10.1086/285308

    • Google Scholar
22. 1. Estrada C
    2. Jiggins CD

    (2008) Interspecific sexual attraction because of convergence in warning colouration: is there a conflict between natural and sexual selection in mimetic species?

    Journal of Evolutionary Biology 21:749–760.

    https://doi.org/10.1111/j.1420-9101.2008.01517.x

    • PubMed
    • Google Scholar
23. 1. Finkbeiner SD
    2. Briscoe AD
    3. Reed RD

    (2014) Warning SIGNALS are seductive: relative contributions of color and pattern to predator avoidance and mate attraction in Heliconius butterflies

    Evolution; International Journal of Organic Evolution 68:3410–3420.

    https://doi.org/10.1111/evo.12524

    • PubMed
    • Google Scholar
24. 1. Giraldo MA
    2. Yoshioka S
    3. Liu C
    4. Stavenga DG

    (2016) Coloration mechanisms and phylogeny of Morpho butterflies

    The Journal of Experimental Biology 219:3936–3944.

    https://doi.org/10.1242/jeb.148726

    • PubMed
    • Google Scholar
25. 1. González-Rojas MF
    2. Darragh K
    3. Robles J
    4. Linares M
    5. Schulz S
    6. McMillan WO
    7. Jiggins CD
    8. Pardo-Diaz C
    9. Salazar C

    (2020) Chemical signals act as the main reproductive barrier between sister and mimetic Heliconius butterflies

    Proceedings. Biological Sciences 287:20200587.

    https://doi.org/10.1098/rspb.2020.0587

    • PubMed
    • Google Scholar
26. 1. Grether GF
    2. Drury JP
    3. Okamoto KW
    4. McEachin S
    5. Anderson CN

    (2020) Predicting evolutionary responses to interspecific interference in the wild

    Ecology Letters 23:221–230.

    https://doi.org/10.1111/ele.13395

    • PubMed
    • Google Scholar
27. 1. Gröning J
    2. Hochkirch A

    (2008) Reproductive interference between animal species

    The Quarterly Review of Biology 83:257–282.

    https://doi.org/10.1086/590510

    • PubMed
    • Google Scholar
28. 1. Gruson H
    2. Andraud C
    3. Daney de Marcillac W
    4. Berthier S
    5. Elias M
    6. Gomez D

    (2019) Quantitative characterization of iridescent colours in biological studies: a novel method using optical theory

    Interface Focus 9:20180049.

    https://doi.org/10.1098/rsfs.2018.0049

    • PubMed
    • Google Scholar
29. 1. Haavie J
    2. Borge T
    3. Bures S
    4. Garamszegi LZ
    5. Lampe HM
    6. Moreno J
    7. Qvarnström A
    8. Török J
    9. Saetre GP

    (2004) Flycatcher song in allopatry and sympatry--convergence, divergence and reinforcement

    Journal of Evolutionary Biology 17:227–237.

    https://doi.org/10.1111/j.1420-9101.2003.00682.x

    • PubMed
    • Google Scholar
30. Software

    1. Hartig F

    (2022) DHARMa: residual diagnostics for hierarchical (multi-level / mixed) regression models

    CRAN.

    https://github.com/florianhartig/dharma
31. 1. Houde AE

    (1987) Mate choice based upon naturally occurring color-pattern variation in a guppy population

    Evolution; International Journal of Organic Evolution 41:1–10.

    https://doi.org/10.1111/j.1558-5646.1987.tb05766.x

    • PubMed
    • Google Scholar
32. 1. Joron M
    2. Wynne IR
    3. Lamas G
    4. Mallet J

    (1999) Variable Selection and the Coexistence of Multiple mimetic forms of the Butterfly Heliconius numata

    Evolutionary Ecology 13:721–754.

    https://doi.org/10.1023/A:1010875213123

    • Google Scholar
33. 1. Keddy PA

    (1992) Assembly and response rules: two goals for predictive community ecology

    Journal of Vegetation Science 3:157–164.

