Here is a list of over 600 publications about Heliconius and related butterfly genera.  No guarantees regarding accuracy, and the list was compiled early in 2008 so is already out of date.

• [2008,article] bibtex
  M. Kronforst, "Gene flow persists millions of years after speciation in Heliconius butterflies," BMC Evolutionary Biology, vol. 8, p. 98, 2008.
  @article{ Author = {Kronforst, M.},
  Title = {Gene flow persists millions of years after speciation in Heliconius butterflies},
  Journal = {BMC Evolutionary Biology},
  Volume = {8},
  Pages = {98},
  Year = {2008}
}
• [2008,article] bibtex
  S. Schulz, C. Estrada, S. Yildizhan, M. Boppre, and L. E. Gilbert, "An antiaphrodisiac in Heliconius melpomene butterflies," Journal of Chemical Ecology, vol. 34, iss. 1, pp. 82-93, 2008.
  @article{ Author = {Schulz, S. and Estrada, C. and Yildizhan, S. and Boppre, M. and Gilbert, L. E.},
  Title = {An antiaphrodisiac in Heliconius melpomene butterflies},
  Journal = {Journal of Chemical Ecology},
  Volume = {34},
  Number = {1},
  Pages = {82-93},
  Note = {256ZM Times Cited:0 Cited References Count:52},
  Abstract = {Gilbert (1976) suggested that male-contributed odors of mated females of Heliconius erato could enforce monogamy. We investigated the pheromone system of a relative, Heliconius melpomene, using chemical analysis, behavioral experiments, and feeding experiments with labeled biosynthetic pheromone precursors. The abdominal scent glands of males contained a complex odor bouquet, consisting of the volatile compound (E)-beta-ocimene together with some trace components and a less volatile matrix made up predominately of esters of common C-16- and C-18-fatty acids with the alcohols ethanol, 2-propanol, 1-butanol, isobutanol, 1-hexanol, and (Z)-3-hexenol. This bouquet is formed during the first days after eclosion, and transferred during copulation to the females. Virgin female scent glands do not contain these compounds. The transfer of ocimene and the esters was shown by analysis of butterflies of both sexes before and after copulation. Additional proof was C-13-labeled ocimene and transferred it to females during copulation. Behavioral tests with ocimene applied to unmated females showed its repellency to males. The esters did not show such activity, but they moderated the evaporation rate of ocimene. Our investigation showed that beta-ocimene is an antiaphrodisiac pheromone of H. melpomene.},
  Keywords = {pheromones heliconius antiaphrodisiacs sperm competition ocimene fatty acid esters labeled pheromone pheromone biosynthesis cyanogenic glucosides floral scents lepidoptera biosynthesis evolution females mimicry glands lotaustralin nymphalidae},
  Year = {2008}
}
• [2008,article] bibtex
  R. Papa, C. M. Morrison, J. R. Walters, B. A. Counterman, R. Chen, G. Halder, L. Roberts, D. D. Kapan, C. D. Jiggins, R. D. Reed, and W. O. McMillan, "Highly conserved gene order and numerous novel repetitive elements in genomic regions linked to wing pattern variation in Heliconius butterflies," BMC Genomics, vol. in review, 2008.
  @article{ Author = {Papa, R. and Morrison, C. M. and Walters, J. R. and Counterman, B. A. and Chen, R. and Halder, G. and Roberts, L. and Kapan, D. D. and Jiggins, C. D. and Reed, R. D. and McMillan, W. O.},
  Title = {Highly conserved gene order and numerous novel repetitive elements in genomic regions linked to wing pattern variation in Heliconius butterflies},
  Journal = {BMC Genomics},
  Volume = {in review},
  Year = {2008}
}
• [2008,article] bibtex
  R. D. Reed, W. O. McMillan, and L. M. Nagy, "Gene expression underlying adaptive variation in Heliconius wing patterns: non-modular regulation of overlapping cinnabar and vermilion prepatterns," Proceedings of the Royal Society B-Biological Sciences, vol. 275, iss. 1630, pp. 37-45, 2008.
  @article{ Author = {Reed, R. D. and McMillan, W. O. and Nagy, L. M.},
  Title = {Gene expression underlying adaptive variation in Heliconius wing patterns: non-modular regulation of overlapping cinnabar and vermilion prepatterns},
  Journal = {Proceedings of the Royal Society B-Biological Sciences},
  Volume = {275},
  Number = {1630},
  Pages = {37-45},
  Note = {242IC Times Cited:0 Cited References Count:28},
  Abstract = {Geographical variation in the mimetic wing patterns of the butterfly Heliconius erato is a textbook example of adaptive polymorphism; however, little is known about how this variation is controlled developmentally. Using microarrays and qPCR, we identified and compared expression of candidate genes potentially involved with a red/yellow forewing band polymorphism in H. erato. We found that transcripts encoding the pigment synthesis enzymes cinnabar and vermilion showed pattern- and polymorphism-related expression patterns, respectively. cinnabar expression was associated with the forewing band regardless of pigment colour, providing the first gene expression pattern known to be correlated with a major Heliconius colour pattern. In contrast, vermilion expression changed spatially over time in red-banded butterflies, but was not expressed at detectable levels in yellow-banded butterflies, suggesting that regulation of this gene may be involved with the red/yellow polymorphism. Furthermore, we found that the yellow pigment, 3-hydroxykynurenine, is incorporated into wing scales from the haemolymph rather than being synthesized in situ. We propose that some aspects of Heliconius colour patterns are determined by spatio-temporal overlap of pigment gene transcription prepatterns and speculate that evolutionary changes in vermilion regulation may in part underlie an adaptive colour pattern polymorphism.},
  Keywords = {heliconius evo-devo cinnabar vermillion pigmentation colour patterns mullerian mimicry warning color linkage map butterflies erato evolution scales},
  Year = {2008}
}
• [2008,article] bibtex
  M. J. Blum, "Ecological and genetic associations across a Heliconius hybrid zone," Journal of Evolutionary Biology, vol. 21, iss. 1, pp. 330-341, 2008.
  @article{ Author = {Blum, M. J.},
  Title = {Ecological and genetic associations across a Heliconius hybrid zone},
  Journal = {Journal of Evolutionary Biology},
  Volume = {21},
  Number = {1},
  Pages = {330-341},
  Note = {242ZV Times Cited:0 Cited References Count:57},
  Abstract = {Differences in habitat use can bridge early and late stages of speciation by initiating assortative mating. Heliconius colour pattern races might select habitats over which each pattern confers a relative fitness advantage because signal efficacy of wing patterns can vary by environment. Thus habitat preferences could serve to promote the evolution of mimetic colour patterns for mate choice. Here I compare colour pattern genotype and phenotype frequencies to environmental variation across the H. erato hydara x H. erato erato hybrid zone in French Guiana to determine whether races exhibit habitat preferences. I found that genotype and phenotype frequencies correspond to differences in land cover moreso than to other environmental factors. Temporal shifts in colour pattern genotypes, phenotypes and land cover also were associated at individual sample sites, which further suggests that H. erato races differ in habitat use and that habitat preferences may promote speciation among Heliconius butterflies.},
  Keywords = {colour pattern evolution habitat preferences hybrid zone movement speciation passion-vine butterflies warning-color habitat preference natural-selection shifting balance mate preference mimicry speciation evolution erato},
  Year = {2008}
}
• [2008,article] bibtex
  C. Salazar, C. D. Jiggins, J. Taylor, M. Kronforst, and M. Linares, "Hybrid speciation and the genealogical history of Heliconius heurippa," BMC Evolutionary Biology, vol. Accepted subject to revision, 2008.
  @article{ Author = {Salazar, C. and Jiggins, C. D. and Taylor, J. and Kronforst, M. and Linares, M.},
  Title = {Hybrid speciation and the genealogical history of Heliconius heurippa},
  Journal = {BMC Evolutionary Biology},
  Volume = {Accepted subject to revision},
  Year = {2008}
}
• [2008,article] bibtex
  M. P. Sison-Mangus, A. D. Briscoe, G. Zaccardi, H. Knuttel, and A. Kelber, "The lycaenid butterfly Polyommatus icarus uses a duplicated blue opsin to see green," Journal of Experimental Biology, vol. 211, iss. 3, pp. 361-369, 2008.
  @article{ Author = {Sison-Mangus, M. P. and Briscoe, A. D. and Zaccardi, G. and Knuttel, H. and Kelber, A.},
  Title = {The lycaenid butterfly Polyommatus icarus uses a duplicated blue opsin to see green},
  Journal = {Journal of Experimental Biology},
  Volume = {211},
  Number = {3},
  Pages = {361-369},
  Note = {263CY Times Cited:0 Cited References Count:48},
  Abstract = {The functional significance of gene duplication is rarely addressed at the level of animal behavior. Butterflies are excellent models in this regard because they can be trained and the use of their opsin-based visual pigments in color vision can be assessed. In the present study, we demonstrate that the lycaenid Polyommatus icarus uses its duplicate blue (B2) opsin, BRh2, in conjunction with its long-wavelength (LW) opsin, LWRh, to see color in the green part of the light spectrum extending up to 560 nm. This is in contrast to butterflies in the genus Papilio, which use duplicate LW opsins to discriminate colors in the long-wavelength range. We also found that P. icarus has a heterogeneously expressed red filtering pigment and red-reflecting ommatidia in the ventral eye region. In behavioural tests, the butterflies could not discriminate colors in the red range (570-640 nm). This finding is significant because we have previously found that the nymphalid butterfly Heliconius erato has filter-pigment mediated color vision in the long wavelength range. Our results suggest that lateral filtering pigments may not always influence color vision in insects.},
  Keywords = {lycaenid color vision visual pigment filter pigment butterfly opsin pieris-rapae-crucivora color-vision visual pigment papilio-xuthus screening pigments messenger-rnas class insecta eye red ultraviolet},
  Year = {2008}
}
• [2008,article] bibtex
  P. Beldade, W. O. McMillan, and A. Papanicolaou, "Butterfly genomics eclosing," Heredity, vol. 100, iss. 2, pp. 150-157, 2008.
  @article{ Author = {Beldade, P. and McMillan, W. O. and Papanicolaou, A.},
  Title = {Butterfly genomics eclosing},
  Journal = {Heredity},
  Volume = {100},
  Number = {2},
  Pages = {150-157},
  Note = {254KP Times Cited:0 Cited References Count:67},
  Abstract = {Technological and conceptual advances of the last decade have led to an explosion of genomic data and the emergence of new research avenues. Evolutionary and ecological functional genomics, with its focus on the genes that affect ecological success and adaptation in natural populations, benefits immensely from a phylogenetically widespread sampling of biological patterns and processes. Among those organisms outside established model systems, butterflies offer exceptional opportunities for multidisciplinary research on the processes generating and maintaining variation in ecologically relevant traits. Here we highlight research on wing color pattern variation in two groups of Nymphalid butterflies, the African species Bicyclus anynana (subfamily Satyrinae) and species of the South American genus Heliconius (subfamily Heliconiinae), which are emerging as important systems for studying the nature and origins of functional diversity. Growing genomic resources including genomic and cDNA libraries, dense genetic maps, high-density gene arrays, and genetic transformation techniques are extending current gene mapping and expression profiling analysis and enabling the next generation of research questions linking genes, development, form, and fitness. Efforts to develop such resources in Bicyclus and Heliconius underscore the general challenges facing the larger research community and highlight the need for a community-wide effort to extend ongoing functional genomic research on butterflies.},
  Keywords = {evolutionary and ecological functional genomics butterfly wing patterns bicyclus heliconius est linkage maps bicyclus-anynana butterflies evo-devo model bombyx-mori wing patterns linkage map heliconius butterflies mullerian mimicry natural-selection developmental constraints color patterns},
  Year = {2008}
}
• [2008,misc] bibtex
  M. G. Lamas, BIBLIOGRAPHY OF BUTTERFLIES. An Annotated Bibliography of the Neotropical Butterflies and Skippers (Lepidoptera: Papilionoidea and Hesperioidea), 2008.
  @misc{ Author = {Lamas, M. G.},
  Title = {BIBLIOGRAPHY OF BUTTERFLIES. An Annotated Bibliography of the Neotropical Butterflies and Skippers (Lepidoptera: Papilionoidea and Hesperioidea)},
  Note = {http://www.andeanbutterflies.org/country/peru/lamas_2004_bibliography_butterflies.doc.},
  Year = {2008}
}
• [2008,article] bibtex
  S. Baxter, R. Papa, N. Chamberlain, S. Humphray, R. ffrench-Constant, W. O. McMillan, and C. D. Jiggins, "Parallel evolution in the genetic basis of Mullerian mimicry in Heliconius butterflies," Genetics, vol. In press, 2008.
  @article{ Author = {Baxter, S. and Papa, R. and Chamberlain, N. and Humphray, S. and ffrench-Constant, R. and McMillan, W. O. and Jiggins, C. D.},
  Title = {Parallel evolution in the genetic basis of Mullerian mimicry in Heliconius butterflies},
  Journal = {Genetics},
  Volume = {In press},
  Year = {2008}
}
• [2008,article] bibtex
  R. Clark, S. M. Brown, S. C. Collins, C. D. Jiggins, D. G. Heckel, and A. P. Vogler, "Colour pattern specification in the Mocker swallowtail Papilio dardanus: the transcription factor invected is a candidate for the mimicry locus H," Proceedings of the Royal Society B-Biological Sciences, vol. 275, iss. 1639, pp. 1181-1188, 2008.
  @article{ Author = {Clark, R. and Brown, S. M. and Collins, S. C. and Jiggins, C. D. and Heckel, D. G. and Vogler, A. P.},
  Title = {Colour pattern specification in the Mocker swallowtail Papilio dardanus: the transcription factor invected is a candidate for the mimicry locus H},
  Journal = {Proceedings of the Royal Society B-Biological Sciences},
  Volume = {275},
  Number = {1639},
  Pages = {1181-1188},
  Note = {280YY Times Cited:0 Cited References Count:33},
  Abstract = {The swallowtail butterfly, Papilio dardanus, is an iconic example of a polymorphic Batesian mimic. The expression of various female-limited colour forms is thought to be controlled by a single autosomal locus, termed H, whose function in determining the wing pattern remains elusive. As a step towards the physical mapping of H, we established a set of 272 polymorphic amplified fragment length polymorphism (AFLP) markers (EcoRI-MseI). Segregation patterns in a 'female-informative' brood (exploiting the absence of crossing over in female Lepidoptera) mapped these AFLPs to 30 linkage groups (putative chromosomes). The difference between the hippocoon and cenea female forms segregating in this family resides on a single one of these linkage groups, defined by 14 AFLPs. In a 'male-informative' cross (markers segregating within a linkage group), a pair of AFLPs co-segregated closely with the two female forms, except in four recombinants out of 19 female offspring. Linkage with these AFLP markers using four further female-informative families demonstrated that the genetic factor determining other morphs (poultoni, lamborni and trimeni) also maps to this same linkage group. The candidate gene invected, obtained in a screen for co-segregation of developmental genes with the colour forms, resides in a 13.9 cM interval flanked by the two AFLP markers. In the male-informative family invected co-segregated perfectly with the hippocoon/cenea factor, despite the four crossovers with the AFLPs. These findings make invected, and possibly its closely linked paralogue engrailed, strong candidates for H. This is supported by their known role in eyespot specification in nymphalid butterfly wings.},
  Keywords = {evolutionary genetics mimicry phenotype-genotype association engrailed candidate genes amplified fragment length polymorphism linkage map heliconius-melpomene evolution resistance genetics genes brown identification synteny erato},
  Year = {2008}
}
• [2008,article] bibtex
  M. R. Kronforst and L. E. Gilbert, "The population genetics of mimetic diversity in Heliconius butterflies," Proceedings of the Royal Society B-Biological Sciences, vol. 275, iss. 1634, pp. 493-500, 2008.
  @article{ Author = {Kronforst, M. R. and Gilbert, L. E.},
  Title = {The population genetics of mimetic diversity in Heliconius butterflies},
  Journal = {Proceedings of the Royal Society B-Biological Sciences},
  Volume = {275},
  Number = {1634},
  Pages = {493-500},
  Note = {256YC Times Cited:0 Cited References Count:60},
  Abstract = {Theory predicts strong stabilizing selection on warning patterns within species and convergent evolution among species in Mullerian mimicry systems yet Heliconius butterflies exhibit extreme wing pattern diversity. One potential explanation for the evolution of this diversity is that genetic drift occasionally allows novel warning patterns to reach the frequency threshold at which they gain protection. This idea is controversial, however, because Heliconius butterflies are unlikely to experience pronounced population subdivision and local genetic drift. To examine the fine-scale population genetic structure of Heliconius butterflies we genotyped 316 individuals from eight Costa Rican Heliconius species with 1428 AFLP markers. Six species exhibited evidence of population subdivision and/or isolation by distance indicating genetic differentiation among populations. Across species, variation in the extent of local genetic drift correlated with the roles different species have played in generating pattern diversity: species that originally generated the diversity of warning patterns exhibited striking population subdivision while species that later radiated onto these patterns had intermediate levels of genetic diversity and less genetic differentiation among populations. These data reveal that Heliconius butterflies possess the coarse population genetic structure necessary for local populations to experience pronounced genetic drift which, in turn, could explain the origin of mimetic diversity.},
  Keywords = {genetic differentiation genetic drift mullerian mimicry population structure mitochondrial-DNA sequences shifting balance theory home-range behavior mullerian mimicry race formation warning color hybrid zones evolution nymphalidae lepidoptera},
  Year = {2008}
}
• [2008,article] bibtex
  N. O. Mega and A. M. de Araujo, "Do caterpillars of Dryas iulia alcionea (Lepidoptera, Nymphalidae) show evidence of adaptive behaviour to avoid predation by ants?," Journal of Natural History, vol. 42, iss. 1-2, pp. 129-137, 2008.
  @article{ Author = {Mega, N. O. and de Araujo, A. M.},
  Title = {Do caterpillars of Dryas iulia alcionea (Lepidoptera, Nymphalidae) show evidence of adaptive behaviour to avoid predation by ants?},
  Journal = {Journal of Natural History},
  Volume = {42},
  Number = {1-2},
  Pages = {129-137},
  Note = {269NS Times Cited:0 Cited References Count:36},
  Abstract = {Plants bearing extrafloral nectaries can attract wasps, bugs, and mainly ants. Ants can defend plants against herbivores by treating the plant as an extension of their own territory. We evaluated the efficiency of resting sites built by Dryas iulia alcionea larvae as protection against ant predation on their host plant Passiflora suberosa. The results showed that ants on P. suberosa have significant influence on the survival of D. iulia caterpillars. The use and construction of resting sites was recorded through the fourth larval stadium. A strong inverse relationship between the number of resting sites constructed per larva and the developmental stage of the larvae was observed. Additional results suggest that the efficacy of resting sites against cursorial predators decreases with larval development. The behaviour of constructing resting sites probably evolved by natural selection. It is likely that this behaviour was important for the ancestral lineage that gave origin to Heliconiinae genera Dryas, Dryadula and Phylaethria.},
  Keywords = {anti-predator adaptation caterpillar resting site heliconiinae heliconius butterflies phylogeny nectaries responses larvae plant},
  Year = {2008}
}
• [2007,article] bibtex
  K. K. Dasmahapatra, A. Silva-Vasquez, J. W. Chung, and J. Mallet, "Genetic analysis of a wild-caught hybrid between non-sister Heliconius butterfly species," Biology Letters, vol. 3, iss. 6, pp. 660-663, 2007.
  @article{ Author = {Dasmahapatra, K. K. and Silva-Vasquez, A. and Chung, J. W. and Mallet, J.},
  Title = {Genetic analysis of a wild-caught hybrid between non-sister Heliconius butterfly species},
  Journal = {Biology Letters},
  Volume = {3},
  Number = {6},
  Pages = {660-663},
  Note = {230VE Times Cited:0 Cited References Count:18},
  Abstract = {Interspecific hybridization occurs regularly in wild Heliconius butterflies, although hybrid individuals are usually very rare. However, hybridization generally occurs only between the most closely related species. We report a rare naturally occurring hybrid between non-sister species and carry out the first genetic analysis of such distant hybridization. Mitochondrial and nuclear genes indicate that the specimen is an F-1 hybrid between a female Heliconius ethilla and a male Heliconius melpomene, originating from a group of 13 species estimated to have diverged over 2.5Myr ago. The presence of such distant natural hybrids, together with evidence for backcrossing, suggests that gene flow across species boundaries can take place long after speciation. Adaptive genes such as those involved in wing coloration could thus be widely shared among members of this highly mimetic genus.},
  Keywords = {heliconius ethilla heliconius melpomene hybridization hybridization zone selection evolution},
  Year = {2007}
}
• [2007,article] bibtex
  E. Ihalainen, L. Lindstrom, and J. Mappes, "Investigating Mullerian mimicry: predator learning and variation in prey defences," Journal of Evolutionary Biology, vol. 20, iss. 2, pp. 780-791, 2007.
  @article{ Author = {Ihalainen, E. and Lindstrom, L. and Mappes, J.},
  Title = {Investigating Mullerian mimicry: predator learning and variation in prey defences},
  Journal = {Journal of Evolutionary Biology},
  Volume = {20},
  Number = {2},
  Pages = {780-791},
  Note = {136SL Times Cited:2 Cited References Count:66},
  Abstract = {Inexperienced predators are assumed to select for similarity of warning signals in aposematic species (Mullerian mimicry) when learning to avoid them. Recent theoretical work predicts that if co-mimic species have unequal defences, predators attack them according to their average unpalatability and mimicry may not be beneficial for the better defended co-mimic. In this study, we tested in a laboratory environment whether a uniform warning signal is superior to a variable one in promoting predator learning, and simultaneously whether co-mimics are preyed upon according to their average unpalatability. There was an interaction of signal variation and unpalatability but inexperienced birds did not select for signal similarity in artificial prey; when the prey was moderately defended a variable signal was even learnt faster than a uniform one. Due to slow avoidance learning, moderately defended prey had higher mortality than highly defended prey (although this was not straightforward), but mixing high and moderate unpalatability did not increase predation compared with high unpalatability. This does not support the view that predators are sensitive to varying unpalatability. The results suggest that inexperienced predators may neither strongly select for accurate Mullerian mimicry nor affect the benefits of mimicry when the co-mimics are unequally defended.},
  Keywords = {aposematism memory palatability spectrum quasi-batesian mimicry coral-snake pattern warning coloration muellerian mimicry heliconius butterflies palatability spectrum dietary conservatism natural-selection batesian mimicry shifting balance evolution},
  Year = {2007}
}
• [2007,article] bibtex
  R. B. Srygley, "Evolution of the wave: aerodynamic and aposematic functions of butterfly wing motion," Proceedings of the Royal Society B-Biological Sciences, vol. 274, iss. 1612, pp. 913-917, 2007.
  @article{ Author = {Srygley, R. B.},
  Title = {Evolution of the wave: aerodynamic and aposematic functions of butterfly wing motion},
  Journal = {Proceedings of the Royal Society B-Biological Sciences},
  Volume = {274},
  Number = {1612},
  Pages = {913-917},
  Note = {138PV Times Cited:0 Cited References Count:30},
  Abstract = {Many unpalatable butterfly species use coloration to signal their distastefulness to birds, but motion cues may also be crucial to ward off predatory attacks. In previous research, captive passion-vine butterflies Heliconius mimetic in colour pattern were also mimetic in motion. Here, I investigate whether wing motion changes with the flight demands of different behaviours. If birds select for wing motion as a warning signal, aposematic butterflies should maintain wing motion independently of behavioural context. Members of one mimicry group ( Heliconius cydno and Heliconius sapho) beat their wings more slowly and their wing strokes were more asymmetric than their sister-species ( Heliconius melpomene and Heliconius erato, respectively), which were members of another mimicry group having a quick and steady wing motion. Within mimicry groups, wing beat frequency declined as its role in generating lift also declined in different behavioural contexts. In contrast, asymmetry of the stroke was not associated with wing beat frequency or behavioural context - strong indication that birds process and store the Fourier motion energy of butterfly wings. Although direct evidence that birds respond to subtle differences in butterfly wing motion is lacking, birds appear to generalize a motion pattern as much as they encounter members of a mimicry group in different behavioural contexts.},
  Keywords = {locomotor mimicry insect flight mimetic behaviour mutualism bird vision mullerian mimicry heliconius butterflies warning signals visual neurons mimicry perception flight morphology models costs},
  Year = {2007}
}
• [2007,article] bibtex
  M. M. Hay-Roe and J. Nation, "Spectrum of cyanide toxicity and allocation in Heliconius erato and Passiflora host plants," Journal of Chemical Ecology, vol. 33, iss. 2, pp. 319-329, 2007.
  @article{ Author = {Hay-Roe, M. M. and Nation, J.},
  Title = {Spectrum of cyanide toxicity and allocation in Heliconius erato and Passiflora host plants},
  Journal = {Journal of Chemical Ecology},
  Volume = {33},
  Number = {2},
  Pages = {319-329},
  Note = {127AQ Times Cited:1 Cited References Count:33},
  Abstract = {The larvae of three races of Heliconius erato were fed various species of Passiflora containing varying levels of cyanoglucosides. The mortality rate of larvae and pupae rose when larvae were fed species of Passiflora capable of releasing larger quantities of cyanide. When larvae were fed species of Passiflora with these properties, the resulting adult butterflies also released higher levels of cyanide. This may serve as a defense mechanism. The compounds responsible for the release of cyanide were not evenly distributed throughout the adult butterfly's body. The thorax contained the highest concentration of cyanogenic substances, followed by the head, wings, and abdomen. The younger tissues of Passiflora plants had higher levels of cyanide-releasing compounds than stems and mature leaves. Cyanogenic glycoside distribution within the plants is consistent with optimal allocation theory. The levels of cyanide-releasing substances in plants varied depending on the season.},
  Keywords = {heliconius erato cyrbia heliconius erato favorinus heliconius erato demophoon passiflora cyanogenic glycosides beta-glucosidase cyanide ecuador peru panama neotropical butterflies cyanogenic glucosides mullerian mimicry palatability biosynthesis consequences lotaustralin lepidoptera coloration glycosides},
  Year = {2007}
}
• [2007,article] bibtex
  M. Elias, R. I. Hill, K. R. Willmott, K. K. Dasmahapatra, A. V. Z. Brower, J. Malllet, and C. D. Jiggins, "Limited performance of DNA barcoding in a diverse community of tropical butterflies," Proceedings of the Royal Society B-Biological Sciences, vol. 274, iss. 1627, pp. 2881-2889, 2007.