    https://doi.org/10.2307/3235676

    • Google Scholar
34. 1. Kjernsmo K
    2. Whitney HM
    3. Scott-Samuel NE
    4. Hall JR
    5. Knowles H
    6. Talas L
    7. Cuthill IC

    (2020) Iridescence as Camouflage

    Current Biology 30:551–555.

    https://doi.org/10.1016/j.cub.2019.12.013

    • PubMed
    • Google Scholar
35. 1. Kneen MA
    2. Annegarn HJ

    (1996) Algorithm for fitting XRF, SEM and PIXE X-ray spectra backgrounds

    Nuclear Instruments and Methods in Physics Research Section B 109–110:209–213.

    https://doi.org/10.1016/0168-583X(95)00908-6

    • Google Scholar
36. 1. Le Roy C
    2. Amadori D
    3. Charberet S
    4. Windt J
    5. Muijres FT
    6. Llaurens V
    7. Debat V

    (2021a) Adaptive evolution of flight in Morpho butterflies

    Science 374:11581162.

    https://doi.org/10.1126/science.abh2620

    • Google Scholar
37. 1. Le Roy C
    2. Roux C
    3. Authier E
    4. Parrinello H
    5. Bastide H
    6. Debat V
    7. Llaurens V

    (2021b) Convergent morphology and divergent phenology promote the coexistence of Morpho butterfly species

    Nature Communications 12:7248.

    https://doi.org/10.1038/s41467-021-27549-1

    • PubMed
    • Google Scholar
38. 1. Li C
    2. Wang H
    3. Chen X
    4. Yao J
    5. Shi L
    6. Zhou C

    (2017) Role of visual and olfactory cues in sex recognition in butterfly Cethosia cyane cyane

    Scientific Reports 7:5033.

    https://doi.org/10.1038/s41598-017-04721-6

    • Google Scholar
39. 1. Llaurens V
    2. Joron M
    3. Théry M

    (2014) Cryptic differences in colour among Müllerian mimics: how can the visual capacities of predators and prey shape the evolution of wing colours?

    Journal of Evolutionary Biology 27:531–540.

    https://doi.org/10.1111/jeb.12317

    • PubMed
    • Google Scholar
40. 1. Llaurens V
    2. Le Poul Y
    3. Puissant A
    4. Blandin P
    5. Debat V

    (2021) Convergence in sympatry: Evolution of blue-banded wing pattern in Morpho butterflies

    Journal of Evolutionary Biology 34:284–295.

    https://doi.org/10.1111/jeb.13726

    • PubMed
    • Google Scholar
41. 1. Maia R
    2. Gruson H
    3. Endler JA
    4. White TE

    (2019) 2: New tools for the spectral and spatial analysis of colour in r

    Methods in Ecology and Evolution 10:e13174.

    https://doi.org/10.1111/2041-210X.13174

    • Google Scholar
42. 1. Maisonneuve L
    2. Smadi C
    3. Llaurens V

    (2024) Which cues are sexy? The evolution of mate preference in sympatric species reveals the contrasted effect of adaptation and reproductive interference

    Evolution Letters 8:283–294.

    https://doi.org/10.1093/evlett/qrad058

    • PubMed
    • Google Scholar
43. 1. Mallet J
    2. Gilbert LE

    (1995) Why are there so many mimicry rings? Correlations between habitat, behaviour and mimicry in Heliconius butterflies

    Biological Journal of the Linnean Society 55:159–180.

    https://doi.org/10.1111/j.1095-8312.1995.tb01057.x

    • Google Scholar
44. 1. Mallet J

    (1999) Causes and consequences of a lack of coevolution in Müllerian mimicry

    Evolutionary Ecology 13:777–806.

    https://doi.org/10.1023/A:1011060330515

    • Google Scholar
45. 1. Marko PB

    (2005)

    An intraspecific comparative analysis of character divergence between sympatric species

    Evolution; International Journal of Organic Evolution 59:554–564.