  @article{ Author = {Elias, M. and Hill, R. I. and Willmott, K. R. and Dasmahapatra, K. K. and Brower, A. V. Z. and Malllet, J. and Jiggins, C. D.},
  Title = {Limited performance of DNA barcoding in a diverse community of tropical butterflies},
  Journal = {Proceedings of the Royal Society B-Biological Sciences},
  Volume = {274},
  Number = {1627},
  Pages = {2881-2889},
  Note = {225YU Times Cited:0 Cited References Count:70},
  Abstract = {DNA 'barcoding' relies on a short fragment of mitochondrial DNA to infer identification of specimens. The method depends on genetic diversity being markedly lower within than between species. Closely related species are most likely to share genetic variation in communities where speciation rates are rapid and effective population sizes are large, such that coalescence times are long. We assessed the applicability of DNA barcoding (here the 5' half of the cytochrome c oxidase 1) to a diverse community of butterflies from the upper Amazon, using a group with a well-established morphological taxonomy to serve as a reference. Only 77% of species could be accurately identified using the barcode data, a figure that dropped to 68% in species represented in the analyses by more than one geographical race and at least one congener. The use of additional mitochondrial sequence data hardly improved species identification, while a fragment of a nuclear gene resolved issues in some of the problematic species. We acknowledge the utility of barcodes when morphological characters are ambiguous or unknown, but we also recommend the addition of nuclear sequence data, and caution that species-level identification rates might be lower in the most diverse habitats of our planet.},
  Keywords = {DNA barcoding amazon biodiversity lepidoptera mimicry parasitoid flies diptera mitochondrial-DNA molecular systematics phylogenetic-relationships heliconius butterflies species delimitation astraptes-fulgerator gene trees lepidoptera identification},
  Year = {2007}
}
• [2007,article] bibtex
  S. H. Eberhard, N. Hrassnigg, K. Crailsheim, and H. W. Krenn, "Evidence of protease in the saliva of the butterfly Heliconius melpomene (L.) (Nymphalidae, Lepidoptera)," Journal of Insect Physiology, vol. 53, iss. 2, pp. 126-131, 2007.
  @article{ Author = {Eberhard, S. H. and Hrassnigg, N. and Crailsheim, K. and Krenn, H. W.},
  Title = {Evidence of protease in the saliva of the butterfly Heliconius melpomene (L.) (Nymphalidae, Lepidoptera)},
  Journal = {Journal of Insect Physiology},
  Volume = {53},
  Number = {2},
  Pages = {126-131},
  Note = {144ST Times Cited:0 Cited References Count:25},
  Abstract = {Butterflies of the genus Heliconius are well known for their peculiar habits of utilizing pollen as a source of amino acids. Saliva plays a major role in the process of extracting amino acids and proteins from the pollen grains. In this investigation, we obtained samples of saliva from adult Heliconius melpomene by placing pumpkin pollen or fine glass-beads on the proboscis, which stimulates the butterflies to release saliva. Proteolytic activity was determined in the saliva by an insoluble protein-dye that turns blue when cleaved by proteases. Its extinction value was measured with a spectrophotometer at 595 nm. Both the saliva sampled with pollen and the saliva obtained from inert glass-beads exhibit proteolytic activity demonstrating that the saliva contains proteases.. The proteolytic activity of the pollen/saliva samples was higher than that of the glass-bead/saliva samples, which we attribute to the stimulating effects of pollen, such as taste, smell, and texture, and not to proteases which might have been liberated from the pollen. This is indicated by the fact that pollen samples without saliva showed only a negligible indication for proteolytic activity. In general, females exhibit higher proteolytic activities than males, presumably due to their greater amino acid investment in reproduction. We present here first evidence for the existence of proteases in the saliva of a butterfly species and suggest that these enzymes are crucial for the use of amino acids and proteins from pollen in Heliconius butterflies. (c) 2006 Elsevier Ltd. All rights reserved.},
  Keywords = {pollen-feeding pollen enzyme proteolytic activity pre-oral digestion apis-mellifera pollen adaptations digestion patterns drones},
  Year = {2007}
}
• [2007,article] bibtex
  J. M. Douglas, T. W. Cronin, T. H. Chiou, and N. J. Dominy, "Light habitats and the role of polarized iridescence in the sensory ecology of neotropical nymphalid butterflies (Lepidoptera : Nymphalidae)," Journal of Experimental Biology, vol. 210, iss. 5, pp. 788-799, 2007.
  @article{ Author = {Douglas, J. M. and Cronin, T. W. and Chiou, T. H. and Dominy, N. J.},
  Title = {Light habitats and the role of polarized iridescence in the sensory ecology of neotropical nymphalid butterflies (Lepidoptera : Nymphalidae)},
  Journal = {Journal of Experimental Biology},
  Volume = {210},
  Number = {5},
  Pages = {788-799},
  Note = {142EU Times Cited:0 Cited References Count:61},
  Abstract = {The exploitation of polarized light may increase perceived visual contrast independent of spectrum and intensity and thus have adaptive value in forest habitats, where illumination varies greatly in brightness and spectral properties. Here we investigate the extent to which Costa Rican butterflies of the family Nymphalidae exhibit polarized wing reflectance and evaluate the types of habitats in which the trait is commonly found. We also examine the degree of polarized reflectance of wing patterns in representative species belonging to the nymphalid subfamilies Charaxinae, Heliconiinae, Morphinae and Nymphalinae. Polarized reflectance was evaluated using museum specimens illuminated with a light source that simulated the spectrum of ambient sunlight and viewed through a polarized filter. Of the 144 species examined, 75 species exhibited polarized reflectance patterns. These species were significantly more likely to occupy forest habitats than open habitats. A concentrated changes test performed on a phylogeny of the Nymphalidae, with the Papilionidae as an outgroup, provides further support for the correlated evolution of polarized iridescence and life in a forest light environment. These results are consistent with the hypothesis that the production and detection of polarized light may have adaptive communicative value in those species inhabiting forest habitats with complex light conditions. The potential utility of polarized iridescence and iridescent wing coloration within differing ambient spectral environments is discussed to provide a basis for future investigation of the polarized light ecology of butterflies.},
  Keywords = {polarized iridescence butterfly light habitat mitochondrial-DNA sequences dependent color-vision dorsal rim area phylogenetic-relationships heliconius butterflies papilio butterflies compound eye sensitivity evolution pattern},
  Year = {2007}
}
• [2007,article] bibtex
  D. W. Franks and T. N. Sherratt, "The evolution of multicomponent mimicry," Journal of Theoretical Biology, vol. 244, iss. 4, pp. 631-639, 2007.
  @article{ Author = {Franks, D. W. and Sherratt, T. N.},
  Title = {The evolution of multicomponent mimicry},
  Journal = {Journal of Theoretical Biology},
  Volume = {244},
  Number = {4},
  Pages = {631-639},
  Note = {141MZ Times Cited:0 Cited References Count:36},
  Abstract = {The relative sizes of phenotypic mutations contributing to evolutionary change has long been the subject of debate. We describe how mimicry research can shed light on this debate, and frame mimicry studies within the general context of macromutationism and micromutationism. and punctuated versus gradual evolution. Balogh and Leimar [Mullerian mimicry: an examination of Fisher's theory of gradual evolutionary change. Proc. Roy. Soc. Lond. B Biol. Sci. 272, 2269-2275] have recently used a model to readdress the question of whether or not mimicry evolves gradually along a single dimension. We extend their approach, and present the first model to consider the effect of predator generalization along multiple components on the evolution of mimicry. We find that the gradual evolution of mimicry becomes increasingly less likely as the number of signal components increases, unless predators generalize widely over all components. However, we show that the contemporary two-step hypothesis (punctuated evolution followed by gradual refinement) can explain the evolution of Mullerian mimicry under all tested conditions. Thus, although the gradual evolution of mimicry is possible, the two-step hypothesis appears more generally applicable. Crown Copyright (c) 2006 Published by Elsevier Ltd. All rights reserved.},
  Keywords = {multicomponent mullerian mimicry gradualism advergence generalization heliconius butterflies mullerian mimicry warning color adaptation genetics radiation paradigm},
  Year = {2007}
}
• [2007,article] bibtex
  H. S. Engler-Chaouat and L. E. Gilbert, "De novo synthesis vs. sequestration: Negatively correlated metabolic traits and the evolution of host plant specialization in cyanogenic butterflies," Journal of Chemical Ecology, vol. 33, iss. 1, pp. 25-42, 2007.
  @article{ Author = {Engler-Chaouat, H. S. and Gilbert, L. E.},
  Title = {De novo synthesis vs. sequestration: Negatively correlated metabolic traits and the evolution of host plant specialization in cyanogenic butterflies},
  Journal = {Journal of Chemical Ecology},
  Volume = {33},
  Number = {1},
  Pages = {25-42},
  Note = {119GZ Times Cited:3 Cited References Count:86},
  Abstract = {Larvae of Heliconius butterflies (Nymphalidae: Heliconiinae) feed exclusively on cyanogenic leaves of Passiflora (passion vine). Most Heliconius manufacture cyanogenic glycosides (cyanogens) and some species sequester cyanogens from host plants. We compare ability to sequester simple monoglycoside cyclopentenyl (SMC) cyanogens and manufacture aliphatic cyanogens in 12 Heliconius species, including larvae that are specialized (single host species) and generalized (many host species). All butterflies tested higher for cyanide concentrations when reared on plants that larvae can sequester from (SMC plants) than when reared on plants that larvae do not sequester from (non-SMC plants). Specialists in the sara-sapho clade sequestered SMC cyanogens from specific host plants at seven times that of Passiflora generalists fed the same hosts. In contrast, sara-sapho clade species reared on non-SMC plants had significantly lower cyanide concentrations from de novo synthesis than generalists fed the same plants. Furthermore, cyanogen analyses indicated that Heliconius sara butterflies reared on an SMC host had a greater proportion of sequestered SMC cyanogens (95.0%) than de novo-synthesized aliphatic cyanogens (5.0%). Thus, sequestration and de novo synthesis are negatively correlated traits. Results suggest that losing the ability to synthesize cyanogens has restricted sara-sapho clade species to specific hosts containing SMC cyanogens and explains dietary restriction in this clade.},
  Keywords = {plant-insect interactions host plant specialization secondary compounds cyanogenic glycosides sequestration de novo synthesis coevolution phytophagous insects genetic-variation cyanohydrin glycosides heliconius butterflies generalist predators leaf beetle trade-offs lepidoptera coevolution nymphalidae},
  Year = {2007}
}
• [2007,article] bibtex
  M. Joron, R. Papa, J. Mallet, W. O. McMillan, and C. D. Jiggins, "Conserved but flexible: Genetic control of mimicry in Heliconius butterfly wing patterns," Journal of Insect Science, vol. 7, 2007.
  @article{ Author = {Joron, M. and Papa, R. and Mallet, J. and McMillan, W. O. and Jiggins, C. D.},
  Title = {Conserved but flexible: Genetic control of mimicry in Heliconius butterfly wing patterns},
  Journal = {Journal of Insect Science},
  Volume = {7},
  Pages = {-},
  Note = {166PS Times Cited:0 Cited References Count:0},
  Abstract = {It is now well known that convergent morphology can evolve via repeated recruitment of the same regulatory genes in different lineages. We here contrast three butterfly species, all classic examples of Müllerian mimicry. We use a genetic linkage map to show that a locus, Yb, controlling the presence of a yellow band in geographic races of H. melpomene maps precisely to the same location as the locus Cr, which has very similar phenotypic effects in its co-mimic H. erato. Furthermore, the same genomic location acts as a 'supergene' determining multiple, sympatric morphs in a third species, H. numata, a species with a very different phenotypic appearance, whose many forms mimic different unrelated ithomiine butterflies in the genus Melinaea. Hence, a single locus from the multilocus colour pattern architecture in H. melpomene and H. erato appears to have gained control of the entire wing-pattern variability in H. numata, presumably as a result of selection for mimetic 'supergene' polymorphism without intermediates. Our results imply that a conserved, yet relatively unconstrained mechanism underlying pattern switching can affect mimicry in radically different ways. We also show that adaptive evolution, both convergent and diversifying, can occur by the repeated involvement of the same genomic regions.},
  Year = {2007}
}
• [2007,article] bibtex
  S. Schulz, S. Yildizhan, K. Stritzke, C. Estrada, and L. E. Gilbert, "Macrolides from the scent glands of the tropical butterflies Heliconius cydno and Heliconius pachinus," Organic & Biomolecular Chemistry, vol. 5, iss. 21, pp. 3434-3441, 2007.
  @article{ Author = {Schulz, S. and Yildizhan, S. and Stritzke, K. and Estrada, C. and Gilbert, L. E.},
  Title = {Macrolides from the scent glands of the tropical butterflies Heliconius cydno and Heliconius pachinus},
  Journal = {Organic & Biomolecular Chemistry},
  Volume = {5},
  Number = {21},
  Pages = {3434-3441},
  Note = {221GP Times Cited:1 Cited References Count:25},
  Abstract = {The four major components present in scent gland extracts of the male Costa Rica longwing butterflies Heliconius cydno and Heliconius pachinus were identified as 12- and 14-membered macrolides containing a C-18-carbon skeleton. By use of micro-reactions and spectrometric examinations, structural proposals were made and subsequently proven by synthesis, using ring-closing-metathesis as the key steps. These macrolides, ( 9Z, 11E, 13S)-octadeca- 9,11-dien-13-olide (5, S-coriolide), (9Z, 11E, 13S, 15Z)-octadeca- 9,11,15-trien-13-olide (6), (9Z, 13S)-octadec-9-en-13-olide(13), and (9Z, 11S)-octadec-9-en-11-olide( 25), are biosynthetically obviously derived from oleic, linoleic, and linolenic acids. Their absolute configurations were determined by gas chromatographic investigations on chiral phases, showing all to possess (S)-configuration.},
  Keywords = {mass-spectrometry double-bond fatty-acid metathesis position lactone},
  Year = {2007}
}
• [2007,article] bibtex
  M. R. Kronforst, C. Salazar, M. Linares, and L. E. Gilbert, "No genomic mosaicism in a putative hybrid butterfly species," Proceedings of the Royal Society B-Biological Sciences, vol. 274, iss. 1615, pp. 1255-1264, 2007.
  @article{ Author = {Kronforst, M. R. and Salazar, C. and Linares, M. and Gilbert, L. E.},
  Title = {No genomic mosaicism in a putative hybrid butterfly species},
  Journal = {Proceedings of the Royal Society B-Biological Sciences},
  Volume = {274},
  Number = {1615},
  Pages = {1255-1264},
  Note = {151PE Times Cited:0 Cited References Count:40},
  Abstract = {Recent descriptions of hybrid animal species have spurred interest in this phenomenon, but little genomic data exist to support it. Here, we use frequency variation for 657 amplified fragment length polymorphism (AFLP) markers and DNA sequence variation from 16 genes to determine whether the genome of Heliconius pachinus, a suspected hybrid butterfly species, is a mixture of the putative parental species, Heliconius cydno and Heliconius melpomene. Despite substantial shared genetic variation among all three species, we show that the genome of H. pachinus is not a mosaic; both AFLP and DNA sequence data overwhelmingly associate H. pachinus with just one of the potential parents, H. cydno. This pattern also applies to the gene wingless, which is tightly linked to the locus that determines forewing colour-one specific H. pachinus trait that has been hypothesized to have originated from H. melpomene. As a whole, the data support a traditional, bifurcating model of speciation in which H. pachinus split from a common ancestor with H. cydno without a genetic contribution from H. melpomene. However, comparison of our data to DNA sequence data for another putative hybrid Heliconius species, Heliconius heurippa, suggests that the H. heurippa genome may be a mosaic.},
  Keywords = {amplified fragment length polymorphism heliconius introgressive hybridization hybrid speciation lepidoptera mitochondrial-DNA sequences multilocus genotype data heliconius butterflies cladistic-analysis nymphalidae speciation lepidoptera population hybridization inference},
  Year = {2007}
}
• [2007,article] bibtex
  M. R. Kronforst, L. G. Young, and L. E. Gilbert, "Reinforcement of mate preference among hybridizing Heliconius butterflies," Journal of Evolutionary Biology, vol. 20, iss. 1, pp. 278-285, 2007.
  @article{ Author = {Kronforst, M. R. and Young, L. G. and Gilbert, L. E.},
  Title = {Reinforcement of mate preference among hybridizing Heliconius butterflies},
  Journal = {Journal of Evolutionary Biology},
  Volume = {20},
  Number = {1},
  Pages = {278-285},
  Note = {117WS Times Cited:1 Cited References Count:34},
  Abstract = {Recent models of mate preference evolution suggest that direct selection on alleles at preference loci and correlated evolution of preference with locally adapted mating cues are more likely to drive the evolution of assortative mate preference than reinforcement. Mate preference evolution in mimetic Heliconius butterflies has been attributed to all three forms of selection, but here we show that reinforcement has been critical. By examining geographical variation in assortative mating and male mate preference among seven populations of three hybridizing Heliconius species from Costa Rica, we found pronounced character displacement of preference such that sexual isolation was enhanced in areas of interspecific contact. Of the different explanations for the evolution of assortative mate preference, only reinforcement is dependent on interspecific contact in this system. Thus, the observed pattern of reproductive character displacement of mate preference is best explained as a product of indirect selection generated by natural selection against nonmimetic hybrids.},
  Keywords = {assortative mating lepidoptera natural selection reproductive character displacement speciation reproductive character displacement mitochondrial-DNA sequences premating isolation natural-selection mullerian mimicry sexual selection gene flow speciation evolution melpomene},
  Year = {2007}
}
• [2007,article] bibtex
  J. A. Addison, "Simultaneous cloning of multiple nuclear genes by pooling PCR products of variable size: a cost-effective method of improving efficiency in large-scale genetic analyses," Molecular Ecology Notes, vol. 7, iss. 3, pp. 389-392, 2007.
  @article{ Author = {Addison, J. A.},
  Title = {Simultaneous cloning of multiple nuclear genes by pooling PCR products of variable size: a cost-effective method of improving efficiency in large-scale genetic analyses},
  Journal = {Molecular Ecology Notes},
  Volume = {7},
  Number = {3},
  Pages = {389-392},
  Note = {164OV Times Cited:0 Cited References Count:14},
  Abstract = {I present a simple approach to overcome the high cost and low efficiency of cloning polymerase chain reaction (PCR) products for individuals in wide-scale population genetic analyses. The methodology reduces the number of cloning reactions per individual by engineering a suite of genetic markers that differ in size and pooling these PCR products prior to cloning. Alleles from each gene are then recovered by screening transformed bacterial colonies and identifying the inserts corresponding to each gene based on size. I demonstrate the utility of this technique by presenting the results I obtained from cloning four nuclear genes in 118 individuals from three species of sea urchins (Strongylocentrotus purpuratus, S. droebachiensis and S. pallidus). Of the 472 different PCR products I cloned, I recovered at least one allele for 432 of them (91.5%) by screening between 16 and 32 bacterial colonies for each individual. There existed a bias with respect to recovery efficiency: the two largest fragments (1130-800 bp) were recovered 100% of the time, while the two smaller fragments (580-650 bp) were recovered in 85.6% and 81.4% of the experiments, respectively. I discuss the promise of this application for wide-scale genetic analyses.},
  Keywords = {diploid DNA polymorphism nuclear gene pcr pooled cloning heliconius butterflies phylogeography populations sequences prospects},
  Year = {2007}
}
• [2007,article] bibtex
  J. Mallet, M. Beltran, W. Neukirchen, and M. Linares, "Natural hybridization in heliconiine butterflies: the species boundary as a continuum," Bmc Evolutionary Biology, vol. 7, 2007.
  @article{ Author = {Mallet, J. and Beltran, M. and Neukirchen, W. and Linares, M.},
  Title = {Natural hybridization in heliconiine butterflies: the species boundary as a continuum},
  Journal = {Bmc Evolutionary Biology},
  Volume = {7},
  Pages = {-},
  Note = {146LX Times Cited:2 Cited References Count:102},
  Abstract = {Background: To understand speciation and the maintenance of taxa as separate entities, we need information about natural hybridization and gene flow among species. Results: Interspecific hybrids occur regularly in Heliconius and Eueides (Lepidoptera: Nymphalidae) in the wild: 26-29% of the species of Heliconiina are involved, depending on species concept employed. Hybridization is, however, rare on a per-individual basis. For one well-studied case of species hybridizing in parapatric contact (Heliconius erato and H. himera), phenotypically detectable hybrids form around 10% of the population, but for species in sympatry hybrids usually form less than 0.05% of individuals. There is a roughly exponential decline with genetic distance in the numbers of natural hybrids in collections, both between and within species, suggesting a simple "exponential failure law" of compatibility as found in some prokaryotes. Conclusion: Hybridization between species of Heliconius appears to be a natural phenomenon; there is no evidence that it has been enhanced by recent human habitat disturbance. In some well-studied cases, backcrossing occurs in the field and fertile backcrosses have been verified in insectaries, which indicates that introgression is likely, and recent molecular work shows that alleles at some but not all loci are exchanged between pairs of sympatric, hybridizing species. Molecular clock dating suggests that gene exchange may continue for more than 3 million years after speciation. In addition, one species, H. heurippa, appears to have formed as a result of hybrid speciation. Introgression may often contribute to adaptive evolution as well as sometimes to speciation itself, via hybrid speciation. Geographic races and species that coexist in sympatry therefore form part of a continuum in terms of hybridization rates or probability of gene flow. This finding concurs with the view that processes leading to speciation are continuous, rather than sudden, and that they are the same as those operating within species, rather than requiring special punctuated effects or complete allopatry. Although not qualitatively distinct from geographic races, nor "real" in terms of phylogenetic species concepts or the biological species concept, hybridizing species of Heliconius are stably distinct in sympatry, and remain useful groups for predicting morphological, ecological, behavioural and genetic characteristics.},
  Keywords = {passion-vine butterflies mitochondrial-DNA hybrid zones gene flow reproductive isolation sympatric speciation adaptive radiation mullerian mimicry close relatives warning color},
  Year = {2007}
}
• [2007,article] bibtex
  M. Kronforst, L. G. Young, and L. E. Gilbert, "Reinforcement of mate preference among hybridizing Heliconius butterflies," Journal of Evolutionary Biology, vol. 20, pp. 278-285, 2007.
  @article{ Author = {Kronforst, M. and Young, L. G. and Gilbert, L. E.},
  Title = {Reinforcement of mate preference among hybridizing Heliconius butterflies },
  Journal = {Journal of Evolutionary Biology},
  Volume = {20},
  Pages = {278-285},
  Year = {2007}
}
• [2007,article] bibtex
  R. J. Parchem, M. W. Perry, and N. H. Patel, "Patterns on the insect wing," Current Opinion in Genetics & Development, vol. 17, iss. 4, pp. 300-308, 2007.
  @article{ Author = {Parchem, R. J. and Perry, M. W. and Patel, N. H.},
  Title = {Patterns on the insect wing},
  Journal = {Current Opinion in Genetics & Development},
  Volume = {17},
  Number = {4},
  Pages = {300-308},
  Note = {209CQ Times Cited:0 Cited References Count:51},
  Abstract = {The evolution of wings and the adaptive advantage they provide have allowed insects to become one of the most evolutionarily successful groups on earth. The incredible diversity of their shape, size, and color patterns is a direct reflection of the important role wings have played in the radiation of insects. In this review, we highlight recent studies on both butterflies and Drosophila that have begun to uncover the types of genetic variations and developmental mechanisms that control diversity in wing color patterns. In butterflies, these analyses are now possible because of the recent development of a suite of genomic and functional tools, such as detailed linkage maps and transgenesis. In one such study, extensive linkage mapping in Heliconius butterflies has shown that surprisingly few, and potentially homologous, loci are responsible for several major pattern variations on the wings of these butterflies. Parallel work on a clade of Drosophila has uncovered how cis-regulatory changes of the same gene correlate with the repeated gain and loss of pigmented wing spots. Collectively, our understanding of formation and evolution of color pattern in insect wings is rapidly advancing because of these recent breakthroughs in several different fields.},
  Keywords = {silkworm bombyx-mori heliconius butterflies pigment patterns linkage map bicyclus-anynana evolution mimicry scales drosophila transformation},
  Year = {2007}
}
• [2007,article] bibtex
  K. Sahara, A. Yoshido, F. Marec, I. Fukova, H. B. Zhang, C. C. Wu, M. R. Goldsmith, and Y. Yasukochi, "Conserved synteny of genes between chromosome 15 of Bombyx mori and a chromosome of Manduca sexta shown by five-color BAC-FISH," Genome, vol. 50, iss. 11, pp. 1061-1065, 2007.
  @article{ Author = {Sahara, K. and Yoshido, A. and Marec, F. and Fukova, I. and Zhang, H. B. and Wu, C. C. and Goldsmith, M. R. and Yasukochi, Y.},
  Title = {Conserved synteny of genes between chromosome 15 of Bombyx mori and a chromosome of Manduca sexta shown by five-color BAC-FISH},
  Journal = {Genome},
  Volume = {50},
  Number = {11},
  Pages = {1061-1065},
  Note = {245IO Times Cited:1 Cited References Count:17},
  Abstract = {The successful assignment of the existing genetic linkage groups (LGs) to individual chromosomes and the second-generation linkage map obtained by mapping a large number of bacterial artificial chromosome (BAC) contigs in the silkworm, Bombyx mori, together with public nucleotide sequence databases, offer a powerful tool for the study of synteny between karyotypes of B. mori and other lepidopteran species. Conserved synteny of genes between particular chromosomes can be identified by comparatively mapping orthologous genes of the corresponding linkage groups with the help of BAC-FISH (fluorescent in situ hybridization). This technique was established in B. mori for 2 differently labeled BAC probes simultaneously hybridized to pachytene bivalents. To achieve higher-throughput comparative mapping using BAC-FISH in Lepidoptera, we developed a protocol for five-color BAC-FISH, which allowed us to map simultaneously 6 different BAC probes to chromosome 15 in B. mori. We identified orthologs of 6 B. mori LG15 genes (RpP0, RpS8, eIF3, RpL7A, RpS23, and Hsc70) for the tobacco hornworm, Manduca sexta, and selected the ortholog-containing BAC clones from an M. sexta BAC library. All 6 M. sexta BAC clones hybridized to a single M. sexta bivalent in pachytene spermatocytes. Thus, we have confirmed the conserved synteny between the B. mori chromosome 15 and the corresponding M. sexta chromosome (hence provisionally termed chromosome 15).},
  Keywords = {bac-fish bombyx mori chromosome conserved synteny lepidoptera linkage map manduca sexta heliconius-melpomene silkworm lepidoptera sequence insects genome map},
  Year = {2007}
}
• [2007,article] bibtex
  M. Z. Cardoso and L. E. Gilbert, "A male gift to its partner? Cyanogenic glycosides in the spermatophore of longwing butterflies (Heliconius)," Naturwissenschaften, vol. 94, iss. 1, pp. 39-42, 2007.
  @article{ Author = {Cardoso, M. Z. and Gilbert, L. E.},
  Title = {A male gift to its partner? Cyanogenic glycosides in the spermatophore of longwing butterflies (Heliconius)},
  Journal = {Naturwissenschaften},
  Volume = {94},
  Number = {1},
  Pages = {39-42},
  Note = {117EG Times Cited:2 Cited References Count:23},
  Abstract = {Males of several insect species transfer nuptial gifts to females during mating, typically in the form of a protein-rich spermatophore. In chemically defended species, males could potentially enhance such a gift with chemicals that help protect the female, her eggs, or both. This was shown for lepidopteran species that accumulate pyrrolizidine alkaloids. Most Heliconius butterflies are presumably protected from predators by virtue of de novo synthesized and/or sequestered cyanogenic glycosides. Males of Heliconius species are known to transfer nutritional gifts to the females but whether defensive chemicals could also be transferred is not known. To ascertain whether transfer of cyanogens occurs, we dissected freshly mated females from nine different Heliconius species and analyzed spermatophores for cyanogenic glycosides. We found cyanogens in the spermatophores of all nine species. This is the first time cyanogenic glycosides are reported in the spermatophores of arthropods. We discuss the implications of these findings for Heliconius biology and for other cyanogenic insects as well. We suggest that chemically defended species commonly lace their nuptial gifts with defensive chemicals to improve gift quality.},
  Keywords = {nuptial gifts spermatophore heliconius cyanogenic glycoside utetheisa-ornatrix lepidoptera glucosides linamarin biosynthesis lotaustralin fecundity insecta cyanide defense},
  Year = {2007}
}
• [2007,article] bibtex
  A. S. Putnam, J. M. Scriber, and P. Andolfatto, "Discordant divergence times among Z-chromosome regions between two ecologically distinct swallowtail butterfly species," Evolution, vol. 61, iss. 4, pp. 912-927, 2007.
  @article{ Author = {Putnam, A. S. and Scriber, J. M. and Andolfatto, P.},
  Title = {Discordant divergence times among Z-chromosome regions between two ecologically distinct swallowtail butterfly species},
  Journal = {Evolution},
  Volume = {61},
  Number = {4},
  Pages = {912-927},
  Note = {158UD Times Cited:4 Cited References Count:93},
  Abstract = {We investigate multi locus patterns of differentiation between parental populations of two swallowtail butterfly species that differ at a number of ecologically important sex-linked traits. Using a new coalescent-based approach, we show that there is significant heterogeneity in estimated divergence times among five Z-linked markers, rejecting a purely allopatric speciation model. We infer that the Z chromosome is a mosaic of regions that differ in the extent of historical gene flow, potentially due to isolating barriers that prevent the introgression of species-specific traits that result in hybrid incompatibilities. Surprisingly, a candidate region for a strong barrier to introgression, Ldh, does not show a significantly deeper divergence time than other markers on the Z chromosome. Our approach can be used to test alternative models of speciation and can potentially assign chronological order to the appearance of factors contributing to reproductive isolation between species.},
  Keywords = {approximate bayesian computation divergence hybrid zones introgression lepidoptera papilio speciation hybrid male-sterility drosophila-melanogaster papilio-glaucus gene flow population-genetics nuclear genes heliconius butterflies natural-selection DNA polymorphism migration rates},
  Year = {2007}
}
• [2007,article] bibtex
  C. E. G. Pinheiro, "Asynchrony in daily activity patterns of butterfly models and mimics," Journal of Tropical Ecology, vol. 23, pp. 119-123, 2007.
  @article{ Author = {Pinheiro, C. E. G.},
  Title = {Asynchrony in daily activity patterns of butterfly models and mimics},
  Journal = {Journal of Tropical Ecology},
  Volume = {23},
  Pages = {119-123},
  Note = {Part 1 135OE Times Cited:0 Cited References Count:28},
  Keywords = {atlantic forest batesian mimicry brazil insectivorous birds neotropical butterflies rufous-tailed jacamars neotropical butterflies heliconius butterflies locomotor mimicry flight morphology rain-forest prey palatability temperature tyrannidae},
  Year = {2007}
}
• [2007,article] bibtex
  E. G. Pringle, S. W. Baxter, C. L. Webster, A. Papanicolaou, S. F. Lee, and C. D. Jiggins, "Synteny and chromosome evolution in the lepidoptera: Evidence from mapping in Heliconius melpomene," Genetics, vol. 177, iss. 1, pp. 417-426, 2007.
  @article{ Author = {Pringle, E. G. and Baxter, S. W. and Webster, C. L. and Papanicolaou, A. and Lee, S. F. and Jiggins, C. D.},
  Title = {Synteny and chromosome evolution in the lepidoptera: Evidence from mapping in Heliconius melpomene},
  Journal = {Genetics},
  Volume = {177},
  Number = {1},
  Pages = {417-426},
  Note = {218XM Times Cited:4 Cited References Count:45},
  Abstract = {The extent of conservation of synteny and gene order in the Lepidoptera has been investigated previously only by comparing a small subset of linkage groups between the moth Bombyx mori and the butterfly Heliconius melpomene. Here we report the mapping of 64 additional conserved genes in H. melpomene, which contributed 47 markers to a comparative framework of 72 orthologous loci spanning all 21 H. melpomene chromosomes and 27 of the 28 B. mori chromosomes. Comparison of the maps revealed conserved synteny across all chromosomes for the 72 loci, as well as evidence for six cases of chromosome fusion in the Heliconius lineage that contributed to the derived 21-chromosome karyotype. Comparisons of gene order on these fused chromosomes revealed two instances of colinearity between H. melpomene and B. mori, but also one instance of likely chromosomal rearrangement. B. mori is the first lepidopteran species to have its genome sequenced, and the finding that there is conserved synteny and gene order among Lepidoptera indicates that the genomic tools developed in B. mori will be broadly useful in other species.},
  Keywords = {genetic-linkage map evo-devo model bombyx-mori drosophila-melanogaster caenorhabditis-briggsae domesticated silkworm subfamily nymphalinae comparative genomics anopheles-gambiae warning color},
  Year = {2007}
}
• [2007,article] bibtex
  C. D. Jiggins, "The evolutionary importance of mimicry in Heliconius butterflies," Journal of Insect Science, vol. 7, 2007.
  @article{ Author = {Jiggins, C. D.},
  Title = {The evolutionary importance of mimicry in Heliconius butterflies},
  Journal = {Journal of Insect Science},
  Volume = {7},
  Pages = {-},
  Note = {166PS Times Cited:0 Cited References Count:0},
  Abstract = {eliconius butterflies are well known for their bright mimetic colour patterns. Here I describe studies of speciation among the Heliconius melpomene species group. Differences in colour pattern that distinguish species involve the same genetic basis as within-species variation. Furthermore colour patterns are used as cues in mating, and therefore pattern shifts lead to reproductive isolation and contribute to speciation. In one Colombian species, Heliconius heurippa it has been suggested that a novel pattern may have arisen as a result of hybridisation between H. melpomene and H. cydno. By means of crosses we have demonstrated that the H. heurippa pattern can be re-created within three generations of hybridisation in the laboratory. Furthermore, both elements of the hybrid pattern are necessary for mate recognition, such that the hybrid trait is clearly causing reproductive isolation between natural populations. Other examples suggest that a hybrid origin for novel patterns may be relatively common in these butterflies. A definitive test of this hypothesis awaits identification of the genes at a molecular basis.},
  Year = {2007}
}
• [2007,article] bibtex
  E. J. Baack and L. H. Rieseberg, "A genomic view of introgression and hybrid speciation," Current Opinion in Genetics & Development, vol. 17, iss. 6, pp. 513-518, 2007.
  @article{ Author = {Baack, E. J. and Rieseberg, L. H.},
  Title = {A genomic view of introgression and hybrid speciation},
  Journal = {Current Opinion in Genetics & Development},
  Volume = {17},
  Number = {6},
  Pages = {513-518},
  Note = {244HL Times Cited:0 Cited References Count:54},
  Abstract = {Hybridization in plants and animals is more common and has more complex outcomes than previously realized. Genome-wide analyses of introgression in organisms ranging from oaks to sunflowers to fruit flies show that a substantial fraction of their genomes are permeable to alleles from related species. Hybridization can lead to rapid genomic changes, including chromosomal rearrangements, genome expansion, differential gene expression, and gene silencing, some of which are mediated by transposable elements. These genomic changes may lead to beneficial new phenotypes, and selection for fertility and ecological traits may in turn alter genome structure. Dramatic increases in the availability of genomic tools will produce a new understanding of the genetic nature of species and will resolve a century-old debate over the basis of hybrid vigor, while the natural recombinants found in hybrid zones will permit genetic mapping of species differences and reproductive barriers in nonmodel organisms.},
  Keywords = {recurrent gene flow ancestral polymorphism heliconius butterflies hybridization expression evolution heterosis loci drosophila patterns},
  Year = {2007}
}
• [2007,article] bibtex
  S. W. Baxter, N. Chamberlain, R. Papa, S. J. Humphray, R. H. Ffrench-Constant, W. O. McMillan, and C. D. Jiggins, "Identifying DNA markers close to quantitative traits in lepidopteran genomes: Using wing colour variation in Heliconius butterflies as a model," Journal of Insect Science, vol. 7, 2007.
  @article{ Author = {Baxter, S. W. and Chamberlain, N. and Papa, R. and Humphray, S. J. and Ffrench-Constant, R. H. and McMillan, W. O. and Jiggins, C. D.},
  Title = {Identifying DNA markers close to quantitative traits in lepidopteran genomes: Using wing colour variation in Heliconius butterflies as a model},
  Journal = {Journal of Insect Science},
  Volume = {7},
  Pages = {-},
  Note = {166PS Times Cited:0 Cited References Count:0},
  Abstract = {Identifying a gene responsible for a trait of interest within Lepidoptera is a multi-step process. Commonly, this involves performing crosses to enable trait segregation, using progeny to create linkage maps, identifying the genome region responsible for variation and then genetic mapping of candidate genes to test for associations. If candidate genes tested do not map to the chromosome or locus of interest, sequencing genomic BAC clones spanning the genome region is required. Here we present a molecular method for identifying DNA markers tightly linked to a quantitative trait, which can be used to screen BAC libraries, using wing colour variation in Heliconius butterflies as a model. Heliconius melpomene and H. erato are co-mimics that share the same warningly coloured wing patterns where they cohabit throughout Central and South America. Approximately 30 races of wing colour variants have been characterised and identifying the colour determining genes will be of value to developmental and evolutionary biology, and population genetics. We have used AFLP methods to identify the genetic locus of a strong visual trait in these sympatric species – red wing patches. Two dominant loci on the same chromosome of H. melpomene have previously been found to control red wing colouration: “B” is a large red spot on the forewing and “Dennis” (D) includes ray patterns on the hind wing and red colour on the proximal forewing. Crosses were performed between B/D heterozygotes (BbDd x BbDd) then AFLPs were performed on two progeny bulks; i) homozygotes lacking the B spot (bbD-) or ii) lacking the Dennis pattern (B-dd). When AFLP analysis was performed, it was expected that both bulks would share identical banding patterns, except in areas tightly linked to the B or D loci. AFLP analysis was performed using fluorescently labelled primers on a 3730 DNA Analyzer (Applied Biosystems). From approximately 300 primer combinations, nine bands were identified and confirmed linked the B or D locus. AFLPs were sequenced and used to create genotyping assays for additional broods, which found B and D to be tightly linked. Multiple of the AFLP markers were within 1 centimorgan (~180kb) of this region, and were used as probes to screen a H. melpomene BAC library. A similar strategy has been used to identify an AFLP band close to the Dry locus in H. erato, which controls the same phenotype as B and D in H. melpomene. Mapping a gene close to the H. erato AFLP in H. melpomene, has shown homologous genetic regions are responsible for similar red wing phenotypes in these two species. Performing AFLP analysis on bulks of progeny, selected for the presence or absence of a quantitative trait, could be applied to other lepidopteran species.},
  Year = {2007}
}
• [2007,article] bibtex
  M. Beltran, C. D. Jiggins, A. V. Z. Brower, E. Bermingham, and J. Mallet, "Do pollen feeding, pupal-mating and larval gregariousness have a single origin in Heliconius butterflies? Inferences from multilocus DNA sequence data," Biological Journal of the Linnean Society, vol. 92, iss. 2, pp. 221-239, 2007.
  @article{ Author = {Beltran, M. and Jiggins, C. D. and Brower, A. V. Z. and Bermingham, E. and Mallet, J.},
  Title = {Do pollen feeding, pupal-mating and larval gregariousness have a single origin in Heliconius butterflies? Inferences from multilocus DNA sequence data},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {92},
  Number = {2},
  Pages = {221-239},
  Note = {211ZG Times Cited:0 Cited References Count:79},
  Abstract = {Phylogenetic information is useful in understanding the evolutionary history of adaptive traits. Here, we present a well-resolved phylogenetic hypothesis for Heliconius butterflies and related genera. We use this tree to investigate the evolution of three traits, pollen feeding, pupal-mating behaviour and larval gregariousness. Phylogenetic relationships among 60 Heliconiina species (86% of the subtribe) were inferred from partial DNA sequences of the mitochondrial genes cytochrome oxidase I, cytochrome oxidase II and 16S rRNA, and fragments of the nuclear genes elongation factor-1 alpha, apterous, decapentaplegic and wingless (3834 bp in total). The results corroborate previous hypotheses based on sequence data in showing that Heliconius is paraphyletic, with Laparus doris and Neruda falling within the genus, demonstrating a single origin for pollen feeding but with a loss of the trait in Neruda. However, different genes are not congruent in their placement of Neruda; therefore, monophyly of the pollen feeding species cannot be ruled out. There is also a highly supported monophyletic 'pupal-mating clade' suggesting that pupal mating behaviour evolved only once in the Heliconiina. Additionally, we observed at least three independent origins for larval gregariousness from a solitary ancestor, showing that gregarious larval behaviour arose after warning coloration. (C) 2007 The Linnean Society of London.},
  Keywords = {bayesian analysis ef1 alpha mimicry mtdna parsimony phylogeny passion-vine butterflies mitochondrial-DNA phylogenetic analysis elongation factor-1-alpha nymphalid butterflies bayesian-inference cladistic-analysis nuclear gene linkage map lepidoptera},
  Year = {2007}
}
• [2007,article] bibtex
  N. Wahlberg and A. V. L. Freitas, "Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera : Nymphalidae)," Molecular Phylogenetics and Evolution, vol. 44, iss. 3, pp. 1257-1272, 2007.
  @article{ Author = {Wahlberg, N. and Freitas, A. V. L.},
  Title = {Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera : Nymphalidae)},
  Journal = {Molecular Phylogenetics and Evolution},
  Volume = {44},
  Number = {3},
  Pages = {1257-1272},
  Note = {215YR Times Cited:1 Cited References Count:78},
  Abstract = {The historical biogeography of insects in South America is largely unknown, as dated phylogenies have not been available for most groups. We have studied the phylogenetic relationships and historical biogeography of a subtribe of butterflies, Phyciodina in the family Nymphalidae, based on one mitochondrial gene (COI) and two nuclear gene regions (EF-1 alpha and wingless). The subtribe comprises 89 species mainly found in tropical South America, with a few species in North America and the Greater Antilles. We find that the enigmatic genus Antillea is sister to the rest of Phyciodina, and suggest that it should be included in the subtribe. Several genera are found to be polyphyletic or nested within another genus, and are proposed to be synonymised. These are Dagon, Castilia, Telenassa and Janatella, which we propose should be synonymised with Eresia. Brazilian "Ortilia" form an independent lineage and require a new genus name. The diversification of Phyciodina has probably taken place over the past about 34 MYA. The ancestral phyciodine colonised South America from North America through a possible landspan that connected the Greater Antilles to South America about 34 MYA. A vicariance event left the ancestral Antillea on the Greater Antilles, while the ancestral 0e on South America colonised the Guyanan Shield and soon after the Brazilian Shield. We hypothesise that the Brazilian Shield was an important area for the diversification of Phyciodina. From there, the ancestor of Anthanassa, Eresia and Tegosa colonised NW South America, where especially Eresia diversified in concert with the rising of the Andes beginning about 20 MYA. Central America was colonised from NW South America about 15 MYA by the ancestors of Anthanassa and Phyciodes. Our study is the first to use a dated phylogeny to study the historical biogeography of a group of South American species of butterflies. (c) 2007 Elsevier Inc. All rights reserved.},
  Keywords = {molecular systematics historical biogeography butterfly lepidoptera melitaeini nymphalidae mitochondrial-DNA sequences higher-level phylogeny heliconius butterflies historical biogeography divergence times branch support nuclear gene data sets evolution incongruence},
  Year = {2007}
}
• [2007,article] bibtex
  V. Y. Vedenina, N. K. Kulygina, and A. K. Panyutin, "Isolation mechanisms in the closely related grasshopper species, Chorthippus albomarginatus and Ch. oschei (Orthoptera, Acrididae)," Zoologichesky Zhurnal, vol. 86, iss. 5, pp. 537-546, 2007.
  @article{ Author = {Vedenina, V. Y. and Kulygina, N. K. and Panyutin, A. K.},
  Title = {Isolation mechanisms in the closely related grasshopper species, Chorthippus albomarginatus and Ch. oschei (Orthoptera, Acrididae)},
  Journal = {Zoologichesky Zhurnal},
  Volume = {86},
  Number = {5},
  Pages = {537-546},
  Note = {180CW Times Cited:0 Cited References Count:36},
  Abstract = {Two closely related grasshopper species, Chorthippus albomarginatus and Ch. oschei, are known to hybridize at a narrow contact zone in the territory of Ukraine and Moldova. Different isolation mechanisms providing a reproductive isolation of these species were studied. In the mating experiments, females of both species demonstrated a strong assortative mating (80-90% preference for conspecific males). The comparison of the parental and hybrid viability revealed a reduced hatching and an increased larval mortality in F-1 and F-2 hybrids. In the choice mating experiments, the females mated less assortatively than parental females. The data obtained demonstrated the existence of pre- and post-mating isolation in Ch. albomarginatus and Ch. oschei. A possible fate of the hybrid zone is discussed.},
  Keywords = {gomphocerus-rufus l hybrid zone reproductive isolation acoustic communication jacobsi orthoptera heliconius-cydno sexual selection gene exchange speciation courtship},
  Year = {2007}
}
• [2006,article] bibtex
  C. D. Jiggins, R. Mallarino, K. R. Willmott, and E. Bermingham, "The phylogenetic pattern of speciation and wing pattern change in neotropical Ithomia butterflies (Lepidoptera : Nymphalidae)," Evolution, vol. 60, iss. 7, pp. 1454-1466, 2006.
  @article{ Author = {Jiggins, C. D. and Mallarino, R. and Willmott, K. R. and Bermingham, E.},
  Title = {The phylogenetic pattern of speciation and wing pattern change in neotropical Ithomia butterflies (Lepidoptera : Nymphalidae)},
  Journal = {Evolution},
  Volume = {60},
  Number = {7},
  Pages = {1454-1466},
  Note = {072OI Times Cited:5 Cited References Count:65},
  Abstract = {Species level phylogenetic hypotheses can be used to explore patterns of divergence and speciation. In the tropics, speciation is commonly attributed to either vicariance, perhaps within climate-induced forest refugia, or ecological speciation caused by niche adaptation. Mimetic butterflies have been used to identify forest refugia as well as in studies of ecological speciation, so they are ideal for discriminating between these two models. The genus Ithomia contains 24 species of warningly colored mimetic butterflies found in South and Central America, and here we use a phylogenetic hypothesis based on seven genes for 23 species to investigate speciation in this group. The history of wing color pattern evolution in the genus was reconstructed using both parsimony and likelihood. The ancestral pattern for the group was almost certainly a transparent butterfly, and there is strong evidence for convergent evolution due to mimicry. A punctuationist model of pattern evolution was a significantly better fit to the data than a gradualist model, demonstrating that pattern changes above the species level were associated with cladogenesis and supporting a model of ecological speciation driven by mimicry adaptation. However, there was only one case of sister species unambiguously differing in pattern, suggesting that some recent speciation events have occurred without pattern shifts. The pattern of geographic overlap between clades over time shows that closely related species are mostly sympatric or, in one case, parapatric. This is consistent with modes of speciation with ongoing gene flow, although rapid range changes following allopatric speciation could give a similar pattern. Patterns of lineage accumulation through time differed significantly from that expected at random, and show that most of the extant species were present by the beginning of the Pleistocene at the latest. Hence Pleistocene refugia are unlikely to have played a major role in Ithomia diversification.},
  Keywords = {comparative analysis lepidoptera mimicry pleistocene refugia speciation swallowtail butterflies reproductive isolation molecular phylogenies ecological factors color pattern evolution heliconius diversification mimicry biogeography},
  Year = {2006}
}
• [2006,article] bibtex
  V. Bull, M. Beltran, C. D. Jiggins, W. O. McMillan, E. Bermingham, and J. Mallet, "Polyphyly and gene flow between non-sibling Heliconius species," Bmc Biology, vol. 4, 2006.
  @article{ Author = {Bull, V. and Beltran, M. and Jiggins, C. D. and McMillan, W. O. and Bermingham, E. and Mallet, J.},
  Title = {Polyphyly and gene flow between non-sibling Heliconius species},
  Journal = {Bmc Biology},
  Volume = {4},
  Pages = {-},
  Note = {055RD Times Cited:15 Cited References Count:83},
  Abstract = {Background: The view that gene flow between related animal species is rare and evolutionarily unimportant largely antedates sensitive molecular techniques. Here we use DNA sequencing to investigate a pair of morphologically and ecologically divergent, non-sibling butterfly species, Heliconius cydno and H. melpomene (Lepidoptera: Nymphalidae), whose distributions overlap in Central and Northwestern South America. Results: In these taxa, we sequenced 30-45 haplotypes per locus of a mitochondrial region containing the genes for cytochrome oxidase subunits I and II (CoI/CoII), and intron-spanning fragments of three unlinked nuclear loci: triose-phosphate isomerase (Tpi), mannose-6-phosphate isomerase (Mpi) and cubitus interruptus (Ci) genes. A fifth gene, dopa decarboxylase (Ddc) produced sequence data likely to be from different duplicate loci in some of the taxa, and so was excluded. Mitochondrial and Tpi genealogies are consistent with reciprocal monophyly, whereas sympatric populations of the species in Panama share identical or similar Mpi and Ci haplotypes, giving rise to genealogical polyphyly at the species level despite evidence for rapid sequence divergence at these genes between geographic races of H. melpomene. Conclusion: Recent transfer of Mpi haplotypes between species is strongly supported, but there is no evidence for introgression at the other three loci. Our results demonstrate that the boundaries between animal species can remain selectively porous to gene flow long after speciation, and that introgression, even between non-sibling species, can be an important factor in animal evolution. Interspecific gene flow is demonstrated here for the first time in Heliconius and may provide a route for the transfer of switch-gene adaptations for Mullerian mimicry. The results also forcefully demonstrate how reliance on a single locus may give an erroneous picture of the overall genealogical history of speciation and gene flow.},
  Keywords = {mitochondrial-DNA sequences drosophila-pseudoobscura phylogenetic utility population-genetics dopa decarboxylase cladistic-analysis anopheles-gambiae mullerian mimicry hybrid sterility close relatives},
  Year = {2006}
}
• [2006,article] bibtex
  M. Joron, C. D. Jiggins, A. Papanicolaou, and W. O. McMillan, "Heliconius wing patterns: an evo-devo model for understanding phenotypic diversity," Heredity, vol. 97, iss. 3, pp. 157-167, 2006.
  @article{ Author = {Joron, M. and Jiggins, C. D. and Papanicolaou, A. and McMillan, W. O.},
  Title = {Heliconius wing patterns: an evo-devo model for understanding phenotypic diversity},
  Journal = {Heredity},
  Volume = {97},
  Number = {3},
  Pages = {157-167},
  Note = {077GC Times Cited:9 Cited References Count:77},
  Abstract = {Evolutionary Developmental Biology aims for a mechanistic understanding of phenotypic diversity, and present knowledge is largely based on gene expression and interaction patterns from a small number of well-known model organisms. However, our understanding of biological diversification depends on our ability to pinpoint the causes of natural variation at a micro-evolutionary level, and therefore requires the isolation of genetic and developmental variation in a controlled genetic background. The colour patterns of Heliconius butterflies (Nymphalidae: Heliconiinae) provide a rich suite of naturally occurring variants with striking phenotypic diversity and multiple taxonomic levels of variation. Diversification in the genus is well known for its dramatic colour-pattern divergence between races or closely related species, and for Murllerian mimicry convergence between distantly related species, providing a unique system to study the development basis of colour-pattern evolution. A long history of genetic studies has showed that pattern variation is based on allelic combinations at a surprisingly small number of loci, and recent developmental evidence suggests that pattern development in Heliconius is different from the eyespot determination of other butterflies. Fine-scale genetic mapping studies have shown that a shared toolkit of genes is used to produce both convergent and divergent phenotypes. These exciting results and the development of new genomic resources make Heliconius a very promising evo-devo model for the study of adaptive change.},
  Keywords = {linkage mapping positional cloning expressed sequence tags signalling pathways melanin ommochrome mimicry quantitative trait loci shifting balance theory genetic-linkage map mullerian mimicry warning-color morphological evolution natural-selection bicyclus-anynana hybrid zone distal-less},
  Year = {2006}
}
• [2006,article] bibtex
  M. R. Kronforst, D. D. Kapan, and L. E. Gilbert, "Parallel genetic architecture of parallel adaptive radiations in mimetic heliconius butterflies," Genetics, vol. 174, iss. 1, pp. 535-539, 2006.