    • PubMed
    • Google Scholar
46. 1. Martin A
    2. Reed RD

    (2010) Wingless and aristaless2 define a developmental ground plan for moth and butterfly wing pattern evolution

    Molecular Biology and Evolution 27:2864–2878.

    https://doi.org/10.1093/molbev/msq173

    • PubMed
    • Google Scholar
47. 1. Mérot C
    2. Frérot B
    3. Leppik E
    4. Joron M

    (2015) Beyond magic traits: Multimodal mating cues in Heliconius butterflies

    Evolution; International Journal of Organic Evolution 69:2891–2904.

    https://doi.org/10.1111/evo.12789

    • PubMed
    • Google Scholar
48. 1. Merrill RM
    2. Wallbank RWR
    3. Bull V
    4. Salazar PCA
    5. Mallet J
    6. Stevens M
    7. Jiggins CD

    (2012) Disruptive ecological selection on a mating cue

    Proceedings of the Royal Society B 279:4907–4913.

    https://doi.org/10.1098/rspb.2012.1968

    • Google Scholar
49. 1. Merrill RM
    2. Chia A
    3. Nadeau NJ

    (2014) Divergent warning patterns contribute to assortative mating between incipient Heliconius species

    Ecology and Evolution 4:911–917.

    https://doi.org/10.1002/ece3.996

    • PubMed
    • Google Scholar
50. 1. Murali G
    2. Kodandaramaiah U

    (2020) Size and unpredictable movement together affect the effectiveness of dynamic flash coloration

    Animal Behaviour 162:87–93.

    https://doi.org/10.1016/j.anbehav.2020.02.002

    • Google Scholar
51. 1. Muschick M
    2. Indermaur A
    3. Salzburger W

    (2012) Convergent evolution within an adaptive radiation of cichlid fishes

    Current Biology 22:2362–2368.

    https://doi.org/10.1016/j.cub.2012.10.048

    • PubMed
    • Google Scholar
52. 1. Myers OD
    2. Sumner SJ
    3. Li S
    4. Barnes S
    5. Du X

    (2017) One step forward for reducing false positive and false negative compound identifications from mass spectrometry metabolomics data: New algorithms for constructing extracted ion chromatograms and detecting chromatographic peaks

    Analytical Chemistry 89:8696–8703.

    https://doi.org/10.1021/acs.analchem.7b00947

    • PubMed
    • Google Scholar
53. Software

    1. Oksanen J
    2. Simpson GL
    3. Blanchet FG
    4. Kindt R
    5. Legendre P
    6. Minchin PR
    7. O’Hara RB
    8. Solymos P
    9. Stevens MHH
    10. Szoecs E
    11. Wagner H
    12. Barbour M
    13. Bedward M
    14. Bolker B
    15. Borcard D
    16. Carvalho G
    17. Chirico M
    18. Caceres M
    19. Durand S
    20. Weedon J

    (2001) Vegan: community ecology package

    CRAN.

    https://cran.r-project.org/web/packages/vegan/index.html
54. 1. Oliver JC
    2. Robertson KA
    3. Monteiro A

    (2009) Accommodating natural and sexual selection in butterfly wing pattern evolution

    Proceedings. Biological Sciences 276:2369–2375.

    https://doi.org/10.1098/rspb.2009.0182

    • PubMed
    • Google Scholar
55. 1. Ômura H
    2. Noguchi T
    3. Ohta S

    (2020) The male swallowtail butterfly, Papilio polytes, uses cuticular hydrocarbons for mate discrimination

    Animal Behaviour 170:133–145.

    https://doi.org/10.1016/j.anbehav.2020.10.016

    • Google Scholar
56. 1. Páez E
    2. Valkonen JK
    3. Willmott KR
    4. Matos-Maraví P
    5. Elias M
    6. Mappes J

    (2021) Hard to catch: experimental evidence supports evasive mimicry

    Proceedings of the Royal Society B 288:20203052.

    https://doi.org/10.1098/rspb.2020.3052

    • Google Scholar
57. 1. Pérez-Rodríguez L
    2. Jovani R
    3. Stevens M

    (2017) Shape matters: animal colour patterns as signals of individual quality

    Proceedings of the Royal Society B 284:20162446.