  @article{ Author = {Kronforst, M. R. and Kapan, D. D. and Gilbert, L. E.},
  Title = {Parallel genetic architecture of parallel adaptive radiations in mimetic heliconius butterflies},
  Journal = {Genetics},
  Volume = {174},
  Number = {1},
  Pages = {535-539},
  Note = {092YT Times Cited:3 Cited References Count:33},
  Abstract = {It is unknown whether homologous loci underlie the independent and parallel wing pattern radiations of Heliconius butterflies. By comparing the locations of color patterning genes on linkage maps we show that three loci that act similarly in the two radiations are in similar positions on homologous chromosomes.},
  Keywords = {linkage map warning color evolution erato patterns nymphalidae lepidoptera drosophila melpomene mimicry},
  Year = {2006}
}
• [2006,article] bibtex
  T. J. Simonsen, N. Wahlberg, A. V. Z. Brower, and R. de Jong, "Morphology, molecules and fritillaries: approaching a stable phylogeny for Argynnini (Lepidoptera : Nymphalidae)," Insect Systematics & Evolution, vol. 37, iss. 4, pp. 405-418, 2006.
  @article{ Author = {Simonsen, T. J. and Wahlberg, N. and Brower, A. V. Z. and de Jong, R.},
  Title = {Morphology, molecules and fritillaries: approaching a stable phylogeny for Argynnini (Lepidoptera : Nymphalidae)},
  Journal = {Insect Systematics & Evolution},
  Volume = {37},
  Number = {4},
  Pages = {405-418},
  Note = {138AZ Times Cited:2 Cited References Count:43},
  Abstract = {We examine the phylogenetic relationships among 29 species of Argynnini based on 141 previously published morphological characters and new data from the mitochondrial gene COI and the two nuclear genes EF-1a and wingless. We investigate the stability and robustness of the resulting phylogenetic hypotheses through various combinations of the 4 functionally separate datasets. Increasing the number of datasets in combined analyses led to increased support for clades, sometimes substantially. We find that the tribe Argynnini is a well-supported, robust, monophyletic clade with the following internal subtribal structure: (Euptoietina (Yrameina (Boloriina Argynnina))). Our analyses support the classification of argynnine species into six robust and stable genera: Euptoieta, Yramea, Boloria, Issoria, Brenthis and Argynnis. We suggest that for moderate amounts of data, a total evidence approach is always best.},
  Keywords = {mitochondrial-DNA sequences higher-level phylogeny heliconius butterflies hawaiian drosophila characters evolution support gene incongruence congruence},
  Year = {2006}
}
• [2006,article] bibtex
  Y. Yasukochi, L. A. Ashakumary, K. Baba, A. Yoshido, and K. Sahara, "A second-generation integrated map of the silkworm reveals synteny and conserved gene order between lepidopteran insects," Genetics, vol. 173, iss. 3, pp. 1319-1328, 2006.
  @article{ Author = {Yasukochi, Y. and Ashakumary, L. A. and Baba, K. and Yoshido, A. and Sahara, K.},
  Title = {A second-generation integrated map of the silkworm reveals synteny and conserved gene order between lepidopteran insects},
  Journal = {Genetics},
  Volume = {173},
  Number = {3},
  Pages = {1319-1328},
  Note = {071UM Times Cited:10 Cited References Count:25},
  Abstract = {A second-generation linkage map was constructed for the silkworm, Bombyx mori, focusing on mapping Bombyx sequences appearing in public nucleotide databases and bacterial artificial chromosome (BAC) contigs. A total of 874 BAC contigs containing 5067 clones (22% of the library) were constructed by PCR-based screening with sequence-tagged sites (STSs) derived from whole-genome shotgun (WGS) sequences. A total of 523 BAC contigs, including 342 independent genes registered in public databases and 85 expressed sequence tags (ESTs), were placed onto the linkage map. We found significant synteny and conserved gene order between B. mori and a nymphalid butterfly, Heliconius melpomene, in four linkage groups (LGs), strongly suggesting that using B. mori as a reference for comparative genomics in Lepidotera is highly feasible.},
  Keywords = {bombyx-mori linkage map domesticated silkworm sequence chromosomes markers polymorphisms construction genome},
  Year = {2006}
}
• [2006,article] bibtex
  M. Joron, R. Papa, M. Beltran, N. Chamberlain, J. Mavarez, S. Baxter, M. Abanto, E. Bermingham, S. J. Humphray, J. Rogers, H. Beasley, K. Barlow, R. H. Ffrench-Constant, J. Mallet, W. O. McMillan, and C. D. Jiggins, "A conserved supergene locus controls colour pattern diversity in Heliconius butterflies," Plos Biology, vol. 4, iss. 10, pp. 1831-1840, 2006.
  @article{ Author = {Joron, M. and Papa, R. and Beltran, M. and Chamberlain, N. and Mavarez, J. and Baxter, S. and Abanto, M. and Bermingham, E. and Humphray, S. J. and Rogers, J. and Beasley, H. and Barlow, K. and Ffrench-Constant, R. H. and Mallet, J. and McMillan, W. O. and Jiggins, C. D.},
  Title = {A conserved supergene locus controls colour pattern diversity in Heliconius butterflies},
  Journal = {Plos Biology},
  Volume = {4},
  Number = {10},
  Pages = {1831-1840},
  Note = {095WC Times Cited:10 Cited References Count:54},
  Abstract = {We studied whether similar developmental genetic mechanisms are involved in both convergent and divergent evolution. Mimetic insects are known for their diversity of patterns as well as their remarkable evolutionary convergence, and they have played an important role in controversies over the respective roles of selection and constraints in adaptive evolution. Here we contrast three butterfly species, all classic examples of Mullerian mimicry. We used a genetic linkage map to show that a locus, Yb, which controls the presence of a yellow band in geographic races of Heliconius melpomene, maps precisely to the same location as the locus Cr, which has very similar phenotypic effects in its co-mimic H. erato. Furthermore, the same genomic location acts as a "supergene", determining multiple sympatric morphs in a third species, H. numata. H. numata is a species with a very different phenotypic appearance, whose many forms mimic different unrelated ithomiine butterflies in the genus Melinaea. Other unlinked colour pattern loci map to a homologous linkage group in the co-mimics H. melpomene and H. erato, but they are not involved in mimetic polymorphism in H. numata. Hence, a single region from the multilocus colour pattern architecture of H. melpomene and H. erato appears to have gained control of the entire wing-pattern variability in H. numata, presumably as a result of selection for mimetic "supergene" polymorphism without intermediates. Although we cannot at this stage confirm the homology of the loci segregating in the three species, our results imply that a conserved yet relatively unconstrained mechanism underlying pattern switching can affect mimicry in radically different ways. We also show that adaptive evolution, both convergent and diversifying, can occur by the repeated involvement of the same genomic regions.},
  Keywords = {cis-regulatory evolution genetic-linkage maps parallel evolution mullerian mimicry natural-selection papilio-dardanus computer package warning color chance caught melpomene},
  Year = {2006}
}
• [2006,article] bibtex
  D. D. Kapan, N. S. Flanagan, A. Tobler, R. Papa, R. D. Reed, J. A. Gonzalez, M. R. Restrepo, L. Martinez, K. Maldonado, C. Ritschoff, D. G. Heckel, and W. O. McMillan, "Localization of Mullerian mimicry genes on a dense linkage map of Heliconius erato," Genetics, vol. 173, iss. 2, pp. 735-757, 2006.
  @article{ Author = {Kapan, D. D. and Flanagan, N. S. and Tobler, A. and Papa, R. and Reed, R. D. and Gonzalez, J. A. and Restrepo, M. R. and Martinez, L. and Maldonado, K. and Ritschoff, C. and Heckel, D. G. and McMillan, W. O.},
  Title = {Localization of Mullerian mimicry genes on a dense linkage map of Heliconius erato},
  Journal = {Genetics},
  Volume = {173},
  Number = {2},
  Pages = {735-757},
  Note = {058FM Times Cited:11 Cited References Count:68},
  Abstract = {We report a dense genetic linkage map of Heliconius erato, a neotropical butterfly that has undergone a remarkable adaptive radiation in warningly colored mimetic wing patterns. Our study exploited natural variation segregating in a cross between H. erato etylus and H. himera to localize wing color pattern loci on a dense linkage map containing amplified fragment length polymorphisms (AFLP), microsatellites, and single-copy nuclear loci. We unambiguously identified all 20 autosomal linkage groups and the sex chromosome (Z). The map spanned a total of 1430 Haldane cM and linkage groups varied in size from 26.3 to 97.8 cM. The average distance between markers was 5.1 cM. Within this framework, we localized two major color pattern loci to narrow regions of the genome. The first gene, D, responsible for red/orange elements, had a most likely placement in a 6.7-cM region flanked by two AFLP markers on the end of a large 87.5-cM linkage group. The second locus, Sd, affects the melanic pattern on the forewing and was found within a 6.3-cM interval between flanking AFLP loci. This study complements recent linkage analysis of H. erato's comimic, H. melpomene, and forms the basis for marker-assisted physical mapping and for Studies into the comparative genetic architecture of wing-pattern mimicry in Heliconius.},
  Keywords = {pattern-formation chromosome-numbers race formation hybrid zones bombyx-mori distal-less butterflies evolution color nymphalidae},
  Year = {2006}
}
• [2006,article] bibtex
  I. Kawaguchi and A. Sasaki, "The wave speed of intergradation zone in two-species lattice Mullerian mimicry model," Journal of Theoretical Biology, vol. 243, iss. 4, pp. 594-603, 2006.
  @article{ Author = {Kawaguchi, I. and Sasaki, A.},
  Title = {The wave speed of intergradation zone in two-species lattice Mullerian mimicry model},
  Journal = {Journal of Theoretical Biology},
  Volume = {243},
  Number = {4},
  Pages = {594-603},
  Note = {115YI Times Cited:0 Cited References Count:19},
  Abstract = {A spatially explicit model is studied to analyse the movement of coupled clines in two-species Millierian mimicry system as exemplified by the comimicking helicoiine butterflies in Central-South America Heliconius erato and Heliconius melpomene. In this system, a pair of comimicking wing patterns of two species (mimicry ring) is found in a geographical region but another pair of wing patterns is found in a different geographical region. The distribution of mimicry rings thus forms a spatial mosaic in a large geographical scale, and the mechanism responsible for their stable maintenance has been a long-standing question in evolutionary biology. We here examine the speed of the movement of boundaries that divide the regions inhabited by different mimetic morphs in each comimicking species, by assuming coupled two-state stochastic cellular automatons where the flipping rate of the site occupied by a mimetic morph depends on the local density of the same morph and of the comimicking morph in the other species. The speed of cline movement shows a complex dependence on the coupling parameter between mimetic species-greater coupling of comimicking morphs between species slows down the cline movement only when the reduction in predation rate exhibits diminishing return to the increase of local mimetic morph density. The analytical predictions are confirmed by the results of Monte Carlo simulations. The speed of advance is quite different from that predicted from the conventional reaction-diffusion model, indicating that demographic stochasticity plays a critical role in determining the speed of cline movement. We also examine if the spatial heterogeneity in migration rate can stably maintain clines. (c) 2006 Elsevier Ltd. All rights reserved.},
  Keywords = {mullerian mimicry mimicry ring lattice model spatial mosaic pair-edge approximation hybrid zones shifting balance heliconius speciation adaptation evolution diversity invasion dynamics},
  Year = {2006}
}
• [2006,article] bibtex
  S. M. Kerpel, E. Soprano, and G. R. P. Moreira, "Effect of nitrogen on Passiflora suberosa L. (Passifloraceae) and consequences for larval performance and oviposition in Heliconius erato phyllis (Fabricius) (Lepidoptera : nymphalidae)," Neotropical Entomology, vol. 35, iss. 2, pp. 192-200, 2006.
  @article{ Author = {Kerpel, S. M. and Soprano, E. and Moreira, G. R. P.},
  Title = {Effect of nitrogen on Passiflora suberosa L. (Passifloraceae) and consequences for larval performance and oviposition in Heliconius erato phyllis (Fabricius) (Lepidoptera : nymphalidae)},
  Journal = {Neotropical Entomology},
  Volume = {35},
  Number = {2},
  Pages = {192-200},
  Note = {042LD Times Cited:2 Cited References Count:44},
  Abstract = {The present study examines the influence of nitrogen on growth rates, morphological and nutritional characteristics of Passiflora suberosa L., and the corresponding consequences for Heliconius erato phyllis (Fabricius) growth and oviposition. The treatments consisted of different nitrogen doses (0, 150 and 300 mg L-1) applied to the soil. Newly emerged larvae were individually reared in the laboratory (25 +/- 1 degrees C) on branches of P suberosa obtained from plants grown under the three nitrogen levels, and field collected females were submitted to oviposition choice tests. P suberosa growth rates, leaf area and intemode length increased significantly when grown with nitrogen enriched soil. P suberosa presented significantly greater leaf toughness when grown without nitrogen addition. Leaves of this species also presented lower water content and toughness. Nitrogen addition increased the concentration of nitrogen and potassium on plant leaf tissues. The greater nitrogen concentration on plants did not affect H. erato phyllis larval survival, but led to a reduction in larval development time and increased adult size. Females significantly preferred to lay eggs in plants cultivated in nitrogen enriched soil. This choice was positively correlated with larval performance, which could be a possible explanation for the maintenance of that behaviour, as well as the choice of larger branches as reported in previous studies.},
  Keywords = {leaf toughness insect-plant interaction insect development rate insect body size and plant nutritional value host-plant herbivory growth specialization caterpillars defenses insects choice},
  Year = {2006}
}
• [2006,article] bibtex
  T. N. Sherratt, "Spatial mosaic formation through frequency-dependent selection in Mullerian mimicry complexes," Journal of Theoretical Biology, vol. 240, iss. 2, pp. 165-174, 2006.
  @article{ Author = {Sherratt, T. N.},
  Title = {Spatial mosaic formation through frequency-dependent selection in Mullerian mimicry complexes},
  Journal = {Journal of Theoretical Biology},
  Volume = {240},
  Number = {2},
  Pages = {165-174},
  Note = {046RH Times Cited:1 Cited References Count:40},
  Abstract = {Although contemporary models of Mullerian mimicry have considered the movement of interfacial boundaries between two distinct mimetic forms. and even the possibility of polymorphisms in two patch systems, no model has considered how multiple forms of Mullerian mimics might evolve and be maintained over large geographical areas. A spatially explicit individual-based model for the evolution of Mullerian mimicry is presented, in which two unpalatable species are distributed over discrete cells within a regular lattice. Populations in each cell are capable of genetic drift and experience localized dispersal as well as frequency-dependent selection by predators. When each unpalatable prey species was introduced into a random cell and allowed to spread, then mimicry evolved throughout the system in the form of a spatial mosaic of phenotypes, separated by narrow "hybrid zones". The primary mechanism generating phenotypic diversity was the occasional establishment of new mutant forms in unoccupied cells and their subsequent maintenance (and spread) through frequency-dependent selection. The mean number of discrete clusters of the same morph that formed in the lattice was higher the higher the intensity of predation, and higher the lower the dispersal rate of unpalatable prey. Under certain conditions the hybrid zones moved, in a direction dependent on the curvature of their interfacial boundaries. However, the mimetic mosaics were highly stable when the intensity of predation was high and the rate of prey dispersal was low. Overall, this model highlights how a stable mosaic of different mimetic forms can evolve from a range of starting conditions through a combination of chance effects and localized frequency-dependent selection. (c) 2005 Elsevier Ltd. All rights reserved.},
  Keywords = {anti-apostatic selection polymorphism predation genetic drift heliconius heliconius butterflies warning-color shifting balance phylogenetic evidence natural-selection hybrid zones evolution hypothesis radiation movement},
  Year = {2006}
}
• [2006,article] bibtex
  N. Janz, S. Nylin, and N. Wahlberg, "Diversity begets diversity: host expansions and the diversification of plant-feeding insects," Bmc Evolutionary Biology, vol. 6, 2006.
  @article{ Author = {Janz, N. and Nylin, S. and Wahlberg, N.},
  Title = {Diversity begets diversity: host expansions and the diversification of plant-feeding insects},
  Journal = {Bmc Evolutionary Biology},
  Volume = {6},
  Pages = {-},
  Note = {015KF Times Cited:17 Cited References Count:104},
  Abstract = {Background: Plant-feeding insects make up a large part of earth's total biodiversity. While it has been shown that herbivory has repeatedly led to increased diversification rates in insects, there has been no compelling explanation for how plant-feeding has promoted speciation rates. There is a growing awareness that ecological factors can lead to rapid diversification and, as one of the most prominent features of most insect-plant interactions, specialization onto a diverse resource has often been assumed to be the main process behind this diversification. However, specialization is mainly a pruning process, and is not able to actually generate diversity by itself. Here we investigate the role of host colonizations in generating insect diversity, by testing if insect speciation rate is correlated with resource diversity. Results: By applying a variant of independent contrast analysis, specially tailored for use on questions of species richness (MacroCAIC), we show that species richness is strongly correlated with diversity of host use in the butterfly family Nymphalidae. Furthermore, by comparing the results from reciprocal sister group selection, where sister groups were selected either on the basis of diversity of host use or species richness, we find that it is likely that diversity of host use is driving species richness, rather than vice versa. Conclusion: We conclude that resource diversity is correlated with species richness in the Nymphalidae and suggest a scenario based on recurring oscillations between host expansions - the incorporation of new plants into the repertoire - and specialization, as an important driving force behind the diversification of plant-feeding insects.},
  Keywords = {phylogenetic-relationships phytophagous insects genetic differentiation nymphalidae lepidoptera heliconius butterflies evolutionary dynamics cladistic-analysis mitochondrial-DNA chemical defense range evolution},
  Year = {2006}
}
• [2006,article] bibtex
  G. Zaccardi, A. Kelber, M. P. Sison-Mangus, and A. D. Briscoe, "Color discrimination in the red range with only one long-wavelength sensitive opsin," Journal of Experimental Biology, vol. 209, iss. 10, pp. 1944-1955, 2006.
  @article{ Author = {Zaccardi, G. and Kelber, A. and Sison-Mangus, M. P. and Briscoe, A. D.},
  Title = {Color discrimination in the red range with only one long-wavelength sensitive opsin},
  Journal = {Journal of Experimental Biology},
  Volume = {209},
  Number = {10},
  Pages = {1944-1955},
  Note = {038QA Times Cited:8 Cited References Count:57},
  Abstract = {The basic precondition for color vision is the presence of at least two receptor types with different spectral sensitivities. The sensitivity of a receptor is mostly defined by the opsin-based visual pigment expressed in it. We show here, through behavioral experiments, that the nymphalid butterfly Heliconius erato, although it expresses short and medium wavelength opsins and only one long wavelength opsin, discriminates colors in the long-wavelength range (590 nm, 620 nm and 640 nm), whereas another nymphalid, Vanessa atalanta, despite having color vision, is unable to do so. In the eyes of H. erato we identified filtering pigments very close to the rhabdom which differ between ommatidia and produce the yellow and red ommatidial reflection seen under orthodromic illumination. The eyes of V. atalanta lack the filtering pigments, and reflect a homogeneous orange. We hypothesize that the filtering pigments found in the eyes of H. erato may shift the spectral sensitivity peak of the long wavelength receptors in some ommatidia towards longer wavelengths. The comparison of the signals between the two new receptor types makes color discrimination in the red range possible. To our knowledge, this is the first behavioral proof of color vision based on receptors expressing the same opsin.},
  Keywords = {color vision opsin filter pigment insect butterfly heliconius erato pieris-rapae-crucivora butterfly papilio-xuthus visual pigment absorption-spectra compound eye vision photoreceptors receptors retina green},
  Year = {2006}
}
• [2006,article] bibtex
  S. P. Mullen, "Wing pattern evolution and the origins of mimicry among North American admiral butterflies (Nymphalidae : Limenitis)," Molecular Phylogenetics and Evolution, vol. 39, iss. 3, pp. 747-758, 2006.
  @article{ Author = {Mullen, S. P.},
  Title = {Wing pattern evolution and the origins of mimicry among North American admiral butterflies (Nymphalidae : Limenitis)},
  Journal = {Molecular Phylogenetics and Evolution},
  Volume = {39},
  Number = {3},
  Pages = {747-758},
  Note = {051IR Times Cited:4 Cited References Count:68},
  Abstract = {The evolution of wing pattern diversity in butterflies has emerged as a model system for understanding the origins and maintenance of adaptive phenotypic novelty. Admiral butterflies (genus Limenitis) are an attractive system for studying wing pattern diversity because mimicry is common among the North American species and hybrid zones occur wherever mimetic and non-mimetic wing pattern races meet. However, the utility of this system has been limited because the evolutionary relationships among these butterflies remain unclear. Here I present a robust species-level phylogeny of Limenitis based on 1911 bp of two mitochondrial genes (COI and COII) and 904 bp of EF1-alpha for all five of the Nearctic species/wing pattern races, the majority of the Palearctic species, and three outgroup genera; Athyma, Moduza (Limenitidini), and Neptis (Limenitidinae: Neptini). Maximum-likelihood and Bayesian analyses indicate that the North American species are a well-supported, monophyletic lineage that is most closely related to the widespread, Palearctic, Poplar admiral (L. populi). Within North America, the Viceroy (L. archippus) is the basal lineage while the relationships among the remaining species are not well resolved. A combined maximum-likelihood analysis, however, indicates that the two western North America species (L. lorquini and L. weidemeyerii) are sister taxa and closely related to the wing pattern subspecies of the polytypic Limenitis arthemis species complex. These results are consistent with (1) an ancestral host-shift to Salicaceae by the common ancestor of the Poplar admiral and the Nearctic admiral lineage, (2) a single colonization of the Nearctic, and (3) a subsequent radiation of the North American forms leading to at least three independent origins of mimicry. (c) 2006 Elsevier Inc. All rights reserved.},
  Keywords = {mimicry wing pattern evolution host-plant shift molecular phylogeny lepidoptera limenitis admiral butterflies cricket hybrid zone natural-selection reproductive isolation astyanax butterflies heliconius-melpomene bering strait warning-color speciation lepidoptera phylogeny},
  Year = {2006}
}
• [2006,article] bibtex
  G. Zaccardi, A. Kelber, M. P. Sison-Mangus, and A. D. Briscoe, "Opsin expression in the eyes of Heliconius erato," Perception, vol. 35, pp. 142-143, 2006.
  @article{ Author = {Zaccardi, G. and Kelber, A. and Sison-Mangus, M. P. and Briscoe, A. D.},
  Title = {Opsin expression in the eyes of Heliconius erato},
  Journal = {Perception},
  Volume = {35},
  Pages = {142-143},
  Note = {Suppl. S 127PR Times Cited:0 Cited References Count:0},
  Abstract = {The requisite for colour vision is the assessment of at least two inputs (receptors) with different spectral sensitivities. The opsin-based visual pigment expressed in these receptors mostly defines their sensitivity. In a recent paper, we demonstrated through behavioural experiments that the nymphalid butterfly Heliconius erato discriminates colours in the long-wavelength range (590 nm, 620 nm, and 640 nm). Since the known L, M, and S opsins found in this species (L receptor peak sensitivity 570 nm) are not sufficient to explain this, we started to search for a second L opsin. This search was not successful. To our surprise, a new S2 opsin that is co-expressed with the already described S1 opsin was found. The in-situ hybridisation performed on the eyes of H. erato revealed that the L opsin expresses in six of the eight cells. The remaining two cells expressed S1, S2, and M opsins. Three classes of ommatidia expressed either S in both, M in both, or each opsin in one of the receptors. The distribution of these two types of ommatidia does not show any dorso-ventral differences.},
  Year = {2006}
}
• [2006,article] bibtex
  M. R. Kronforst, L. G. Young, L. M. Blume, and L. E. Gilbert, "Multilocus analyses of admixture and introgression among hybridizing Heliconius butterflies," Evolution, vol. 60, iss. 6, pp. 1254-1268, 2006.
  @article{ Author = {Kronforst, M. R. and Young, L. G. and Blume, L. M. and Gilbert, L. E.},
  Title = {Multilocus analyses of admixture and introgression among hybridizing Heliconius butterflies},
  Journal = {Evolution},
  Volume = {60},
  Number = {6},
  Pages = {1254-1268},
  Note = {062UJ Times Cited:20 Cited References Count:71},
  Abstract = {Introgressive hybridization is an important evolutionary process and new analytical methods provide substantial power to detect and quantify it. In this study we use variation in the frequency of 657 AFLP fragments and DNA sequence variation from 15 genes to measure the extent of admixture and the direction of interspecific gene flow among three Heliconius butterfly species that diverged recently as a result of natural selection for Mullerian mimicry, and which continue to hybridize. Bayesian clustering based on AFLP genotypes correctly delineated the three species and identified four H. cydno, three H. pachinus, and three H. melpomene individuals that were of mixed ancestry. Gene genealogies revealed substantial shared DNA sequence variation among all three species and coalescent simulations based on the Isolation with Migration (IM) model pointed to interspecific gene flow as its cause. The IM simulations further indicated that interspecific gene flow was significantly asymmetrical, with greater gene flow from H. pachinus into H. cydno (2Nm = 4.326) than the reverse (2Nm = 0.502), and unidirectional gene flow from H. cydno and H. pachinus into H. melpomene (2Nm = 0.294 and 0.252, respectively). These asymmetries are in the directions expected based on the genetics of wing patterning and the probability that hybrids of various phenotypes will survive and reproduce in different mimetic environments. This empirical demonstration of extensive interspecific gene flow is in contrast to a previous study which found little evidence of gene flow between another pair of hybridizing Heliconius species, H. himera and H. erato, and it highlights the critical role of natural selection in maintaining species diversity. Furthermore, these results lend support to the hypotheses that phenotypic diversification in the genus Heliconius has been fueled by introgressive hybridization and that reinforcement has driven the evolution of assortative mate preferences.},
  Keywords = {admixture gene flow heliconius hybridization introgression isolation with migration lepidoptera molecular population-genetics mitochondrial-DNA sequences mullerian mimicry natural-selection pattern-formation race formation genotype data nuclear gene evolution speciation},
  Year = {2006}
}
• [2006,article] bibtex
  A. Meyer, "Repeating patterns of mimicry," Plos Biology, vol. 4, iss. 10, pp. 1675-1677, 2006.
  @article{ Author = {Meyer, A.},
  Title = {Repeating patterns of mimicry},
  Journal = {Plos Biology},
  Volume = {4},
  Number = {10},
  Pages = {1675-1677},
  Note = {095WC Times Cited:1 Cited References Count:15},
  Keywords = {mitochondrial-DNA sequences cichlid fishes heliconius butterflies mullerian mimicry lake victoria evolution origin},
  Year = {2006}
}
• [2006,article] bibtex
  A. A. Ferreira, R. N. Garcia, and A. M. de Araujo, "Pupal melanization in Heliconius erato phyllis (Lepidoptera; Nymphalidae): genetic and environmental effects," Genetica, vol. 126, iss. 1-2, pp. 133-140, 2006.
  @article{ Author = {Ferreira, A. A. and Garcia, R. N. and de Araujo, A. M.},
  Title = {Pupal melanization in Heliconius erato phyllis (Lepidoptera; Nymphalidae): genetic and environmental effects},
  Journal = {Genetica},
  Volume = {126},
  Number = {1-2},
  Pages = {133-140},
  Note = {015SY Times Cited:0 Cited References Count:35},
  Abstract = {This paper deals with estimates of heritability and the effect of two different kinds of environment on the melanization of pupae in the butterfly Heliconius erato phyllis. The results for heritability showed greater differences when the offspring were regressed on male or female values for pupal colour. Pupal colour followed a system of discrete scores, from 2 for the light pupae to 5 for the darkest one. Estimate of the heritability (h(2)) was 0.44 when the average score of the siblings were regressed on the mid-parental value. When regressed on the male parent, h(2) was larger (0.54), being very low when regressed on the female parent (0.09). Estimate by the analysis of variance was also 0.44 when both male and female sibs were included in the analysis ( for male sibs, h(2) = 0.43 and for female sibs, 0.47). To test for the effect of the environment, each brood was divided in three, one being the control, the second subjected to a black environment when reaching the fifth instar and the third to a white environment. Those in the black environment originated dark pupae that scored 5 or 4; for those in the white environment, there was no difference with the controls. Caterpillars when entering the prepupal stage were also subjected to the black treatment ( early and late prepupal stage, respectively, for treatments called black A and black B). A significant effect was observed only when early prepupal stage was subjected to the black environment.},
  Keywords = {butterfly environmental effect heritability melanization pupal colour inachis-io l swallowtail butterflies reducing factor pieris-brassicae lutein incorporation natural-selection adult morphology color dimorphism pupation site heritability},
  Year = {2006}
}
• [2006,article] bibtex
  J. Mavarez, C. A. Salazar, E. Bermingham, C. Salcedo, C. D. Jiggins, and M. Linares, "Speciation by hybridization in Heliconius butterflies," Nature, vol. 441, iss. 7095, pp. 868-871, 2006.
  @article{ Author = {Mavarez, J. and Salazar, C. A. and Bermingham, E. and Salcedo, C. and Jiggins, C. D. and Linares, M.},
  Title = {Speciation by hybridization in Heliconius butterflies},
  Journal = {Nature},
  Volume = {441},
  Number = {7095},
  Pages = {868-871},
  Note = {052SL Times Cited:29 Cited References Count:29},
  Abstract = {Speciation is generally regarded to result from the splitting of a single lineage. An alternative is hybrid speciation, considered to be extremely rare, in which two distinct lineages contribute genes to a daughter species. Here we show that a hybrid trait in an animal species can directly cause reproductive isolation. The butterfly species Heliconius heurippa is known to have an intermediate morphology and a hybrid genome(1), and we have recreated its intermediate wing colour and pattern through laboratory crosses between H. melpomene, H. cydno and their F-1 hybrids. We then used mate preference experiments to show that the phenotype of H. heurippa reproductively isolates it from both parental species. There is strong assortative mating between all three species, and in H. heurippa the wing pattern and colour elements derived from H. melpomene and H. cydno are both critical for mate recognition by males.},
  Keywords = {homoploid hybrid speciation melpomene linnaeus mimicry evolution population selection origin cydno},
  Year = {2006}
}
• [2006,article] bibtex
  M. R. Kronforst, L. G. Young, D. D. Kapan, C. McNeely, R. J. O’Neill, and L. E. Gilbert, "Linkage of butterfly mate preference and wing color preference cue at the genomic location of wingless," Proceedings of the National Academy of Sciences of the United States of America, vol. 103, iss. 17, pp. 6575-6580, 2006.
  @article{ Author = {Kronforst, M. R. and Young, L. G. and Kapan, D. D. and McNeely, C. and O'Neill, R. J. and Gilbert, L. E.},
  Title = {Linkage of butterfly mate preference and wing color preference cue at the genomic location of wingless},
  Journal = {Proceedings of the National Academy of Sciences of the United States of America},
  Volume = {103},
  Number = {17},
  Pages = {6575-6580},
  Note = {037MM Times Cited:27 Cited References Count:54},
  Abstract = {Sexual isolation is a critical form of reproductive isolation in the early stages of animal speciation, yet little is known about the genetic basis of divergent mate preferences and preference cues in young species. Heliconius butterflies, well known for their diversity of wing color patterns, mate assortatively as a result of divergence in male preference for wing patterns. Here we show that the specific cue used by Heliconius cydno and Heliconius pachinus males to recognize conspecific females is the color of patches on the wings. in addition, male mate preference segregates with forewing color in hybrids, indicating a genetic association between the loci responsible for preference and preference cue. Quantitative trait locus mapping places a preference locus coincident with the locus that determines forewing color, which itself is perfectly linked to the wing patterning candidate gene, wingless. Furthermore, yellow-colored males of the polymorphic race H. cydno alithea prefer to court yellow females, indicating that wing color and color preference are controlled by loci that are located in an inversion or are pleiotropic effects of a single locus. Tight genetic associations between preference and preference cue, although rare, make divergence and speciation particularly likely because the effects of natural and sexual selection on one trait are transferred to the other, leading to the coordinated evolution of mate recognition. This effect of linkage on divergence is especially important in Heliconius because differentiation of wing color patterns in the genus has been driven and maintained by natural selection for Mullerian mimicry.},
  Keywords = {heliconius lepidoptera sexual isolation speciation quantitative trait loci reproductive isolation genetic-linkage heliconius-melpomene premating isolation mullerian mimicry natural-selection speciation drosophila evolution},
  Year = {2006}
}
• [2006,article] bibtex
  J. Mavarez and M. Gonzalez, "A set of microsatellite markers for Heliconius melpomene and closely related species," Molecular Ecology Notes, vol. 6, iss. 1, pp. 20-23, 2006.
  @article{ Author = {Mavarez, J. and Gonzalez, M.},
  Title = {A set of microsatellite markers for Heliconius melpomene and closely related species},
  Journal = {Molecular Ecology Notes},
  Volume = {6},
  Number = {1},
  Pages = {20-23},
  Note = {017XC Times Cited:4 Cited References Count:8},
  Abstract = {The butterflies in the genus Heliconius offer an exceptional opportunity for the study of the ecology and genetics of an adaptive radiation due to their extensive intra- and interspecific variation in wing colour patterns and mimetic associations. Here, we characterize 22 polymorphic microsatellite loci in Heliconius melpomene that have been shown to be useful for linkage mapping and population studies in this and other species. Levels of variation were high, although heterozygosity deficiencies were found in most loci, probably due to null alleles. The loci showed broad amplification success on six other species across the genus.},
  Keywords = {heliconius lepidoptera microsatellites nymphalidae butterflies mimicry},
  Year = {2006}
}
• [2006,article] bibtex
  O. Niehuis, S. H. Yen, C. M. Naumann, and B. Misof, "Higher phylogeny of zygaenid moths (Insecta : Lepidoptera) inferred from nuclear and mitochondrial sequence data and the evolution of larval cuticular cavities for chemical defence," Molecular Phylogenetics and Evolution, vol. 39, iss. 3, pp. 812-829, 2006.
  @article{ Author = {Niehuis, O. and Yen, S. H. and Naumann, C. M. and Misof, B.},
  Title = {Higher phylogeny of zygaenid moths (Insecta : Lepidoptera) inferred from nuclear and mitochondrial sequence data and the evolution of larval cuticular cavities for chemical defence},
  Journal = {Molecular Phylogenetics and Evolution},
  Volume = {39},
  Number = {3},
  Pages = {812-829},
  Note = {051IR Times Cited:5 Cited References Count:95},
  Abstract = {Zygaenid moths are capable of releasing hydrogen cyanide in their defense by enzymatic break-down of cyanoglucosides, but only larvae of chalcosiine and zygaenine moths store cyanogenic compounds in cuticular cavities and thus are able to discharge defense droplets, which effectively deter potential predators. A previously proposed phylogeny of Zygaenidae hypothesized a sister group relationship of chalcosiine and zygaenine moths because of their similar larval defense system. Not all chalcosiine taxa possess cuticular cavities, however, and a comparable defense mechanism has been reported in larvae of the zygaenoid family Heterogynidae. Considering sequence data of seven molecular loci, the present study estimates the posterior probability of phylogenetic hypotheses explaining the occurrence of larval cuticular cavities. The molecular data confirm the previous exclusion of Himantopteridae from Zygaenidae and suggest their close affinity to Somabrachyidae. The sequence data also corroborate the recently proposed exclusion of the Phaudinae from the Zygaenidae, because this subfamily is recovered in a reasonably well supported species cluster consisting of members of the families Lacturidae, Limacodidae, Himantopteridae, and Somabrachyidae. We consequently agree to raise Phaudinae to family rank. Within Zygaenidae, the subfamilies Callizygaeninae, Chalcosiinae, and Procridinae most likely constitute a monophyletic group, which is sister to the Zygaeninae. Our results imply that cuticular cavities were probably present in the larvae of the most recent common ancestor of Zygaenidae. Heterogynidae cannot be confirmed as sister taxon to this family, but appear at the very first split of the Zygaenoidea, although with poor support. The specific pattern of taxa in the molecular phylogeny showing larval cuticular cavities opens the possibility that these structures could have been already present in the most recent common ancestor of the Zygaenoidea. (c) 2006 Elsevier Inc. All rights reserved.},
  Keywords = {zygaenoidea nd1 mt trna-leu(cun) mt trna-val 16s rrna 18s rrna 28s rrna secondary structure models bayes factors doublet substitution models partition specific substitution models character evolution rna secondary structures last instar larva ribosomal-rna DNA-sequences heliconius butterflies maximum-likelihood bayesian-inference ditrysian-lepidoptera cladistic-analysis morphology},
  Year = {2006}
}
• [2006,article] bibtex
  G. M. Langham, "Rufous-tailed jacamars and aposematic butterflies: do older birds attack novel prey?," Behavioral Ecology, vol. 17, iss. 2, pp. 285-290, 2006.
  @article{ Author = {Langham, G. M.},
  Title = {Rufous-tailed jacamars and aposematic butterflies: do older birds attack novel prey?},
  Journal = {Behavioral Ecology},
  Volume = {17},
  Number = {2},
  Pages = {285-290},
  Note = {014AS Times Cited:4 Cited References Count:33},
  Abstract = {Although avian predators are thought, to drive the evolution of warning-color mimicry in butterflies, Few empirical studies directly address this assumption front the predator's perspective. Heliconius butterflies are textbook examples of Mullerian mimicry, with perhaps the most remarkable example being die Heliconius erato and Heliconius melpomene mimicry complex. Rufous-tailed jacamars, Galbula ruficauda (Galbulidae), are well-known butterfly predators and provide an excellent study organism to investigate patterns of attack behavior in warning-colored butterflies. I investigated patterns of attack behavior by presenting three aposematic butterflies to in a cage trial in Venezuela. I presented 80,jacamars with three Helicornius butterflies: all unaltered wing pattern (local morph) and two altered wing patterns (novel morphs). Twenty-one of 40 males and 8 of 40 females attacked a butterfly with a novel wing pattern. Of the morphological variables measure-cl, tail length was the only significant. predictor of attack behavior. Individuals with relatively longer tails attacked novel butterflies more frequently than shorter tailed individuals. Because tail length tended to increase between seasons, results suggest. that older birds are more likely to attack novel aposematic prey than arc young birds, contrary to the expectations; that younger adult. birds (i.e., more likely to be naive) would attack novel Heliconius more frequently than older birds. Overall results support. the role of specialized avian predators, like jacamars, as important agents in the evolution of warning-color mimicry in butterflies. and the need to investigate different age classes of birds in mimicry studies.},
  Keywords = {aposematism galbula ruficauda heliconius jacamar mullerian mimicry mullerian mimicry neotropical butterflies heliconius butterflies natural-selection warning-color tyrannidae responses palatability predators diversity},
  Year = {2006}
}
• [2006,article] bibtex
  I. R. Leal, E. Fischer, C. Kost, M. Tabarelli, and R. Wirth, "Ant protection against herbivores and nectar thieves in Passiflora coccinea flowers," Ecoscience, vol. 13, iss. 4, pp. 431-438, 2006.
  @article{ Author = {Leal, I. R. and Fischer, E. and Kost, C. and Tabarelli, M. and Wirth, R.},
  Title = {Ant protection against herbivores and nectar thieves in Passiflora coccinea flowers},
  Journal = {Ecoscience},
  Volume = {13},
  Number = {4},
  Pages = {431-438},
  Note = {124CY Times Cited:1 Cited References Count:45},
  Abstract = {In this study we describe the ant assemblage visiting both foliar and bracteal extrafloral nectaries (EFNs) of Passiflora coccinea, investigate the protective role provided by ants against both herbivores and nectar-thief butterflies, and analyze possible fitness benefits for the plant (i.e., fruit and seed production) resulting from the presence of ants. Experiments and observations were performed in two Amazonian terra firme forest sites. Twenty-two ant species were recorded visiting the EFNs of P coccinea. Camponotus aff. blandus was the most frequent and subordinate among the EFN-visiting ant species, accounting for 20% of the ant attacks on simulated herbivores (termites) and 50% on butterflies. Both the number of ant individuals and the number of species were positively correlated with the combined number of buds and flowers of R coccinea. Moreover, ant protection behaviour against termites was stronger on flowers than on leaves. Finally, seed set almost doubled among fruits originating from ant-visited flowers as compared to flowers from which ants were artificially excluded. Our results suggest a protective role of ants for flowers of P coccinea against both herbivores and nectar thieves, improving plant reproductive success.},
  Keywords = {amazon forest ant-plant interaction extrafloral nectaries flower protection nectar thieves plant fitness extra-floral nectaries extrafloral nectaries fruit-set plant defense vochysiaceae heliconius deterrence mutualism patterns},
  Year = {2006}
}
• [2005,article] bibtex
  M. Joron, "Polymorphic mimicry, microhabitat use, and sex-specific behaviour," Journal of Evolutionary Biology, vol. 18, iss. 3, pp. 547-556, 2005.
  @article{ Author = {Joron, M.},
  Title = {Polymorphic mimicry, microhabitat use, and sex-specific behaviour},
  Journal = {Journal of Evolutionary Biology},
  Volume = {18},
  Number = {3},
  Pages = {547-556},
  Note = {916PE Times Cited:3 Cited References Count:67},
  Abstract = {In order to assess the adaptive importance of microhabitat segregation for the maintenance of mimetic diversity, I explore how flight height varies between the sympatric forms of the polymorphic butterfly Heliconius numata and their respective models in the genus Melinaea. There is no evidence for vertical stratification of mimicry rings in these tiger-patterned butterflies, but males of H. numata tend to fly significantly higher than females and the Melinaea models. This difference in microhabitat preference likely results from females searching for host plants whereas males are patrolling for mates. I then present an extension of Muller's mimicry model for the case of partial behavioural or spatial segregation of sexes. The analysis suggests that sex-specific behaviours can make mimicry more beneficial, simply by reducing the effective population size participating in mimicry. The interaction between mimicry and sex-specific behaviours may therefore facilitate the evolution of polymorphism via enhanced, fine-scale local adaptation.},
  Keywords = {heliconius host search ithomiinae lepidoptera mate location melinaea microhabitat mullerian mimicry nymphalidae polymorphism mullerian mimicry neotropical butterflies natural-selection color pattern experimental manipulation heliconius butterflies differential selection genetic-polymorphism papilio polyxenes shifting balance},
  Year = {2005}
}
• [2005,article] bibtex
  A. Whinnett, A. V. Z. Brower, M. M. Lee, K. R. Willmott, and J. Mallet, "Phylogenetic utility of Tektin, a novel region for inferring systematic relationships among Lepidoptera," Annals of the Entomological Society of America, vol. 98, iss. 6, pp. 873-886, 2005.
  @article{ Author = {Whinnett, A. and Brower, A. V. Z. and Lee, M. M. and Willmott, K. R. and Mallet, J.},
  Title = {Phylogenetic utility of Tektin, a novel region for inferring systematic relationships among Lepidoptera},
  Journal = {Annals of the Entomological Society of America},
  Volume = {98},
  Number = {6},
  Pages = {873-886},
  Note = {983DD Times Cited:4 Cited References Count:48},
  Abstract = {Rapidly evolving nuclear coding sequences are highly desirable for phylogenetic studies of closely related species. Here, we investigated an 807-bp region, homologous to the testis-specific Tektin gene from Bombyx mori (L.), in 34 nymphalid butterfly taxa in the subfamilies Ithomiinae, Danainae, and Heliconiinae. Within Ithomiinae, relationships inferred from Tektin sequence data were. remarkably similar to those in trees based on combined morphological and ecological data. Partitioned Bremer analysis, with mitochondrial cytochrome oxidase I and II, and nuclear wingless and elongation factor 1-alpha sequences, revealed Tektin to have the greatest utility for inferring relationships at the genus, tribe, and subfamily levels across the studied taxa. We think Tektin will provide a useful source of molecular characters for inference of relationships among other butterflies, and perhaps among other insect taxa.},
  Keywords = {ithomiinae lepidoptera molecular phylogenetics tektin horizontal gene-transfer flagellar microtubules mitochondrial-DNA molecular systematics species trees heliconius butterflies nymphalidae population sequences tubulin},
  Year = {2005}
}
• [2005,article] bibtex
  Y. C. Golding, M. Edmunds, and A. R. Ennos, "Flight behaviour during foraging of the social wasp Vespula vulgaris (Hymenoptera : Vespidae) and four mimetic hoverfiies (Diptera : Syrphidae) Sericomyia silentis, Myathropa florea, Helophilus sp and Syrphus sp.," Journal of Experimental Biology, vol. 208, iss. 23, pp. 4523-4527, 2005.
  @article{ Author = {Golding, Y. C. and Edmunds, M. and Ennos, A. R.},
  Title = {Flight behaviour during foraging of the social wasp Vespula vulgaris (Hymenoptera : Vespidae) and four mimetic hoverfiies (Diptera : Syrphidae) Sericomyia silentis, Myathropa florea, Helophilus sp and Syrphus sp.},
  Journal = {Journal of Experimental Biology},
  Volume = {208},
  Number = {23},
  Pages = {4523-4527},
  Note = {999UA Times Cited:0 Cited References Count:38},
  Abstract = {Many hoverfly species show specific or non specific morphological resemblance to wasps (Vespula sp.) and it has been suggested that they also show similar flight behaviour. In this study we therefore compared the flight behaviour of wasps with that of four mimetic hoverflies, Sericomyia silentis, Myathropa florea, Helophilus sp. and Syrphus sp., by filming insects while they were foraging on an artificial array of flowers. Films were analysed to determine the routes taken, time spent hovering and flight speed. Of the four flies, only the non specific mimic, Syrphus, showed similar flight behaviour to the wasps; it flew more slowly, and with more roundabout routes than the other flies, hesitating before landing. These results suggest that in hoverflies there is little reason to expect strict correlation between morphological and behavioural mimicry; insects may acquire the similarities to their model more-or-less independently.},
  Keywords = {flight mimicry syrphidae behaviour locomotor mimicry heliconius butterflies apis-mellifera rain-forest kinematics aerodynamics temperature morphology mass},
  Year = {2005}
}
• [2005,article] bibtex
  A. Whinnett, K. R. Willmott, A. V. Z. Brower, F. Simpson, M. Zimmermann, G. Lamas, and J. Mallet, "Mitochondrial DNA provides an insight into the mechanisms driving diversification in the ithomiine butterfly Hyposcada anchiala (Lepidoptera : Nymphalidae : Ithomiinae)," European Journal of Entomology, vol. 102, iss. 4, pp. 633-639, 2005.
  @article{ Author = {Whinnett, A. and Willmott, K. R. and Brower, A. V. Z. and Simpson, F. and Zimmermann, M. and Lamas, G. and Mallet, J.},
  Title = {Mitochondrial DNA provides an insight into the mechanisms driving diversification in the ithomiine butterfly Hyposcada anchiala (Lepidoptera : Nymphalidae : Ithomiinae)},
  Journal = {European Journal of Entomology},
  Volume = {102},
  Number = {4},
  Pages = {633-639},
  Note = {992RK Times Cited:0 Cited References Count:31},
  Abstract = {Geographic Subspecies of several ithomiine butterflies on the lower east Andean slopes display a black and orange V melanic tiger" aposematic wing pattern that occurs from Colombia to Bolivia, while geographically adjacent lowland subspecies typically bear a coloured, "tiger" pattern. However, it is not clear whether subspecies with similar wing patterns in different regions have arisen through independent events of convergent adaptation, possibly through parapatric differentiation, or result from allopatric differentiation, as proposed by the refuge hypothesis. Here, we examine geographic patterns of divergence ill the widespread and common ithomiine butterfly Hyposcada anchiala. We present phylogenetic hypotheses for 5 subspecies of H. anchiala, based on 1567 bp mitochondrial DNA. All topologies indicated that a single switch in mimetic pattern best explained the wing patterning of the H. anchiala studied here. This finding suggests that the subspecies of H. anchiala studied here result from at least two stages of differentiation, and is consistent with a single colonisation into a novel altitudinal zone coincident with a wing pattern switch, followed by subsequent divergence within, rather than across altitudinal zones. The subspecies divergences indicated diversifications Were consistent with the Pleistocene. Furthermore, the lowland subspecies were more recently derived than the montane taxa, in contrast to predictions of the "Andean species pump" hypothesis.},
  Keywords = {mitochondrial DNA andes diversification ithomiinae hyposcada mimicry refuge hypothesis phylogenetic hypothesis heliconius butterflies species trees gene trees mimicry evolution systematics speciation sequences patterns},
  Year = {2005}
}
• [2005,article] bibtex
  A. Whinnett, M. Zimmermann, K. R. Willmott, N. Herrera, R. Mallarino, F. Simpson, M. Joron, G. Lamas, and J. Mallet, "Strikingly variable divergence times inferred across an Amazonian butterfly ‘suture zone’," Proceedings of the Royal Society B-Biological Sciences, vol. 272, iss. 1580, pp. 2525-2533, 2005.
  @article{ Author = {Whinnett, A. and Zimmermann, M. and Willmott, K. R. and Herrera, N. and Mallarino, R. and Simpson, F. and Joron, M. and Lamas, G. and Mallet, J.},
  Title = {Strikingly variable divergence times inferred across an Amazonian butterfly 'suture zone'},
  Journal = {Proceedings of the Royal Society B-Biological Sciences},
  Volume = {272},
  Number = {1580},
  Pages = {2525-2533},
  Note = {990NY Times Cited:11 Cited References Count:48},
  Abstract = {'Suture zones' are areas where hybrid and contact zones of multiple taxa are clustered. Such zones have been regarded as strong evidence for allopatric divergence by proponents of the Pleistocene forest refugia theory, a vicariance hypothesis frequently used to explain diversification in the Amazon basin. A central prediction of the refugia and other vicariance theories is that the taxa should have a common history so that divergence times should be coincident among taxa. A suture zone for Ithomiinae butterflies near Tarapoto, NE Peru, was therefore studied to examine divergence times of taxa in contact across the zone. We sequenced 161.9 bp of the mitochondrial COI/COII region in 172 individuals of 31 species from across the suture zone. Inferred divergence times differed remarkably, with divergence between some pairs of widespread species (each of which may have two or more subspecies interacting in the zone, as in the genus Melinaea) being considerably less than that between hybridizing subspecies in other genera (for instance in Oleria). Our data therefore strongly refute a simple hypothesis of simultaneous vicariance and suggest that ongoing parapatric or other modes of differentiation in continuous forest may be important in driving diversification in Amazonia.},
  Keywords = {lepidoptera mitochondrial DNA phylogeography speciation pleistocene refuge hypothesis DNA barcodes mitochondrial-DNA heliconius butterflies molecular clocks evolution lepidoptera speciation diversity patterns differentiation biogeography},
  Year = {2005}
}
• [2005,article] bibtex
  C. D. Jiggins, J. Mavarez, M. Beltran, W. O. McMillan, J. S. Johnston, and E. Bermingham, "A genetic linkage map of the mimetic butterfly Heliconius melpomene," Genetics, vol. 171, iss. 2, pp. 557-570, 2005.
  @article{ Author = {Jiggins, C. D. and Mavarez, J. and Beltran, M. and McMillan, W. O. and Johnston, J. S. and Bermingham, E.},
  Title = {A genetic linkage map of the mimetic butterfly Heliconius melpomene},
  Journal = {Genetics},
  Volume = {171},
  Number = {2},
  Pages = {557-570},
  Note = {982XF Times Cited:23 Cited References Count:44},
  Abstract = {Heliconius melpomene is a mimetic butterfly that exhibits great geographic variation in color pattern. We present here a genetic linkage map based on analysis of genetic markers in 73 individuals from a single F-2 family, offspring of a cross between H. m. cythera. from western Ecuador and H. m. melpomene from French Guiana. A novel "three-step method" is described for the analysis of dominant markers in an F,, cross, using outbred parental strains and taking advantage of the lack of crossing over in female Lepidoptera. This method is likely to prove useful for future mapping studies in outbred species with crossing over restricted to one sex, such as the Lepidoptera and Drosophila. The resulting linkage map has 21 linkage groups corresponding to the 21 chromosomes of H. melpomene and includes 219, AFLP markers, 23 microsatellites, 19 singlecopy nuclear genes, and the color pattern switch genes Yb and Sb. The marker density is high, averaging > 1/7 cM. The total map length is 1616 cM and the average chromosome length is 77 cM. The genome size of H. melpomene was estimated to be 292 Mb, giving a relationship of physical-to-map distance of 180 kb/cM. This map forms the basis for future comparative linkage analysis of color pattern evolution in Heliconius.},
  Keywords = {warning-color microsatellite loci adaptive radiation mullerian mimicry natural-selection pattern-formation bombyx-mori evolution nymphalidae lepidoptera},
  Year = {2005}
}
• [2005,article] bibtex
  A. K. Hundsdoerfer, I. J. Kitching, and M. Wink, "A molecular phylogeny of the hawkmoth genus Hyles (Lepidoptera : Sphingidae, Macroglossinae)," Molecular Phylogenetics and Evolution, vol. 35, iss. 2, pp. 442-458, 2005.