    https://doi.org/10.1098/rspb.2016.2446

    • Google Scholar
58. 1. Pfennig KS
    2. Pfennig DW

    (2009) Character displacement: ecological and reproductive responses to a common evolutionary problem

    The Quarterly Review of Biology 84:253–276.

    https://doi.org/10.1086/605079

    • PubMed
    • Google Scholar
59. 1. Pinheiro CEG
    2. Freitas AVL

    (2014) Some possible cases of escape mimicry in neotropical butterflies

    Neotropical Entomology 43:393–398.

    https://doi.org/10.1007/s13744-014-0240-y

    • Google Scholar
60. 1. Pinheiro CEG
    2. Freitas AVL
    3. Campos VC
    4. DeVries PJ
    5. Penz CM

    (2016) Both palatable and unpalatable butterflies use bright colors to signal difficulty of capture to predators

    Neotropical Entomology 45:107–113.

    https://doi.org/10.1007/s13744-015-0359-5

    • PubMed
    • Google Scholar
61. 1. Pirih P
    2. Ilić M
    3. Meglič A
    4. Belušič G

    (2022) Opponent processing in the retinal mosaic of nymphalid butterflies

    Philosophical Transactions of the Royal Society B 377:20210275.

    https://doi.org/10.1098/rstb.2021.0275

    • Google Scholar
62. 1. Pluskal T
    2. Castillo S
    3. Villar-Briones A
    4. Oresic M

    (2010) MZmine 2: modular framework for processing, visualizing, and analyzing mass spectrometry-based molecular profile data

    BMC Bioinformatics 11:395.

    https://doi.org/10.1186/1471-2105-11-395

    • PubMed
    • Google Scholar
63. 1. Puissant A
    2. Chotard A
    3. Condamine FL
    4. Llaurens V

    (2023) Convergence in sympatric swallowtail butterflies reveals ecological interactions as a key driver of worldwide trait diversification

    PNAS 120:e2303060120.

    https://doi.org/10.1073/pnas.2303060120

    • Google Scholar
64. 1. Reimchen TE
    2. Nosil P

    (2004) Variable predation regimes predict the evolution of sexual dimorphism in a population of threespine stickleback

    Evolution; International Journal of Organic Evolution 58:1274–1281.

    https://doi.org/10.1111/j.0014-3820.2004.tb01706.x

    • PubMed
    • Google Scholar
65. 1. Reynolds RG
    2. Fitzpatrick BM

    (2007) Assortative mating in poison-dart frogs based on an ecologically important trait

    Evolution; International Journal of Organic Evolution 61:2253–2259.

    https://doi.org/10.1111/j.1558-5646.2007.00174.x

    • PubMed
    • Google Scholar
66. 1. Ruxton GD
    2. Speed M
    3. Sherratt TN

    (2004) Evasive mimicry: when (if ever) could mimicry based on difficulty of capture evolve?

    Proceedings of the Royal Society of London. Series B 271:2135–2142.

    https://doi.org/10.1098/rspb.2004.2816

    • Google Scholar
67. 1. Schulz S
    2. Estrada C
    3. Yildizhan S
    4. Boppré M
    5. Gilbert LE

    (2008) An antiaphrodisiac in Heliconius melpomene butterflies

    Journal of Chemical Ecology 34:82–93.

    https://doi.org/10.1007/s10886-007-9393-z

    • PubMed
    • Google Scholar
68. 1. Seehausen O
    2. Terai Y
    3. Magalhaes IS
    4. Carleton KL
    5. Mrosso HDJ
    6. Miyagi R
    7. van der Sluijs I
    8. Schneider MV
    9. Maan ME
    10. Tachida H
    11. Imai H
    12. Okada N

    (2008) Speciation through sensory drive in cichlid fish

    Nature 455:620–626.

    https://doi.org/10.1038/nature07285

    • PubMed
    • Google Scholar
69. 1. Servedio MR

    (2004) The evolution of premating isolation: local adaptation and natural and sexual selection against hybrids

    Evolution; International Journal of Organic Evolution 58:913–924.

    https://doi.org/10.1111/j.0014-3820.2004.tb00425.x

    • PubMed
    • Google Scholar
70. 1. Servedio MR
    2. Boughman JW

    (2017) The role of sexual selection in local adaptation and speciation

    Annual Review of Ecology, Evolution, and Systematics 48:85–109.

    https://doi.org/10.1146/annurev-ecolsys-110316-022905

    • Google Scholar
71. 1. Siddique RH
    2. Diewald S
    3. Leuthold J
    4. Hölscher H

    (2013) Theoretical and experimental analysis of the structural pattern responsible for the iridescence of Morpho butterflies

    Optics Express 21:14351–14361.

    https://doi.org/10.1364/OE.21.014351

    • PubMed
    • Google Scholar
72. 1. Simpson RK
    2. McGraw KJ

    (2019) Interspecific covariation in courtship displays, iridescent plumage, solar orientation, and their interactions in hummingbirds

    The American Naturalist 194:441–454.

    https://doi.org/10.1086/704774

    • PubMed
    • Google Scholar
73. 1. Soni SP
    2. Apte V
    3. Joshi P
    4. Cyriac VP

    (2025) Barking up the wrong frog: global prevalence of misdirected amplexus in anuran amphibians

    Biological Journal of the Linnean Society 145:blae062.

    https://doi.org/10.1093/biolinnean/blae062

    • Google Scholar
74. 1. Stoffel MA
    2. Nakagawa S
    3. Schielzeth H

    (2017) rptR: repeatability estimation and variance decomposition by generalized linear mixed‐effects models

    Methods in Ecology and Evolution 8:1639–1644.

    https://doi.org/10.1111/2041-210X.12797

    • Google Scholar
75. 1. Stuckert AMM
    2. Venegas PJ
    3. Summers K

    (2014) Experimental evidence for predator learning and Müllerian mimicry in Peruvian poison frogs (Ranitomeya, Dendrobatidae)

    Evolutionary Ecology 28:413–426.

    https://doi.org/10.1007/s10682-013-9685-4

    • Google Scholar
76. 1. terHorst CP
    2. Zee PC
    3. Heath KD
    4. Miller TE
    5. Pastore AI
    6. Patel S
    7. Schreiber SJ
    8. Wade MJ
    9. Walsh MR

    (2018) Evolution in a community context: Trait responses to multiple species interactions

    The American Naturalist 191:368–380.

    https://doi.org/10.1086/695835

    • Google Scholar
77. 1. Thomas DHN
    2. Kjernsmo K
    3. Scott-Samuel NE
    4. Whitney HM
    5. Cuthill IC

    (2023) Interactions between color and gloss in iridescent camouflage

    Behavioral Ecology 34:751–758.

    https://doi.org/10.1093/beheco/arad050

    • PubMed
    • Google Scholar
78. 1. van Doorn GS
    2. Edelaar P
    3. Weissing FJ

    (2009) On the origin of species by natural and sexual selection

    Science 326:1704–1707.

    https://doi.org/10.1126/science.1181661

    • PubMed
    • Google Scholar
79. 1. Vieira‐Silva A
    2. Evora GB
    3. Freitas AVL
    4. Oliveira PS

    (2024) The relevance of flash coloration against avian predation in a Morpho butterfly: A field experiment in a tropical rainforest

    Ethology 130:e13517.

    https://doi.org/10.1111/eth.13517

    • Google Scholar
80. 1. Vorobyev M
    2. Brandt R
    3. Peitsch D
    4. Laughlin SB
    5. Menzel R

    (2001) Colour thresholds and receptor noise: behaviour and physiology compared

    Vision Research 41:639–653.

    https://doi.org/10.1016/s0042-6989(00)00288-1

    • PubMed
    • Google Scholar
81. 1. Waitt C
    2. Little AC
    3. Wolfensohn S
    4. Honess P
    5. Brown AP
    6. Buchanan-Smith HM
    7. Perrett DI