  @article{ Author = {Hundsdoerfer, A. K. and Kitching, I. J. and Wink, M.},
  Title = {A molecular phylogeny of the hawkmoth genus Hyles (Lepidoptera : Sphingidae, Macroglossinae)},
  Journal = {Molecular Phylogenetics and Evolution},
  Volume = {35},
  Number = {2},
  Pages = {442-458},
  Note = {917DO Times Cited:5 Cited References Count:63},
  Abstract = {The hawkmoth genus Hyles is one of 15 genera in the subtribe Choerocampina of the subfamily Macroglossinae. Due to a remarkable uniformity, morphological characters usually used to identify and classify Lepidoptera at the species level cannot be used in this genus. Instead, we used DNA sequences comprising about 2300 bp derived from the mitochondrial genes COX 1, COX 11, and tRNA-leucine to elucidate the phylogeny of Hyles. The results corroborate the monophyly of Hyles but conflict with previous internal classifications of the genus based on morphology. Hyles seems to have evolved in the Neotropics during the Oligocene/Eocene epochs and the molecular data (which evolved clock-like) confirm the hypothesis that it is a very young genus that radiated on a global scale rather quickly. We hypothesize its sister group to be one of the genera Deilephila, Theretra or Xylophanes. The Nearctic may have been colonized rapidly by Hyles once the land bridge formed during the Pliocene, since within this same Epoch, the invasion of the Palaearctic appears to have proceeded from the East, via the Bering route. The colonization of Australia appears to have occurred rather early in Hyles radiation, although the route is not clear. We propose that the radiation of the Hyles euphorbiae-complex s. str. (HEC) occurred as recently as the Pliocene/Pleistocene boundary and that its roots can still be reconstructed in Asia. Hyles dahlii is closely related to the HEC, but a sister group relationship to the HEC s. str. cannot be corroborated unequivocally. HEC population ranges appear to have tracked climate oscillations during the Pleistocene Ice Ages, resulting in hybridization around the Mediterranean Sea as they repeatedly intermingled. Comparison of the phylogeny with food plant affiliations leads us to hypothesize that Euphorbia monophagy evolved at least two times independently within Hyles. (c) 2005 Elsevier Inc. All rights reserved.},
  Keywords = {cox i and ii molecular phylogeny molecular clock hyles sphingidae biogeography cytochrome-oxidase-i mitochondrial-DNA butterflies lepidoptera heliconius butterflies nuclear gene evolution nymphalidae sequences likelihood congruence},
  Year = {2005}
}
• [2005,article] bibtex
  M. A. Peterson, K. J. Monsen, H. Pedersen, T. McFarland, and J. Bearden, "Direct and indirect analysis of the fitness of Chrysochus (Coleoptera : Chrysomelidae) hybrids," Biological Journal of the Linnean Society, vol. 84, iss. 2, pp. 273-286, 2005.
  @article{ Author = {Peterson, M. A. and Monsen, K. J. and Pedersen, H. and McFarland, T. and Bearden, J.},
  Title = {Direct and indirect analysis of the fitness of Chrysochus (Coleoptera : Chrysomelidae) hybrids},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {84},
  Number = {2},
  Pages = {273-286},
  Note = {896YO Times Cited:4 Cited References Count:94},
  Abstract = {The chrysomelid beetles Chrysochus auratus and C. cobaltinus form a narrow hybrid zone in western North America. We used a combination of direct and indirect analyses to examine the fitness of Chrysochus hybrids. For the direct analyses, we compared the mating frequency, longevity, fecundity and fertility of hybrid females and parentals. For the indirect approach, we tested predictions of multilocus genotype frequencies at a focal site in the hybrid zone, based on the frequencies of mating combinations during the previous generation. Hybrid females produced fewer eggs than did parentals and the eggs they produced in the lab failed to hatch, in contrast to those of parental females. In addition, contrary to predictions that 15.8% of the individuals at the focal site would have multilocus genotypes other than those expected of parentals or F-1 individuals, we found no such genotypes at this site. This hybrid zone appears to be an example of a classic tension zone, with endogenous selection against hybrid individuals. We discuss the implications of low hybrid fitness for the evolution of premating barriers in this system, and argue that the integration of direct and indirect approaches is a powerful means of assessing the relative fitness of hybrids, particularly for species in which mate choices are easy to observe in the field. (C) 2005 The Linnean Society of London.},
  Keywords = {endogenous selection fecundity hybrid zone introgression multilocus genotypes postzygotic barriers reinforcement speciation tension zone blue milkweed beetle haldanes rule reproductive isolation mitochondrial-DNA natural hybridization premating isolation heliconius-erato mate choice gene flow tell us},
  Year = {2005}
}
• [2005,article] bibtex
  C. A. Salazar, C. D. Jiggins, C. F. Arias, A. Tobler, E. Bermingham, and M. Linares, "Hybrid incompatibility is consistent with a hybrid origin of Heliconius heurippa Hewitson from its close relatives, Heliconius cydno Doubleday and Heliconius melpomene Linnaeus," Journal of Evolutionary Biology, vol. 18, iss. 2, pp. 247-256, 2005.
  @article{ Author = {Salazar, C. A. and Jiggins, C. D. and Arias, C. F. and Tobler, A. and Bermingham, E. and Linares, M.},
  Title = {Hybrid incompatibility is consistent with a hybrid origin of Heliconius heurippa Hewitson from its close relatives, Heliconius cydno Doubleday and Heliconius melpomene Linnaeus},
  Journal = {Journal of Evolutionary Biology},
  Volume = {18},
  Number = {2},
  Pages = {247-256},
  Note = {897PP Times Cited:8 Cited References Count:49},
  Abstract = {Shared ancestral variation and introgression complicates the reconstruction of phylogenetic relationships among closely related taxa. Here we use overall genomic compatibility as an alternative estimate of species relationships in a group where divergence is rapid and genetic exchange is common. Heliconius heurippa, a butterfly species endemic to Colombia, has a colour pattern genetically intermediate between H. cydno and H. melpomene: its hindwing is nearly indistinguishable from that of H. melpomene and its forewing band is an intermediate phenotype between both species. This observation has lead to the suggestion that the pattern of H. heurippa arose through hybridization. We present a genetic analysis of hybrid compatibility in crosses between the three taxa. Heliconius heurippa x H. cydno and female H. melpomene x male H. heurippa yield fertile and viable F-1 hybrids, but male H. melpomene x female H. heurippa crosses yield sterile F-1 females. In contrast, Haldane's rule has previously been detected between H. melpomene and H cydno in both directions. Therefore, H. heurippa is most closely related to H. cydno, with some evidence for introgression of genes from H. melpomene. The results are compatible with the hypothesis of a hybrid origin for H. heurippa. In addition, backcrosses using F-1 hybrid males provide evidence for a large Z(X)-chromosome effect on sterility and for recessive autosomal sterility factors as predicted by Dominance Theory.},
  Keywords = {dominance theory haldane's rule heliconius hybrid speciation lepidoptera haldanes rule postzygotic isolation reproductive isolation drosophila-simulans genetic dissection speciation butterflies evolution sterility dominance},
  Year = {2005}
}
• [2005,article] bibtex
  M. R. Kronforst, "Primers for the amplification of nuclear introns in Heliconius butterflies," Molecular Ecology Notes, vol. 5, iss. 1, pp. 158-162, 2005.
  @article{ Author = {Kronforst, M. R.},
  Title = {Primers for the amplification of nuclear introns in Heliconius butterflies},
  Journal = {Molecular Ecology Notes},
  Volume = {5},
  Number = {1},
  Pages = {158-162},
  Note = {905VJ Times Cited:5 Cited References Count:9},
  Abstract = {I described the development of degenerate polymerase chain reaction (PCR) primers for intron-containing regions of nine candidate wing patterning and pigmentation genes in Heliconius butterflies. Primers were developed by comparing sequence data from Drosophila melanogaster, Precis coenia, and a variety of other insects so they are likely to be applicable widely among the butterfly family Nymphalidae and perhaps Lepidoptera in general. The amplified regions are highly variable and. should be useful for inferring relationships among closely related species and estimating the phylogeographical and population genetic structure of individual species.},
  Keywords = {degenerate primers lepidoptera nymphalidae pigmentation wing pattern gene},
  Year = {2005}
}
• [2005,article] bibtex
  L. P. Moreno-Fonseca, S. C., B. M., B. E., C. D. Jiggins, and M. Linares, "The failure of mitochondrial DNA barcodes to provide reliable species diagnosis among Heliconius butterflies," Proc. R. Soc. Lond. B, vol. xx, p. xxx-xxx, 2005.
  @article{ Author = {Moreno-Fonseca, L. P. and Salazar C. and Béltran M. and Bermingham E. and Jiggins, C. D. and Linares, M.},
  Title = {The failure of mitochondrial DNA barcodes to provide reliable species diagnosis among Heliconius butterflies},
  Journal = {Proc. R. Soc. Lond. B},
  Volume = {xx},
  Pages = {xxx-xxx},
  Year = {2005}
}
• [2005,article] bibtex
  F. Molleman, H. W. Krenn, M. E. Van Alphen, P. M. Brakefleld, P. J. Devries, and B. J. Zwaan, "Food intake of fruit-feeding butterflies: evidence for adaptive variation in proboscis morphology," Biological Journal of the Linnean Society, vol. 86, iss. 3, pp. 333-343, 2005.
  @article{ Author = {Molleman, F. and Krenn, H. W. and Van Alphen, M. E. and Brakefleld, P. M. and Devries, P. J. and Zwaan, B. J.},
  Title = {Food intake of fruit-feeding butterflies: evidence for adaptive variation in proboscis morphology},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {86},
  Number = {3},
  Pages = {333-343},
  Note = {984ZX Times Cited:3 Cited References Count:37},
  Abstract = {Adult butterflies feed from a variety of substrates and have appropriate adaptations. We examined proboscis morphology in a community of fruit-feeding butterflies (Nymphalidae) in a tropical forest in Uganda. These data were supplemented with behavioural observations and measurements of intake rate on natural and artificial substrates. We found no sexual dimorphism in proboscis morphology even though puddling behaviour is usually performed by males. Two main feeding techniques could be distinguished on the basis of behaviour and morphology: the piercing technique, typically found in Charaxinae, and the sweeping technique employed by both Nymphalinae and Satyrinae. These techniques, distinguished in previous studies, are described in more detail and their relative efficiencies are discussed in the context of sexual dimorphism, food-choice and life history evolution. (c) 2005 The Linnean Society of London.},
  Keywords = {charaxinae intake rate lepidoptera life history nymphalinae post-ingestive behaviour puddling satyrinae tropical forest nymphalidae lepidoptera heliconius butterflies functional-morphology insecta papilionoidea systematics behavior sensilla search moths},
  Year = {2005}
}
• [2005,article] bibtex
  L. Mendoza-Cuenca and R. Macias-Ordonez, "Foraging polymorphism in Heliconius charitonia (Lepidoptera : Nymphalidae): morphological constraints and behavioural compensation," Journal of Tropical Ecology, vol. 21, pp. 407-415, 2005.
  @article{ Author = {Mendoza-Cuenca, L. and Macias-Ordonez, R.},
  Title = {Foraging polymorphism in Heliconius charitonia (Lepidoptera : Nymphalidae): morphological constraints and behavioural compensation},
  Journal = {Journal of Tropical Ecology},
  Volume = {21},
  Pages = {407-415},
  Note = {Part 4 952MY Times Cited:0 Cited References Count:64},
  Abstract = {Sexes and also within sex phenotypes. frequently differ in morphological traits associated with efficiency and performance in foraging and mating behaviours. In butterflies and other flying animals, phenotypic differences in wing size and traits associated with flight are involved in flight performance and individual fitness, but explorations of links among two or more traits and intrasexual differences are scarce. Foraging patterns were studied in a population of Heliconius charitonia butterflies having three phenotypes (females and two male phenotypes) which differ in their wing morphology and reproductive behaviour. As in previous studies, intersexual differences in foraging patterns were found; more interestingly, intrasexual differences were found between alternative male mating strategies. Using morphological and behavioural data, as well as data from previous flight analyses in Heliconius butterflies, we show that intrasexual differences may be explained by the energetic demands of each phenotype. Energetic expenditure is partially related to phenotypic variability in flight morphology and efficiency, and at least in both male phenotypes, differences may also be related to the energetic demands of alternative mating strategies.},
  Keywords = {aerodynamics alternative mating strategies body size flight performance foraging polymorphism heliconius mate-locating behaviour pollen feeding wing size flight performance locomotor mimicry nectar production amino-acids trade-off butterflies efficiency patterns size sex},
  Year = {2005}
}
• [2005,article] bibtex
  H. A. Merchan, C. D. Jiggins, and M. Linares, "A narrow Heliconius cydno (Nymphalidae; Heliconiini) hybrid zone with differences in morph sex ratios," Biotropica, vol. 37, iss. 1, pp. 119-128, 2005.
  @article{ Author = {Merchan, H. A. and Jiggins, C. D. and Linares, M.},
  Title = {A narrow Heliconius cydno (Nymphalidae; Heliconiini) hybrid zone with differences in morph sex ratios},
  Journal = {Biotropica},
  Volume = {37},
  Number = {1},
  Pages = {119-128},
  Note = {903HS Times Cited:0 Cited References Count:53},
  Abstract = {In the western cordillera of the Cauca valley, Colombia, there is a narrow hybrid zone between two races of Heliconius cydno, one of which is polymorphic. Three large broods show that most of the phenotypic variation observed can be explained by four loci of major effect, named Sb, Yb, L, and K Two further loci, G and Wo, were identified that control more minor phenotypic variation. Sb, Yb, and Wo are linked and the latter is differentially expressed between the sexes. The transition between H. c. cydnides in the north and H. c. weymeri to the south occurs over approximately 15 km. Collections from a single site near the center of the hybrid zone show that gene frequencies have been stable over the 10 yr from 1991 to 2001 and that color-pattern genes are in Hardy Weinberg equilibrium, with little evidence for linkage disequilibrium. This suggests that mating is random between color-pattern forms. Nonetheless, there is evidence for variation in the sex ratio, with parental phenotypes and the most melanic hybrid phenotypes showing a fairly even sex ratio compared to a strong male bias observed in the remaining hybrids. We hypothesize that this might be explained by differential selection between sexes and phenotypes, phenomena that could have important implications for hybrid zone analysis and the evolution of Mollerian mimicry.},
  Keywords = {aposematism cauca valley colombia heliconius cydno hybrid zones lower montane forest mullerian mimicry sex ratio warning-color neotropical butterflies natural-selection linkage disequilibrium female butterflies mimicry speciation melpomene lepidoptera evolution},
  Year = {2005}
}
• [2005,article] bibtex
  D. M. O’Brien, C. L. Boggs, and M. L. Fogel, "The amino acids used in reproduction by butterflies: A comparative study of dietary sources using compound-specific stable isotope analysis," Physiological and Biochemical Zoology, vol. 78, iss. 5, pp. 819-827, 2005.
  @article{ Author = {O'Brien, D. M. and Boggs, C. L. and Fogel, M. L.},
  Title = {The amino acids used in reproduction by butterflies: A comparative study of dietary sources using compound-specific stable isotope analysis},
  Journal = {Physiological and Biochemical Zoology},
  Volume = {78},
  Number = {5},
  Pages = {819-827},
  Note = {959FG Times Cited:5 Cited References Count:54},
  Abstract = {It is a nutritional challenge for nectar-feeding insects to meet the amino acid requirements of oviposition. Here we investigate whether egg amino acids derive from larval diet or are synthesized from nectar sugar in four species of butterfly: Colias eurytheme, Speyeria mormonia, Euphydryas chalcedona, and Heliconius charitonia. These species exhibit a range of life history and differ in degree of shared phylogeny. We use C-13 differences among plants to identify dietary sources of amino acid carbon, and we measure amino acid C-13 using compound-specific stable isotope analysis. Egg essential amino acids derived solely from the larval diet, with no evidence for metabolic carbon remodeling. Carbon in nonessential amino acids from eggs derived primarily from nectar sugars, with consistent variation in amino acid turnover. There was no relationship between the nonessential amino acids of eggs and host plants, demonstrating extensive metabolic remodeling. Differences between species in carbon turnover were reflected at the molecular level, particularly by glutamate and aspartate. Essential amino acid C-13 varied in a highly consistent pattern among larval host plants, reflecting a common isotopic "fingerprint" associated with plant biosynthesis. These data demonstrate conservative patterns of amino acid metabolism among Lepidoptera and the power of molecular stable isotope analyses for evaluating nutrient metabolism in situ.},
  Keywords = {life-history carbon nectar fractionation evolution turnover aphid metamorphosis stoichiometry biochemicals},
  Year = {2005}
}
• [2005,article] bibtex
  H. W. Krenn, J. D. Plant, and N. U. Szucsich, "Mouthparts of flower-visiting insects," Arthropod Structure & Development, vol. 34, iss. 1, pp. 1-40, 2005.
  @article{ Author = {Krenn, H. W. and Plant, J. D. and Szucsich, N. U.},
  Title = {Mouthparts of flower-visiting insects},
  Journal = {Arthropod Structure & Development},
  Volume = {34},
  Number = {1},
  Pages = {1-40},
  Note = {915NA Times Cited:9 Cited References Count:274},
  Abstract = {This review deals with the morphology and function of adult insect mouthparts modified to feed on nectar, pollen or petals. Specialization to nectar-feeding is evident in formation of proboscides of various lengths and designs. Proboscides of many Hymenoptera and Diptera function according to adhesion mechanisms that load nectar onto extensible apical mouthpart regions before fluid is conveyed along the food canal to the mouth by capillarity and suction. Predominantly suctorial proboscides evolved once in Lepidoptera, probably twice in Coleoptera, variously in some Hymenoptera and several times with similar design in Diptera. Many of them are particularly long and have sealed food tubes, specialized apical regions, new proboscis resting positions and modified feeding movements. Mouthparts of obligate pollen-feeding insects can be characterized by modified mandibles, specialized bristles for pollen manipulation and elaborate feeding movements. Often saliva is crucial for pollen retention and ingestion. In Coleoptera, intact pollen is gathered by sweeping movements of mouthparts; in Diptera, it is suspended in saliva prior to suction. Pollen is crushed by asymmetrical mandibles in aglossatan Lepidoptera and one group of basal Hymenoptera. Pollen-piercing mouthparts occur in Thysanoptera and one group of Diptera. Some butterflies and few Diptera extract nutrients from pollen by mixing it externally with saliva on their mouthparts. No mouthpart specializations to petal-feeding are reported in flower-visiting insects. (c) 2004 Elsevier Ltd. All rights reserved.},
  Keywords = {mouthparts proboscis anthophilous insects nectar pollen feeding morphology evolution vanessa-cardui nymphalidae lower hymenoptera insecta heliconius butterflies comparative morphology lepidoptera insecta bombyliidae diptera bees hymenoptera hover-flies yucca moths functional-morphology},
  Year = {2005}
}
• [2005,article] bibtex
  J. A. Randrianalijaona, P. A. R. Ramanoelina, J. R. E. Rasoarahona, and E. M. Gaydou, "Seasonal and chemotype influences on the chemical composition of Lantana camara L. – Essential oils from Madagascar," Analytica Chimica Acta, vol. 545, iss. 1, pp. 46-52, 2005.
  @article{ Author = {Randrianalijaona, J. A. and Ramanoelina, P. A. R. and Rasoarahona, J. R. E. and Gaydou, E. M.},
  Title = {Seasonal and chemotype influences on the chemical composition of Lantana camara L. - Essential oils from Madagascar},
  Journal = {Analytica Chimica Acta},
  Volume = {545},
  Number = {1},
  Pages = {46-52},
  Note = {941OY Times Cited:6 Cited References Count:28},
  Abstract = {Yellow-orange and pink-violet colours of flower plants of Lantana camara from Madagascar were studied with a focus on essential oil (EO) in order to characterize chemotype EO variability. The chemical composition of 73 samples of aerial part EO of L. camara collected each month of the year, at various location, have been characterized by gas chromatography-mass spectrometry (GC-MS). Among the 50 peaks characterized, 43 were identified. The main components changed-within the two flower colour types. EO from yellow-orange colour of flowers compared to pink-violet flowers is characterized by high content in terpenic compounds and lower amount in oxygenated compounds. Similar results were observed during seasons, showing that the chemical composition is relatively stable all year long. The components characterizing the pink-violet flower chemotype are: sabinene (9.4-11.3%), 1,8-cineole (3.7-4.6%), linalool (4.8-6.1%), beta-caryophyllene (11.3-13.6%), alpha-humulene (4.4-5.2%), beta-bisabolene (1.7-2.3%), gamma-cadinene (0. 1-0.4%), ar-curcumene (1.0-1.6%), caryophyllene oxide (1.2-0.7%) and davanone (22.6-25.9%). The components characterizing the yellow-orange flower chemotype are: sabinene (9.0-14.3%),1,8-cineole (0.8-1.0%), linalool (0.4-1.4%), beta-caryophyllene (25.8-30.8%), alpha-humulene (2.4-2.6%), beta-bisabolene (13.6-14.9%), gamma-cadinene (0.6-5.2%), ar-curcumene (0.7-2.8%), caryophyllene oxide (0. 1-0.4%) and davanone (0.0-0.6%). Chemical composition of nine L. camara industrial EO show that they are composed of mixture of yellow-orange and pink-violet colour of flower chemotypes. Davanone is found in all samples with a mean of 12.4%, linalool (5.4%) and 1,8-cineole (4.1%). For sesquiterpenes, the main are beta-caryophyllene (15.9%), beta-bisabolene (1.8%) and gamma-muurolene (1.4%). (c) 2005 Elsevier B.V. All rights reserved.},
  Keywords = {lantana camara verbenaceae chemotype davanone beta-caryophyllene beta-bisabolene sabinene linalool butterfly heliconius-melpomene responses india leaf},
  Year = {2005}
}
• [2005,article] bibtex
  J. S. Richards, J. N. Stanley, and P. C. Gregg, "Viability of cotton and canola pollen on the proboscis of Helicoverpa armigera: implications for spread of transgenes and pollination ecology," Ecological Entomology, vol. 30, iss. 3, pp. 327-333, 2005.
  @article{ Author = {Richards, J. S. and Stanley, J. N. and Gregg, P. C.},
  Title = {Viability of cotton and canola pollen on the proboscis of Helicoverpa armigera: implications for spread of transgenes and pollination ecology},
  Journal = {Ecological Entomology},
  Volume = {30},
  Number = {3},
  Pages = {327-333},
  Note = {930PN Times Cited:1 Cited References Count:38},
  Abstract = {1. Pollen can be transported thousands of kilometres by insects but its viability after long-distance transport is not known. Knowing the potential for this mechanism to cause outcrossing of transgenes from genetically modified (GM) plants is important for risk assessments. 2. The viability of pollen from cotton (Gossypium hirsutum L.) and canola (Brassica napus L.) was determined after placing it on the proboscis of Helicoverpa armigera moths for intervals of up to 32 h. Viability of both cotton and canola pollen declined at a much greater rate when in contact with the moth proboscis. Most was non-viable by 8 h compared with 16 h for control cotton pollen or 32 h for canola pollen. 3. There was no significant difference in the rate of decline of pollen viability between the five conventional cotton varieties, or between these and the one GM cotton variety used in these experiments. 4. The number of canola pollen grains remaining on the proboscis declined over time. Very few cotton pollen grains were retained on the proboscis. 5. The reduction in pollen viability during contact with the proboscis might indicate partial ingestion of the pollen via the proboscis. 6. The points above suggest that pollen is unlikely to remain attached or remain viable when carried over large distances by H. armigera. The implications for spread of pollen from transgenic plants and for pollination ecology in general are discussed.},
  Keywords = {canola cotton genetic modification helicoverpa armigera pollen viability pollination ecology proboscis transgenes heliconius butterflies nectar plants honey-bees lepidoptera noctuidae dispersal movement quality biology marker},
  Year = {2005}
}
• [2005,article] bibtex
  S. M. Kerpel and G. R. P. Moreira, "Absence of learning and local specialization on host plant selection by Heliconius erato," Journal of Insect Behavior, vol. 18, iss. 3, pp. 433-452, 2005.
  @article{ Author = {Kerpel, S. M. and Moreira, G. R. P.},
  Title = {Absence of learning and local specialization on host plant selection by Heliconius erato},
  Journal = {Journal of Insect Behavior},
  Volume = {18},
  Number = {3},
  Pages = {433-452},
  Note = {922QZ Times Cited:3 Cited References Count:44},
  Abstract = {There is considerable interspecific variation in larval host plants (Passifloraceae) used among Heliconius erato (Lepidoptera: Nymphalidae) populations. This study evaluates the role of learning and the influence of interspecific variation in host plant attributes on such local specialization in H. erato host plant choices. Experiments were conducted under laboratory, insectary, and field conditions, with the two most widely used host plants in southern Brazil (Passiflora suberosa Linnaeus and Passiflora misera Humbold, Bonpland et Kunth). Larval feeding preference and induction were evaluated through choice tests for all instars. Oviposition was evaluated in relation to host plant preference, Hopkins host-selection principle, and conditioning time (for 3, 7, 11, or 15 days). Also, oviposition choice was tested regarding density, intemode length, and presence of intact terminal bud on P. suberosa and P. misera shoots. Both larvae and adults of H. erato phyllis showed preference for P. misera compared to P. suberosa, under all conditions. Larval feeding preference could not be induced for most instars. The Hopkins effect was not detected and oviposition choice could not be conditioned. Females alternated use of host plant species as a function of variation in either density or presence of terminal buds on shoots. Thus, our data indicate host plant preference in H. erato phyllis is not learned but innate. Therefore, we concluded that variation in local use of host plant by this butterfly in southern Brazil results from qualitative and quantitative variation of the passion vine species.},
  Keywords = {heliconians passion vines induction of preference hopkins' host effect postimaginal conditioning oviposition behavior lepidoptera preferences butterflies herbivore phenology principle},
  Year = {2005}
}
• [2005,article] bibtex
  T. H. Fleming, D. Serrano, and J. Nassar, "Dynamics of a subtropical population of the zebra longwing butterfly Heliconius charithonia (Nymphalidae)," Florida Entomologist, vol. 88, iss. 2, pp. 169-179, 2005.
  @article{ Author = {Fleming, T. H. and Serrano, D. and Nassar, J.},
  Title = {Dynamics of a subtropical population of the zebra longwing butterfly Heliconius charithonia (Nymphalidae)},
  Journal = {Florida Entomologist},
  Volume = {88},
  Number = {2},
  Pages = {169-179},
  Note = {946PB Times Cited:0 Cited References Count:28},
  Abstract = {We studied the population dynamics of the zebra longwing butterfly, Heliconius charithonia (Nymphalidae), in a 0.05 ha garden in Miami, Florida, for 2 years to answer the following questions: (1) How stable is a suburban, subtropical population of this widespread neotropical butterfly? (2) What are the major factors influencing its population dynamics? (3) What are the implications of adult mobility regarding gene flow within and between fragmented urban populations of this species? A mark-recapture study indicated that adult population size averaged 59 individuals (range: 9-115 individuals). Peak numbers occurred in mid-wet season (September) in 1996 and in the late dry and early wet seasons (April through June) in 1997. Fluctuations in size of the adult population paralleled changes in biomass of the larval food plant, Passiflora incense. Population sex ratio was consistently male-biased (68% males). Reproduction occurred year-round, and parasitism by a trichogrammatid wasp killed about 50% of Heliconius eggs throughout the year. Recruitment of adults from chrysalises produced in the garden and deaths, rather than immigration and emigration, accounted for most numerical changes. Both males and females apparently adjust their home range locations in response to changes in the biomass of Passiflora plants. Females search these plants for suitable oviposition sites, and males search them for female pupae (mates). In addition to relatively high mortality from egg parasitism, fluctuations in the number of suitable oviposition sites and amount of larval food limited recruitment into the adult butterfly population. High adult mobility probably results in substantial gene flow within and between populations of this butterfly in urban south Florida.},
  Keywords = {population dynamics food limitation egg parasitism heliconius charithonia home-range behavior marked animals survival dispersal lepidoptera},
  Year = {2005}
}
• [2005,article] bibtex
  A. Papanicolaou, M. Joron, W. O. Mcmillan, M. L. Blaxter, and C. D. Jiggins, "Genomic tools and cDNA derived markers for butterflies," Molecular Ecology, vol. 14, iss. 9, pp. 2883-2897, 2005.
  @article{ Author = {Papanicolaou, A. and Joron, M. and Mcmillan, W. O. and Blaxter, M. L. and Jiggins, C. D.},
  Title = {Genomic tools and cDNA derived markers for butterflies},
  Journal = {Molecular Ecology},
  Volume = {14},
  Number = {9},
  Pages = {2883-2897},
  Note = {946LL Times Cited:16 Cited References Count:67},
  Abstract = {The Lepidoptera have long been used as examples in the study of evolution, but some questions remain difficult to resolve due to a lack of molecular genetic data. However, as technology improves, genomic tools are becoming increasingly available to tackle unanswered evolutionary questions. Here we have used expressed sequence tags (ESTs) to develop genetic markers for two Mullerian mimic species, Heliconius melpomene and Heliconius erato. In total 1363 ESTs were generated, representing 330 gene objects in H. melpomene and 431 in H. erato. User-friendly bioinformatic tools were used to construct a nonredundant database of these putative genes (available at http://www.heliconius.org), and annotate them with (BLAST) similarity searches, InterPro matches and Gene Ontology terms. This database will be continually updated with EST sequences for the Papilionideae as they become publicly available, providing a tool for gene finding in the butterflies. Alignments of the Heliconius sequences with putative homologues derived from Bombyx mori or other public data sets were used to identify conserved PCR priming sites, and develop 55 markers that can be amplified from genomic DNA in both H. erato and H. melpomene. These markers will be used for comparative linkage mapping in Heliconius and will have applications in other phylogenetic and genomic studies in the Lepidoptera.},
  Keywords = {cdna library colour pattern evolution ecological genomics heliconius marker development mimicry expressed sequence tags heliconius butterflies bombyx-mori color pattern lepidoptera drosophila evolution mimicry nymphalidae silkworm},
  Year = {2005}
}
• [2005,phdthesis] bibtex
  A. P. Papanicolaou, "Comparative genomics and genetic linkage mapping between the neotropical butterflies, Heliconius melpomene (Liinaeus) and Heliconius erato (Linnaeus)," PhD Thesis , 2005.
  @phdthesis{ Author = {Papanicolaou, A. P.},
  Title = {Comparative genomics and genetic linkage mapping between the neotropical butterflies, Heliconius melpomene (Liinaeus) and Heliconius erato (Linnaeus)},
  School = {University of Edinburgh},
  Type = {MSc},
  Year = {2005}
}
• [2005,article] bibtex
  M. Joron and Y. Iwasa, "The evolution of a Mullerian mimic in a spatially distributed community," Journal of Theoretical Biology, vol. 237, iss. 1, pp. 87-103, 2005.
  @article{ Author = {Joron, M. and Iwasa, Y.},
  Title = {The evolution of a Mullerian mimic in a spatially distributed community},
  Journal = {Journal of Theoretical Biology},
  Volume = {237},
  Number = {1},
  Pages = {87-103},
  Note = {965EB Times Cited:2 Cited References Count:73},
  Abstract = {Strong positive density-dependence should lead to a loss of diversity, but warning-colour and Mullerian mimicry systems show extraordinary levels of diversity. Here, we propose an analytical model to explore the dynamics of two forms of a Mullerian mimic in a heterogeneous environment with two alternative model species. Two connected populations of a dimorphic, chemically defended mimic are allowed to evolve and disperse. The proportions of the respective model species vary spatially. We use a nonlinear approximation of Muller's number-dependent equations to model a situation where the mortality for either form of the mimic decreases hyberbolically when its local density increases. A first non-spatial analysis confirms that the positive density-dependence makes coexistence of mimetic forms unstable in a single isolated patch, but shows that mimicry of the rarer model can be stable once established. The two-patch analysis shows that when models have different abundance in different places, local mimetic diversity in the mimic is maintained only if spatial heterogeneity is strong, or, more interestingly, if the mimic is not too strongly distasteful. Therefore, mildly toxic species can become polymorphic in a wider range of ecological settings. Spatial dynamics thus reveal a region of Mullerian polymorphism separating classical. Batesian polymorphism and Mullerian monomorphism along the mimic's palatability spectrum. Such polymorphism-palatability relationship in a spatial environment provides a parsimonious hypothesis accounting for the observed Mullerian polymorphism that does not require quasi-Batesian dynamics. While the stability of coexistence depends on all factors, only the migration rate and strength of selection appear to affect the level of diversity at the polymorphic equilibrium. Local adaptation is predicted in most polymorphic cases. These results are in very good accordance with recent empirical findings on the polymorphic butterflies Heliconius numata and H. cydno. (c) 2005 Elsevier Ltd. All rights reserved.},
  Keywords = {mullerian mimicry toxicity spatial heterogeneity density-dependence local adaptation frequency-dependent selection heliconius butterflies gene flow heterogeneous environments neotropical butterflies shifting balance warning color hybrid zones polymorphism diversity},
  Year = {2005}
}
• [2005,article] bibtex
  B. N. Danforth, C. P. Lin, and J. Fang, "How do insect nuclear ribosomal genes compare to protein-coding genes in phylogenetic utility and nucleotide substitution patterns?," Systematic Entomology, vol. 30, iss. 4, pp. 549-562, 2005.
  @article{ Author = {Danforth, B. N. and Lin, C. P. and Fang, J.},
  Title = {How do insect nuclear ribosomal genes compare to protein-coding genes in phylogenetic utility and nucleotide substitution patterns?},
  Journal = {Systematic Entomology},
  Volume = {30},
  Number = {4},
  Pages = {549-562},
  Note = {972ZF Times Cited:11 Cited References Count:88},
  Abstract = {The expanding data set on insect molecular systematics allows examination of phylogenetic performance and molecular evolution of different types of gene. Studies combining more than one gene in the same analysis allow examination of the relative contribution and performance of each gene partition and can help inform gene choice for resolving deep and/or problematic divergences. We compared results obtained from analyses of twelve insect data sets in which authors combined one or more nuclear ribosomal genes (28S and/or 18S) with one or more protein-coding genes [elongation factor-1 alpha (EF-1 alpha), histone H3, carbamoylphosphate synthetase domain (CPS domain of CAD, or rudimentary), long-wavelength rhodopsin (LW opsin), glucose-6-phosphate dehydrogenase (G(6)pd), phosphoenolpyruvate carboxykinase (PEPCK), arginine kinase, and white]. Data sets examined spanned eight orders of insects (Odonata, Ephemeroptera, Hemiptera, Coleoptera, Trichoptera, Lepidoptera, Diptera and Hymenoptera), providing a broad range of divergence times and taxonomic levels. We estimated the phylogenetic utility of the individual genes (using parsimony methods) and characterized the nucleotide substitution patterns (using Bayesian methods) to ask which type of data is preferable for phylogenetic analysis in insects. Nuclear ribosomal and protein coding genes differed little in our measures of phylogenetic performance and patterns of nucleotide substitution. We recommend combining nuclear ribosomal gene data with nuclear protein-coding gene data because each data set has distinct advantages. We do not recommend using mitochondrial genes for higher-level studies of insect phylogeny because reviewed studies demonstrate substitution patterns that lead to high levels of homoplasy.},
  Keywords = {length difference test higher-level phylogenetics 28s rdna sequences bumble bees bombus elongation factor-1-alpha dopa-decarboxylase molecular systematics bayesian-inference historical biogeography heliconius butterflies},
  Year = {2005}
}
• [2005,article] bibtex
  M. Z. Cardoso, "New localities for Heliconius besckei Menetries and Heliconius sara (Fabricius) (Lepidoptera : Nymphalidae) in Bahia, Brazil," Neotropical Entomology, vol. 34, iss. 6, pp. 1007-1008, 2005.
  @article{ Author = {Cardoso, M. Z.},
  Title = {New localities for Heliconius besckei Menetries and Heliconius sara (Fabricius) (Lepidoptera : Nymphalidae) in Bahia, Brazil},
  Journal = {Neotropical Entomology},
  Volume = {34},
  Number = {6},
  Pages = {1007-1008},
  Note = {000KT Times Cited:0 Cited References Count:3},
  Abstract = {New occurrences for two Heliconius Kluk species are reported for sites in inland Bahia, Brazil. New inland sites are reported for Heliconius sara (Fabricius), a species normally found at coastal regions. To our knowledge, Heliconius besckei Menetries, a species restricted to cool mountain habitats, is reported for Northeastern Brazil for the first time.},
  Keywords = {heliconiinae geographical distribution brazilian northeast atlantic rainforest dispersal},
  Year = {2005}
}
• [2005,article] bibtex
  S. H. Eberhard and H. W. Krenn, "Anatomy of the oral valve in nymphalid butterflies and a functional model for fluid uptake in Lepidoptera," Zoologischer Anzeiger, vol. 243, iss. 4, pp. 305-312, 2005.
  @article{ Author = {Eberhard, S. H. and Krenn, H. W.},
  Title = {Anatomy of the oral valve in nymphalid butterflies and a functional model for fluid uptake in Lepidoptera},
  Journal = {Zoologischer Anzeiger},
  Volume = {243},
  Number = {4},
  Pages = {305-312},
  Note = {944AN Times Cited:1 Cited References Count:27},
  Abstract = {The food canal of the proboscis of Lepidoptera serves for the uptake of nutrient fluids and the discharge of saliva. A valve was discovered at the entrance to the sucking pump in the head that separates these countercurrent flows in nymphalid butterflies. Three species of Nymphalidae were examined by dissections and light microscopic serial semithin sections. The sucking pump is a unit composed of three structures: (1) the oral valve, which is a projection of the epipharynx extending into the anterior cibarial lumen, (2) the expandable lumen, and (3) the posterior sphincter valve which controls influx into the oesophagus. Based on the microanatomical results, a functional model is presented to account for the uptake and swallowing of fluids and for the control of the salivary flow into the food canal of the proboscis. Dilator muscles of the sucking pump expand the lumen by pulling on the muscular dorso-anterior side. This opens the oral valve and fluid can be drawn into the lumen from the food canal of the proboscis. Circular compressor muscles which attach to both sides of the sclerotized ventro-posterior wall of the sucking pump reduce the size of the lumen; passively they close the oral valve and press fluid through the relaxed posterior sphincter opening into the oesophagus. According to this model saliva can be discharged into the food canal during the swallowing phase. The oral valve and pumping unit are similar in all studied species despite the fact that saliva presumably plays a special role in the derived pollen-feeding behaviour of one of them, viz. Heliconius melpomene. (c) 2005 Elsevier GmbH. All rights reserved.},
  Keywords = {sucking pump feeding model mouthparts food uptake nymphalidae morphology heliconius biology nectar pollen},
  Year = {2005}
}
• [2005,article] bibtex
  A. Tobler, D. Kapan, N. S. Flanagan, C. Gonzalez, E. Peterson, C. D. Jiggins, J. S. Johntson, D. G. Heckel, and W. O. McMillan, "First-generation linkage map of the warningly colored butterfly Heliconius erato," Heredity, vol. 94, iss. 4, pp. 408-417, 2005.
  @article{ Author = {Tobler, A. and Kapan, D. and Flanagan, N. S. and Gonzalez, C. and Peterson, E. and Jiggins, C. D. and Johntson, J. S. and Heckel, D. G. and McMillan, W. O.},
  Title = {First-generation linkage map of the warningly colored butterfly Heliconius erato},
  Journal = {Heredity},
  Volume = {94},
  Number = {4},
  Pages = {408-417},
  Note = {911EJ Times Cited:21 Cited References Count:52},
  Abstract = {We report the first genetic linkage map of Heliconius erato, a species that shows remarkable variation in its warningly colored wing patterns. We use crosses between H. erato and its sister species, H. himera, to place two major color pattern genes, D and Cr, on a linkage map containing AFLP, allozyme, microsatellite and single-copy nuclear loci. We identified all 21 linkage groups in an initial genetic screen of 22 progeny from an F1 female x male H. himera family. Of the 229 markers, 87 used to identify linkage groups were also informative in 35 progeny from a sibling backcross ( H. himera female x F1 male). With these, and an additional 33 markers informative in the second family, we constructed recombinational maps for 19 of the 21 linkage groups. These maps varied in length from 18.1 to 431.1 centimorgans (cM) and yielded an estimated total length of 2400 cM. The average distance between markers was 23 cM, and eight of the 19 linkage groups, including the sex chromosome (Z) and the chromosome containing the Cr locus, contained two or more codominant anchor loci. Of the three potential candidate genes mapped here, Cubitus interruptus (Ci), Decapentaplegic (Dpp) and Wingless (Wg), only Ci was linked, although loosely, to a known Heliconius color pattern locus. This work is an important first step for constructing a denser genetic map of the H. erato color pattern radiation and for a comparative genomic study of the architecture of mimicry in Heliconius butterflies.},
  Keywords = {color pattern mimicry aflp microsatellites single-copy nuclear loci adaptation bombyx-mori mullerian mimicry pattern-formation hybrid zones evolution gene nymphalidae chromosomes speciation silkworm},
  Year = {2005}
}
• [2004,article] bibtex
  D. G. Stavenga, S. Stowe, K. Siebke, J. Zeil, and K. Arikawa, "Butterfly wing colours: scale beads make white pierid wings brighter," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 271, iss. 1548, pp. 1577-1584, 2004.
  @article{ Author = {Stavenga, D. G. and Stowe, S. and Siebke, K. and Zeil, J. and Arikawa, K.},
  Title = {Butterfly wing colours: scale beads make white pierid wings brighter},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {271},
  Number = {1548},
  Pages = {1577-1584},
  Note = {842HU Times Cited:18 Cited References Count:42},
  Abstract = {The wing-scale morphologies of the pierid butterflies Pieris rapae (small white) and Delias nigrina (common jezabel), and the heliconine Heliconius melpomene are compared and related to the wing-reflectance spectra. Light scattering at the wing scales determines the wing reflectance, but when the scales contain an absorbing pigment, reflectance is suppressed in the absorption wavelength range of the pigment. The reflectance of the white wing areas of P. rapae, where the scales are studded with beads, is considerably higher than that of the white wing areas of H. melpomene, which has scales lacking beads. The beads presumably cause the distinct matt-white colour of the wings of pierids and function to increase the reflectance amplitude. This will improve the visual discrimination between conspecific males and females.},
  Keywords = {pieridae heliconius wing pigmentation reflectance colour contrast rapae-crucivora colias-eurytheme heterogeneity pigmentation interference iridescence lepidoptera reflection insects eye},
  Year = {2004}
}
• [2004,article] bibtex
  C. D. Jiggins, C. Estrada, and A. Rodrigues, "Mimicry and the evolution of premating isolation in Heliconius melpomene Linnaeus," Journal of Evolutionary Biology, vol. 17, iss. 3, pp. 680-691, 2004.
  @article{ Author = {Jiggins, C. D. and Estrada, C. and Rodrigues, A.},
  Title = {Mimicry and the evolution of premating isolation in Heliconius melpomene Linnaeus},
  Journal = {Journal of Evolutionary Biology},
  Volume = {17},
  Number = {3},
  Pages = {680-691},
  Note = {817ST Times Cited:14 Cited References Count:71},
  Abstract = {Ecological divergence can cause speciation if adaptive traits have pleiotropic effects on mate choice. In Heliconius butterflies, mimetic patterns play a role in mate detection between sister species, as well as signalling to predators. Here we show that male butterflies from four recently diverged parapatric populations of Heliconius melpomene are more likely to approach and court their own colour patterns as compared with those of other races. A few exceptions, where males were more attracted to patterns other than their own, suggest that some mimetic patterns are sub-optimal in mate choice. Genotype frequencies in hybrid zones between races of H. melpomene suggest that mating is random, so reinforcement is unlikely to have played a role in intra-specific divergence. In summary, co-evolved divergence of colour pattern and mate preference occurs rapidly and is likely the first step in Heliconius speciation.},
  Keywords = {lepidoptera mate recognition sensory bias sexual selection speciation female mate choice sexual selection sympatric speciation mullerian mimicry reproductive isolation adaptive radiation relative abundance natural-selection hybrid sterility host races},
  Year = {2004}
}
• [2004,article] bibtex
  N. S. Flanagan, A. Tobler, A. Davison, O. G. Pybus, D. D. Kapan, S. Planas, M. Linares, D. Heckel, and W. O. McMillan, "Historical demography of Mullerian mimicry in the neotropical Heliconius butterflies," Proceedings of the National Academy of Sciences of the United States of America, vol. 101, iss. 26, pp. 9704-9709, 2004.
  @article{ Author = {Flanagan, N. S. and Tobler, A. and Davison, A. and Pybus, O. G. and Kapan, D. D. and Planas, S. and Linares, M. and Heckel, D. and McMillan, W. O.},
  Title = {Historical demography of Mullerian mimicry in the neotropical Heliconius butterflies},
  Journal = {Proceedings of the National Academy of Sciences of the United States of America},
  Volume = {101},
  Number = {26},
  Pages = {9704-9709},
  Note = {834IU Times Cited:19 Cited References Count:30},
  Abstract = {We compare the historical demographies of two Mullerian comimetic butterfly species: Heliconius erato and Heliconius melpomene. These species show an extensive parallel geographic divergence in their aposematic wing phenotypes. Recent studies suggest that this coincident mosaic results from simultaneous demographic processes shaped by extrinsic forces over Pleistocene climate fluctuations. However, DNA sequence variation at two rapidly evolving unlinked nuclear loci, Mannose phosphate isomerase (Mpi) and Triose phosphate isomerase (Tpi), show that the comimetic species have quite different quaternary demographies. In H. erato, despite ongoing lineage sorting across the Andes, nuclear genealogical estimates showed little geographical structure, suggesting high historical gene flow. Coalescent-based demographic analysis revealed population growth since the Pliocene period. Although these patterns suggest vicariant population subdivision associated with the Andean orogeny, they are not consistent with hypotheses of Pleistocene population fragmentation facilitating allopatric wing phenotype radiation in H. erato. In contrast, nuclear genetic diversity, theta, in H. melpomene was reduced relative to its comimic and revealed three phylogeographical clades. The pattern of coalescent events within regional clades was most consistent with population growth in relatively isolated populations after a recent period of restricted population size. These different demographic histories suggest that the wing-pattern radiations were not coincident in the two species. Instead, larger effective population size (N-e) in H. erato, together with profound population change in H. melpomene, supports an earlier hypothesis that H. erato diversified first as the model species of this remarkable mimetic association.},
  Keywords = {mitochondrial-DNA sequences natural-selection race formation hybrid zone speciation divergence diversity evolution refugia system},
  Year = {2004}
}
• [2004,article] bibtex
  B. Howarth, M. Edmunds, and F. Gilbert, "Does the abundance of hoverfly (syrphidae) mimics depend on the numbers of their hymenopteran models?," Evolution, vol. 58, iss. 2, pp. 367-375, 2004.
  @article{ Author = {Howarth, B. and Edmunds, M. and Gilbert, F.},
  Title = {Does the abundance of hoverfly (syrphidae) mimics depend on the numbers of their hymenopteran models?},
  Journal = {Evolution},
  Volume = {58},
  Number = {2},
  Pages = {367-375},
  Note = {802WS Times Cited:6 Cited References Count:49},
  Abstract = {We tested the prediction that, if hoverflies are Batesian mimics, this may extend to behavioral mimicry such that their numerical abundance at each hour of the day (the daily activity pattern) is related to the numbers of their hymenopteran models. After accounting for site, season, microclimatic responses, and general hoverfly abundance at three sites in northwestern England, the residual numbers of mimics were significantly correlated positively with their models nine times of 17. Sixteen of 17 relationships were positive, itself a highly significant nonrandom pattern. Several eristatine flies showed significant relationships with honeybees even though some of them mimic wasps or bumblebees, perhaps reflecting an ancestral resemblance to honeybees. There was no evidence that good and poor mimics differed in their daily activity pattern relationships with models. However, the common mimics showed significant activity pattern relationships with their models, whereas the rarer mimics did not. We conclude that many hoverflies show behavioral mimicry of their hymenopteran models.},
  Keywords = {batesian mimicry generalized linear model analysis hoverflies hymenoptera syrphidae vine butterflies heliconius locomotor mimicry imperfect mimicry apis-mellifera evolution diptera flight discrimination behavior habitat},
  Year = {2004}
}
• [2004,article] bibtex
  M. M. Hay-Roe and R. W. Mankin, "Wing-click sounds of Heliconius cydno alithea (Nymphalidae : Heliconiinae) butterflies," Journal of Insect Behavior, vol. 17, iss. 3, pp. 329-335, 2004.
  @article{ Author = {Hay-Roe, M. M. and Mankin, R. W.},
  Title = {Wing-click sounds of Heliconius cydno alithea (Nymphalidae : Heliconiinae) butterflies},
  Journal = {Journal of Insect Behavior},
  Volume = {17},
  Number = {3},
  Pages = {329-335},
  Note = {828GB J INSECT BEHAV},
  Abstract = {Field-collected Heliconius cydno Doubleday females were observed producing audible wing clicks during encounters between conspecifics in greenhouses in a large insectary during the day and at roosting time. Occasionally, these females also were observed producing sounds in aggressive encounters with females of a close relative, H. erato (L). However, the wing- clicks were not observed subsequently from first-generation adults born in the greenhouses. The sounds were produced in short trains of 3-10 wing-clicks at the rate of T 10 clicks/s. The individual clicks had a mean duration of 1.48 ms and a broad frequency spectrum, with a peak near 1075 Hz. This peak lies near the 1200-Hz frequency of maximal sensitivity measured previously for auditory neurons of H. erato. The production of these previously unreported sounds suggests that wing clicks may play a role in both intra- and interspecific communication among Heliconius species.},
  Year = {2004}
}
• [2004,article] bibtex
  C. W. Dick, D. W. Roubik, K. F. Gruber, and E. Bermingham, "Long-distance gene flow and cross-Andean dispersal of lowland rainforest bees (Apidae : Euglossini) revealed by comparative mitochondrial DNA phylogeography," Molecular Ecology, vol. 13, iss. 12, pp. 3775-3785, 2004.
  @article{ Author = {Dick, C. W. and Roubik, D. W. and Gruber, K. F. and Bermingham, E.},
  Title = {Long-distance gene flow and cross-Andean dispersal of lowland rainforest bees (Apidae : Euglossini) revealed by comparative mitochondrial DNA phylogeography},
  Journal = {Molecular Ecology},
  Volume = {13},
  Number = {12},
  Pages = {3775-3785},
  Note = {871RC Times Cited:16 Cited References Count:68},
  Abstract = {Euglossine bees (Apidae; Euglossini) exclusively pollinate hundreds of orchid species and comprise up to 25% of bee species richness in neotropical rainforests. As one of the first studies of comparative phylogeography in a neotropical insect group, we performed a mitochondrial DNA (mtDNA)-based analysis of 14 euglossine species represented by populations sampled across the Andes and/or across the Amazon basin. The mtDNA divergences within species were consistently low; across the 12 monophyletic species the mean intraspecific divergence among haplotypes was 0.9% (range of means, 0-1.9%). The cytochrome oxidase 1 (CO1) divergence among populations separated by the Andes (N = 11 species) averaged 1.1% (range 0.0-2.0%). The mtDNA CO1 data set displayed homogeneous rates of nucleotide substitution, permitting us to infer dispersal across the cordillera long after the final Andean uplift based on arthropod molecular clocks of 1.2-1.5% divergence per million years. Gene flow across the 3000-km breadth of the Amazon basin was inferred from identical cross-Amazon haplotypes found in five species. Although mtDNA haplotypes for 12 of the 14 euglossine species were monophyletic, a reticulate CO1 phylogeny was recovered in Euglossa cognata and E. mixta, suggesting large ancestral populations and recent speciation. Reference to closely related outgroups suggested recent speciation for the majority of species. Phylogeographical structure across a broad spatial scale is weaker in euglossine bees than in any neotropical group previously examined, and may derive from a combination of Quaternary speciation, population expansion and/or long-distance gene flow.},
  Keywords = {cytochrome oxidase 1 molecular clock mtdna neotropical biogeography orchid bees tropical rainforest fresh-water fishes orchid bees costa-rican heliconius butterflies population-structure mullerian mimicry central-america biogeography sequences hymenoptera},
  Year = {2004}
}
• [2004,article] bibtex
  R. B. Srygley, "The aerodynamic costs of warning signals in palatable mimetic butterflies and their distasteful models," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 271, iss. 1539, pp. 589-594, 2004.