    (2003) Evidence from rhesus macaques suggests that male coloration plays a role in female primate mate choice

    Proceedings. Biological Sciences 270 Suppl 2:S144–S146.

    https://doi.org/10.1098/rsbl.2003.0065

    • PubMed
    • Google Scholar
82. 1. White TE
    2. Zeil J
    3. Kemp DJ

    (2015) Signal design and courtship presentation coincide for highly biased delivery of an iridescent butterfly mating signal

    Evolution; International Journal of Organic Evolution 69:14–25.

    https://doi.org/10.1111/evo.12551

    • PubMed
    • Google Scholar
83. 1. Young AM

    (1971) Wing coloration and reflectance in Morpho butterflies as related to reproductive behavior and escape from avian predators

    Oecologia 7:209–222.

    https://doi.org/10.1007/BF00345212

    • PubMed
    • Google Scholar
84. 1. Young AM

    (1973) Studies on comparative ecology and ethology in adult populations of several species of Morpho butterflies (Lepidoptera: Morphidae)

    Studies on Neotropical Fauna 8:17–50.

    https://doi.org/10.1080/01650527309360452

    • Google Scholar
85. 1. Zhou W
    2. Yu L
    3. Kwek BZW
    4. Jin G
    5. Zeng H
    6. Li D

    (2021) Sexual selection on jumping spider color pattern: investigation with a new quantitative approach

    Behavioral Ecology 32:695–706.

    https://doi.org/10.1093/beheco/arab008

    • Google Scholar

Author details

1. Joséphine Ledamoisel

   1. Collège de France, Centre Interdisciplinaire de Recherche en Biologie, Paris, France
   2. Muséum national d'Histoire naturelle, Institut de Systématique, Evolution et Biodiversité (ISYEB UMR 7205), Paris, France
   3. Smithsonian Tropical Research Institute, Panama, Panama
   4. Muséum national d'Histoire naturelle, Centre de Recherche sur la Conservation, Paris, France

   Contribution

   Conceptualization, Resources, Formal analysis, Investigation, Visualization, Methodology, Writing - original draft, Writing – review and editing

   For correspondence

   josephine.ledamoisel@outlook.fr

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-7885-3300

2. Bruno Buatois

   Université de Montpellier, Centre d'Ecologie Fonctionnelle et Evolutive, Montpellier, France

   Contribution

   Methodology, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-3574-8425

3. Rémi Mauxion

   Smithsonian Tropical Research Institute, Panama, Panama

   Contribution

   Resources, Methodology, Writing – review and editing

   Competing interests

   No competing interests declared

4. Christine Andraud

   Muséum national d'Histoire naturelle, Centre de Recherche sur la Conservation, Paris, France

   Contribution

   Resources, Methodology, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-3112-9363

5. Melanie McClure

   Université de Guyane, Laboratoire Écologie, Évolution, Interactions des Systèmes Amazoniens (LEEISA), Cayenne, France

   Contribution

   Resources, Formal analysis, Methodology, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-3590-4002

6. Vincent Debat

   1. Collège de France, Centre Interdisciplinaire de Recherche en Biologie, Paris, France
   2. Muséum national d'Histoire naturelle, Institut de Systématique, Evolution et Biodiversité (ISYEB UMR 7205), Paris, France

   Contribution

   Conceptualization, Supervision, Validation, Methodology, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-0040-1181

7. Violaine Llaurens

   1. Collège de France, Centre Interdisciplinaire de Recherche en Biologie, Paris, France
   2. Muséum national d'Histoire naturelle, Institut de Systématique, Evolution et Biodiversité (ISYEB UMR 7205), Paris, France

   Contribution

   Conceptualization, Supervision, Funding acquisition, Validation, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-1962-7391

• 395

  views

• 21

  downloads

• 1

  citation

Views, downloads and citations are aggregated across all versions of this paper published by eLife.

Citations by DOI

• 1

  citation for Reviewed Preprint v1 https://doi.org/10.7554/eLife.106098.1