  @article{ Author = {Srygley, R. B.},
  Title = {The aerodynamic costs of warning signals in palatable mimetic butterflies and their distasteful models},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {271},
  Number = {1539},
  Pages = {589-594},
  Note = {808KN Times Cited:10 Cited References Count:29},
  Abstract = {Bates hypothesized that some butterfly species that are palatable gain protection from predation by appearing similar to distasteful butterflies. When undisturbed, distasteful butterflies fly slowly and in a straight line, and palatable Batesian mimics also adopt this nonchalant behaviour. When seized by predators, distasteful butterflies are defended by toxic or nauseous chemicals. Lacking chemical defences, Batesian mimics depend on flight to escape attacks. Here, I demonstrate that flight in warning-coloured mimetic butterflies and their distasteful models is more costly than in closely related non-mimetic butterflies. The increased cost is the result of differences in both wing shape and kinematics. Batesian mimics and their models slow the angular velocity of their wings to enhance the colour signal but at an aerodynamic cost. Moreover, the design for flight in Batesian mimics has an additional energetic cost over that of its models. The added cost may cause Batesian mimics to be rare, explaining a general pattern that Bates first observed.},
  Keywords = {locomotor mimicry insect flight aerodynamics mutualism communication mullerian mimicry hovering insect flight locomotor mimicry neotropical butterflies aposematic prey evolution morphology heliconius temperature predation mass},
  Year = {2004}
}
• [2004,article] bibtex
  D. Rodrigues and G. R. P. Moreira, "Seasonal variation in larval host plants and consequences for Heliconius erato (Lepidoptera : Nymphalidae) adult body size," Austral Ecology, vol. 29, iss. 4, pp. 437-445, 2004.
  @article{ Author = {Rodrigues, D. and Moreira, G. R. P.},
  Title = {Seasonal variation in larval host plants and consequences for Heliconius erato (Lepidoptera : Nymphalidae) adult body size},
  Journal = {Austral Ecology},
  Volume = {29},
  Number = {4},
  Pages = {437-445},
  Note = {843EA Times Cited:1 Cited References Count:60},
  Abstract = {Adult body size, a key life history component, varies strongly within and between Heliconius erato phyllis (Lepidoptera: Nymphalidae) populations. In the present study, we determined whether seasonal variation in adult body size is temperature related and/or determined by seasonal changes of host plants (Passifloraceae) used by the larval stage. A population of H. erato phyllis located in a Eucalyptus plantation (Barba Negra Forest, Barra do Ribeiro County, Rio Grande do Sul State, Brazil) was sampled every 45 days from March 1997 to October 1998 to quantify seasonal variation in adult body size and use of larval host plants. In the laboratory, the effects of the following factors on adult body size were quantified: (i) host plant species (Passiflora misera or Passiflora suberosa); (ii) food quantity consumed by larvae (experimentally manipulated for each passion vine species); (iii) winter and summer temperatures (15 and 25degreesC, respectively); and (iv) the interaction between host plant species and temperature. Adults emerging during summer were larger than those emerging in other seasons. Female butterflies oviposited selectively on P. misera even when the dominant passion vine was P. suberosa. They only switched from using P. misera to P. suberosa during later autumn and winter, when P. misera vines were completely defoliated. The laboratory feeding trials with both passion vines showed a strong positive association between food quantity consumed by larvae and adult size. They also confirmed that adults are larger when their larvae are reared on P. misera than on P. suberosa. Temperature during larval development had no effect on H. erato phyllis adult size. Thus, seasonal variation of H. erato phyllis adult size in a given place is primarily determined by the availability and quality of host plant species used by the larval stage.},
  Keywords = {heliconian butterflies host plant selection life history passion vines seasonality size variation temperature life-history plasticity phytophagous insects cycle regulation temperature butterflies ecology growth determinants caterpillars photoperiod},
  Year = {2004}
}
• [2004,article] bibtex
  R. D. Reed, "Evidence for Notch-mediated lateral inhibition in organizing butterfly wing scales," Development Genes and Evolution, vol. 214, iss. 1, pp. 43-46, 2004.
  @article{ Author = {Reed, R. D.},
  Title = {Evidence for Notch-mediated lateral inhibition in organizing butterfly wing scales},
  Journal = {Development Genes and Evolution},
  Volume = {214},
  Number = {1},
  Pages = {43-46},
  Note = {765UA Times Cited:8 Cited References Count:16},
  Abstract = {Here I present gene expression data that implicate a Notch-mediated lateral inhibition process in the spatial organization of butterfly wing scales. During early pupal development the receptor molecule Notch is expressed in a grid-like pattern in the wing epithelium, resulting in parallel rows of uniformly spaced cells with low Notch expression. Previous work has shown that these low-Notch cells express a homolog of the achaete-scute proneural transcription factors and develop into scales. All of these observations are consistent with the Drosophila model of Notch-mediated bristle determination and support the hypothesis that bristles and scales share an underlying patterning mechanism.},
  Keywords = {heliconius bristle achaete-scute evolution development pattern drosophila evolution protein cells},
  Year = {2004}
}
• [2004,article] bibtex
  R. D. Reed and L. E. Gilbert, "Wing venation and Distal-less expression in Heliconius butterfly wing pattern development," Development Genes and Evolution, vol. 214, iss. 12, pp. 628-634, 2004.
  @article{ Author = {Reed, R. D. and Gilbert, L. E.},
  Title = {Wing venation and Distal-less expression in Heliconius butterfly wing pattern development},
  Journal = {Development Genes and Evolution},
  Volume = {214},
  Number = {12},
  Pages = {628-634},
  Note = {884NW Times Cited:12 Cited References Count:15},
  Abstract = {Here we show that major color pattern elements of Heliconius butterfly wings develop independently of wing venation. We recovered a hybrid Heliconius displaying a mutant phenotype with a severe vein deficiency. Although this butterfly lacked most of its wing veins, the large, melanic banding patterns typical of the genus were conserved across the entire wing. The only obvious correlation between vein reduction and pigment patterns was a loss of vein-associated melanin stripes near the distal margin of the wings. We examined the expression of the eyespot-associated transcription factor Distal-less in a banded and a spotted species of Heliconius and found no obvious relationship between protein expression and the band or spot patterns typical of the genus. Together, our results suggest that the melanic bands and spots in Heliconius are unlikely to be derived from an eyespot determination system. We propose that major elements of Heliconius wing pattern formation are based primarily on a complex, whole-wing proximodistal axis system.},
  Keywords = {eyespot evolution nymphalid ground plan mimicry development evolution lepidoptera nymphalidae mimicry genes},
  Year = {2004}
}
• [2004,article] bibtex
  D. Petr and K. W. Stewart, "Comparative morphology of sensilla styloconica on the proboscis of North American Nymphalidae and other selected taxa (Lepidoptera): Systematic and ecological considerations," Transactions of the American Entomological Society, vol. 130, iss. 4, pp. 293-409, 2004.
  @article{ Author = {Petr, D. and Stewart, K. W.},
  Title = {Comparative morphology of sensilla styloconica on the proboscis of North American Nymphalidae and other selected taxa (Lepidoptera): Systematic and ecological considerations},
  Journal = {Transactions of the American Entomological Society},
  Volume = {130},
  Number = {4},
  Pages = {293-409},
  Note = {905BO Times Cited:0 Cited References Count:69},
  Abstract = {Sensilla styloconica are elongated microscopically conspicuous chemo-mechano receptors found exclusively at the tongue tip of many adult Lepidoptera. These unique proboscis sensilla were comparatively studied using the scanning electron microscope in 107 species of North American and tropical butterflies. Focus was on 76 species of North American Nymphalidae representing 45 genera and 11 subfamily groups, and 15 species of tropical Nymphalidae representing additional genera and subfamilies. Observations of adult nymphalid feeding behavior and food preference for correlation with morphological characteristics were made largely in North America and substantially in the Neotropics, where bait traps were used in conjunction with aerial netting. The tongue tips of 16 additional species representing five more butterfly families were also examined for the presence and morphological characteristics of sensilla styloconica. Nomenclature for generalized and specific types of nymphalid sensilla is proposed, with focus on the morphological features of the stylus where great variation in shape, topography, sculpturing and serration was discovered. Platyform was the predominant stylar shape represented, and aspinate (smooth) was the predominant surface topography. Other types were characterized by variable symmetry, ridging, fluting, serration and spination. Morphological features of the stylus were generally consistent for all or most species within genera and often within subfamilies, with specified exceptions. Written descriptions and SEM micrographs are presented for each species studied. The greatest total and average numbers of sensilla on a proboscis were in the subfamily Limenitidinae, with 288 and 89.9, respectively. The greatest density of sensilla in the mid-region of a proboscis tip was 158/linear mm in Megisto cymela (Satyrinae). Other sensillar characteristics presented and discussed include extent of coverage of the proboscis tip, numbers of ridges and shoulder spines, length of stylus and sensory peg and apical shoulder spine lengths. Statistical analysis revealed significant differences for six of eight numerical and measurement characteristics of sensilla styloconica tested between two distinct feeding guilds identified for the North American Nymphalidae species studied. Average number, density, extent of proboscis coverage with sensilla, and their total length, and apical shoulder spine length averages were all significantly greater in the non-nectar feeding guild than in the nectar feeding one, and may indicate adaptation for greater efficiency of their feeding by dabbing the proboscis tip on wet surfaces of various textures. The number of ridges was significantly greater in nectar feeders. No significant difference was detected in the number of shoulder spines per stylus and average sensory peg length between the feeding guilds. The aspinate condition predominated and was correlated with the nectar-feeding guild. The greater frequency of apical shoulder spines in non-nectar feeders may represent adaptation for protection of the delicate sensory pegs from mechanical abrasion during feeding on uneven or rough surfaces, or for anchoring the flexible proboscis tip to the surface during feeding. Correlation analysis indicated that the number of sensilla is positively correlated with number of ridges, number of spines, extent of linear coverage, and shoulder spine length. Seven morphological characters and a total of 19 character states of the stylus were identified in North American Nymphalidae for a preliminary cladistic analysis. Results were not considered to be meaningfully robust because of the low number of characters relative to the number of taxa used, and possibly the use of multistate characters. The ultrastructural stylar characters identified in this study should be useful for future phylogenetic analyses and inferences when included with characters from a variety of other lines of evidence for Nymphalidae and Lepidoptera. The presence of sensilla styloconica in all subfamily groups of Nymphalidae, except Danainae, largely supports Ehrlich's (1958) higher classification concept for the family; the lack of these stylate sensilla, the presence of less conspicuous sensilla in the Danainae and other characteristics are presented as further evidence that they should be reconsidered for full family status. The possibility that, in addition to their sensory function, sensilla may act as extentions for providing greater sensory reach of the proboscis for nymphalid feeding is discussed. Also discussed are the functional considerations of sensilla relative to liquid uptake capability, role in pollen feeding, and involvement in host plant selection.},
  Keywords = {butterflies lepidoptera heliconius butterflies functional-morphology insecta moth papilionoidea mouthparts family},
  Year = {2004}
}
• [2004,article] bibtex
  H. J. Megens, W. J. Van Nes, C. H. M. Van Moorsel, N. E. Pierce, and R. De Jong, "Molecular phylogeny of the Oriental butterfly genus Arhopala (Lycaenidae, Theclinae) inferred from mitochondrial and nuclear genes," Systematic Entomology, vol. 29, iss. 1, pp. 115-131, 2004.
  @article{ Author = {Megens, H. J. and Van Nes, W. J. and Van Moorsel, C. H. M. and Pierce, N. E. and De Jong, R.},
  Title = {Molecular phylogeny of the Oriental butterfly genus Arhopala (Lycaenidae, Theclinae) inferred from mitochondrial and nuclear genes},
  Journal = {Systematic Entomology},
  Volume = {29},
  Number = {1},
  Pages = {115-131},
  Note = {772NX Times Cited:3 Cited References Count:62},
  Abstract = {We present a phylogeny for a selection of species of the butterfly genus Arhopala Boisduval, 1832 based on molecular characters. We sequenced 1778 bases of the mitochondrial genes Cytochrome Oxidase 1 and 2 including tRNA(Leu), and a 393-bp fragment of the nuclear wingless gene for a total of 42 specimens of 33 species, representing all major species groups. Analyses of mtDNA and wingless genes show congruent phylogenetic signal. The phylogeny presented here confirms the monophyly of the centaurus, eumolphus, camdeo and epimuta groups and the amphimuta subgroup. It confirms close relationships between species within the agelastus group, that together with the amphimuta subgroup, centaurus and camdeo groups form a monophyletic group. However, incongruencies with previous taxonomic studies also occur; the amphimuta and silhetensis groups are not monophyletic, as is the genus Arhopala itself. One enigmatic species, A. kinabala, was evaluated further for topology and the support for basal placement of this species is due mainly to the wingless gene. However, in the Parsimony analysis, and subsequent Maximum Likelihood evaluations, certain nodes could not be resolved due to insufficient support. The mtDNA shows extreme AT bias with compositional heterogeneity at 3rd codon positions, which may result in saturation. By contrast, the wingless gene does not show compositional bias, suggesting that poor support is not due solely to saturation. The evaluation of morphological characters used in previous studies on Arhopala systematics on the molecular tree indicates that the macular pattern and the absence of tails at the hind wings show extensive homoplasy. A significant phylogenetic signal (as indicated by T-PTP tests) is present in several of these morphological characters, which are nevertheless of limited use in phylogenetic studies due to their labile nature.},
  Keywords = {cytochrome-oxidase-i heliconius butterflies nucleotide-sequence compositional bias wingless gene lepidoptera evolution DNA nymphalidae inference},
  Year = {2004}
}
• [2004,article] bibtex
  C. P. Lin and B. N. Danforth, "How do insect nuclear and mitochondrial gene substitution patterns differ? Insights from Bayesian analyses of combined datasets," Molecular Phylogenetics and Evolution, vol. 30, iss. 3, pp. 686-702, 2004.
  @article{ Author = {Lin, C. P. and Danforth, B. N.},
  Title = {How do insect nuclear and mitochondrial gene substitution patterns differ? Insights from Bayesian analyses of combined datasets},
  Journal = {Molecular Phylogenetics and Evolution},
  Volume = {30},
  Number = {3},
  Pages = {686-702},
  Note = {774AW Times Cited:57 Cited References Count:89},
  Abstract = {We analyzed 12 combined mitochondrial and nuclear gene datasets in seven orders of insects using both equal weights parsimony (to evaluate phylogenetic utility) and Bayesian methods (to investigate substitution patterns). For the Bayesian analyses we used relatively complex models.(e.g., general time reversible models with rate variation) that allowed us to quantitatively compare relative rates among genes and codon positions, patterns of rate variation among genes, and substitution patterns within genes. Our analyses indicate that nuclear and mitochondrial genes differ in a number of important ways, some of which are correlated with phylogenetic utility. First and most obviously, nuclear genes generally evolve more slowly than mitochondrial genes (except in one case), making them better markers for deep divergences. Second, nuclear genes showed universally high values of CI and (generally) contribute more to overall tree resolution than mitochondrial genes (as measured by partitioned Bremer support). Third, nuclear genes show more homogeneous patterns of among-site rate variation (higher values of alpha than mitochondrial genes). Finally, nuclear genes show more symmetrical transformation rate matrices than mitochondrial genes. The combination of low values of alpha and highly asymmetrical transformation rate matrices may explain the overall poor performance of mitochondrial genes when compared to nuclear genes in the same analysis. Our analyses indicate that some parameters are highly correlated. For example, A/T bias was positively and significantly associated with relative rate and CI was positively and significantly associated with alpha (the shape of the gamma distribution). These results provide important insights into the substitution patterns that might characterized high quality genes for phylogenetic analysis: high values of a, unbiased base composition, and symmetrical transformation rate matrices. We argue that insect molecular systematists should increasingly focus on nuclear rather than mitochondrial gene datasets because nuclear genes do not suffer from the same substitutional biases that characterize mitochondrial genes. (C) 2003 Elsevier Inc. All rights reserved.},
  Keywords = {insect phylogeny molecular evolution bayesian analysis higher-level phylogenetics large carpenter bees elongation factor-1-alpha dopa-decarboxylase molecular evolution sequence data synonymous substitution heliconius butterflies xylocopa hymenoptera lepidoptera insecta},
  Year = {2004}
}
• [2004,article] bibtex
  L. P. Moreno-Fonseca, M. Beltrán, C. Salazar, E. Bermingham, C. D. Jiggins, and M. Linares, "The status of Heliconius tristero Brower (Lepidoptera: Nymphalidae), a doubtful example of mimetic convergence between closely related passion-vine butterflies," Biological Journal of the Linnean Society, vol. In review, 2004.
  @article{ Author = {Moreno-Fonseca, L. P. and Beltrán, M. and Salazar, C. and Bermingham, E. and Jiggins, C. D. and Linares, M.},
  Title = {The status of Heliconius tristero Brower (Lepidoptera: Nymphalidae), a doubtful example of mimetic convergence between closely related passion-vine butterflies},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {In review},
  Year = {2004}
}
• [2004,article] bibtex
  G. M. Langham, "Specialized avian predators repeatedly attack novel color morphs of Heliconius butterflies," Evolution, vol. 58, iss. 12, pp. 2783-2787, 2004.
  @article{ Author = {Langham, G. M.},
  Title = {Specialized avian predators repeatedly attack novel color morphs of Heliconius butterflies},
  Journal = {Evolution},
  Volume = {58},
  Number = {12},
  Pages = {2783-2787},
  Note = {885LE Times Cited:14 Cited References Count:30},
  Abstract = {The persistence of Mullerian mimicry and geographically distinct wing patterns, as observed in many Heliconius species (Lepidoptera: Nymphalidae), is difficult to explain from a predator's perspective: predator selection against locally rare patterns must persist despite avoidance learning. Maintaining spatial color-pattern polymorphism requires local pattern avoidance, fine-scale discrimination among similar wing patterns, and repeated attacks on novel color patterns. I tested for these behaviors by presenting 80 adult rufous-tailed jacamars (Galbula ruficauda) with three morphs of Heliconius butterflies, and then presenting the same suite of butterflies to 46 of these jacamars between four and 429 days later. These trials offer the first direct evidence of the selective predator behavior required to maintain aposematic polymorphism: jacamars avoid local aposematic morphs while repeatedly attacking similar but novel morphs over time.},
  Keywords = {aposematic avian predators frequency-dependent selection hcliconius butterflies jacamars mullerian mimicry frequency-dependent predation mullerian mimicry neotropical butterflies natural-selection aposematic prey warning-color palatability tyrannidae diversity crypsis},
  Year = {2004}
}
• [2004,article] bibtex
  A. V. Z. Brower and M. M. Jeansonne, "Geographical populations and "subspecies" of new world monarch butterflies (Nymphalidae) share a recent origin and are not phylogenetically distinct," Annals of the Entomological Society of America, vol. 97, iss. 3, pp. 519-523, 2004.
  @article{ Author = {Brower, A. V. Z. and Jeansonne, M. M.},
  Title = {Geographical populations and "subspecies" of new world monarch butterflies (Nymphalidae) share a recent origin and are not phylogenetically distinct},
  Journal = {Annals of the Entomological Society of America},
  Volume = {97},
  Number = {3},
  Pages = {519-523},
  Note = {823XF Times Cited:8 Cited References Count:32},
  Abstract = {To test prior results with a more sensitive technique and larger sample sizes, we assessed genetic diversity among far-flung monarch butterfly, Danaus plexippus (L.) (Lepidoptera: Nymphalidae), populations from North and South America by using mitochondrial DNA sequences. Our new data corroborate the previously documented close genetic similarity among individuals and reveal no phylogenetic structure among populations throughout the species' New World range in North and South America. Despite this intraspecific homogeneity, the monarch is clearly distinct from its sister taxon Danaus erippus (Cramer). The evidence suggests that the monarch has colonized its current distribution in relatively recent evolutionary time. Implications for conservation and regulatory policy over interregional transfer are discussed.},
  Keywords = {mtdna danaus plexippus nymphalidae biogeography genetic diversity mitochondrial-DNA lepidoptera heliconius evolution plexippus drosophila sequences patterns},
  Year = {2004}
}
• [2004,article] bibtex
  D. W. Franks and J. Noble, "Batesian mimics influence mimicry ring evolution," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 271, iss. 1535, pp. 191-196, 2004.
  @article{ Author = {Franks, D. W. and Noble, J.},
  Title = {Batesian mimics influence mimicry ring evolution},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {271},
  Number = {1535},
  Pages = {191-196},
  Note = {764BU Times Cited:4 Cited References Count:24},
  Abstract = {Mathematical models of mimicry typically involve artificial prey species with fixed colorations or appearances; this enables a comparison of predation rates to demonstrate the level of protection a mimic might be afforded. Fruitful theoretical results have been produced using this method, but it is also useful to examine the possible evolutionary consequences of mimicry. To that end, we present individual-based evolutionary simulation models where prey colorations are free to evolve. We use the models to examine the effect of Batesian mimics on Mullerian mimics and mimicry rings. Results show that Batesian mimics can potentially incite Mullerian mimicry relationships and encourage mimicry ring convergence.},
  Keywords = {mimicry rings batesian mimicry mullerian mimicry evolutionary simulation modelling muellerian mimicry heliconius butterflies palatability spectrum dynamics},
  Year = {2004}
}
• [2004,article] bibtex
  D. M. O’Brien, C. L. Boggs, and M. L. Fogel, "Making eggs from nectar: the role of life history and dietary carbon turnover in butterfly reproductive resource allocation," Oikos, vol. 105, iss. 2, pp. 279-291, 2004.
  @article{ Author = {O'Brien, D. M. and Boggs, C. L. and Fogel, M. L.},
  Title = {Making eggs from nectar: the role of life history and dietary carbon turnover in butterfly reproductive resource allocation},
  Journal = {Oikos},
  Volume = {105},
  Number = {2},
  Pages = {279-291},
  Note = {805RJ OIKOS},
  Abstract = {The diets of many butterflies and moths change dramatically with development: from herbivory in the larvae to nectarivory in the adults. These diets are nutritionally distinct, and thus are likely to contribute differentially to egg manufacture. We examine the use of dietary resources in egg manufacture by four butterfly species with different patterns of oviposition and lifespan; three in the Nymphalidae (Euphydryas chalcedona, Speyeria mormonia and Heliconius charitonia), and one in the Pieridae (Colias eurytheme). Each species was fed two isotopically distinct adult diets based on sucrose, both of which differed from the larval hostplant in C-13 content. Egg isotopic composition was analyzed to quantify the contribution of carbon from the larval and adult diets to egg manufacture. In all four species, egg C-13 content increased to an asymptotic maximum with time, indicating that adult diet is an increasingly important source of egg carbon . The C-13 increase closely resembled that of a nectar-feeding hawkmoth, and was well-described by a model of carbon flow proposed for that species. This similarity suggests that the turnover from larval to adult dietary support of egg manufacture is conserved among nectar-feeding Lepidoptera. Species varied widely in the maximum % egg carbon that derives from the adult diet, from 44% in E. chalcedona to nearly 80% in S. mormonia. These differences were related both to the extent of oocyte provisioning prior to adult emergence, and to egg composition. A species' lifetime use of larval vs adult resources in egg manufacture reflected both the carbon turnover of the eggs and the timing of oviposition. Thus, the extent to which dietary resources are important in egg manufacture in butterflies depends on development (egg provisioning in teneral adults), behavior (timing of oviposition) and nutritional physiology (nutrient synthesis and turnover).},
  Year = {2004}
}
• [2003,article] bibtex
  S. H. Eberhard and H. W. Krenn, "Salivary glands and salivary pumps in adult Nymphalidae (Lepidoptera)," Zoomorphology, vol. 122, iss. 4, pp. 161-167, 2003.
  @article{ Author = {Eberhard, S. H. and Krenn, H. W.},
  Title = {Salivary glands and salivary pumps in adult Nymphalidae (Lepidoptera)},
  Journal = {Zoomorphology},
  Volume = {122},
  Number = {4},
  Pages = {161-167},
  Note = {744GM Times Cited:3 Cited References Count:20},
  Abstract = {The salivary glands and salivary pumps were investigated by means of dissection and serial semithin sections in order to expose the anatomy and histology of Nymphalidae in relation to feeding ecology. The paired salivary glands are tubular, they begin in the head, and extend through the thorax into the abdomen. The epithelium is a unicellular layer consisting of a single cell type. Despite the uniform composition, each salivary gland can be divided into five anatomically and histologically distinct regions. The bulbous end region of the gland lies within the abdomen and is composed of highly prismatic glandular cells with large vacuoles in their cell bodies. The tubular secretion region extends into the thorax where it forms large loops running backward and forward. It is composed of glandular cells that lack large vacuoles. The salivary duct lies in the thorax and also shows a looped formation but is composed of flat epithelial cells. The salivary reservoir begins in the prothorax and reaches the head. Its cells are hemispherical and bulge out into the large lumen of the tube. In the head the outlet tube connects the left and right halves of the salivary gland, and its epithelial cells are flat. The salivary pump lies in the head ventral to the sucking pump and leads directly into the food canal of the proboscis. It is not part of the salivary gland but is derived from the salivarium. Both the thin cuticle of the roof of the salivary pump and the thick bottom are ventrally arched. Paired muscles extend from the hypopharyngeal ridges and obviously serve as dilators for the pump. A functional interpretation of the salivary pump suggests that when not in use, the dilators are not contracted and the pump is tightly closed due to its own elasticity. When the dilator muscles repeatedly contract, the saliva is forced forward into the food canal of the proboscis. The salivary gland anatomy was found to be similar to other Lepidoptera. Furthermore, the histology of the salivary glands is identical in all examined butterflies, even in the species which exhibit specialized pollen-feeding behavior.},
  Keywords = {salivary gland salivary pump salivarium functional anatomy lepidoptera nymphalidae heliconius butterflies biology},
  Year = {2003}
}
• [2003,article] bibtex
  M. M. Hay-Roe, D. G. Boucias, A. M. Shapiro, and J. J. Becriel, "A newly discovered baculovirus induces reflex bleeding in the butterfly Heliconius himera (Nymphalidae : Heliconiinae)," Journal of Invertebrate Pathology, vol. 84, iss. 1, pp. 59-62, 2003.
  @article{ Author = {Hay-Roe, M. M. and Boucias, D. G. and Shapiro, A. M. and Becriel, J. J.},
  Title = {A newly discovered baculovirus induces reflex bleeding in the butterfly Heliconius himera (Nymphalidae : Heliconiinae)},
  Journal = {Journal of Invertebrate Pathology},
  Volume = {84},
  Number = {1},
  Pages = {59-62},
  Note = {726PD J INVERTEBR PATHOL},
  Year = {2003}
}
• [2003,article] bibtex
  N. Wahlberg, E. Weingartner, and S. Nylin, "Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera : Papilionoidea)," Molecular Phylogenetics and Evolution, vol. 28, iss. 3, pp. 473-484, 2003.
  @article{ Author = {Wahlberg, N. and Weingartner, E. and Nylin, S.},
  Title = {Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera : Papilionoidea)},
  Journal = {Molecular Phylogenetics and Evolution},
  Volume = {28},
  Number = {3},
  Pages = {473-484},
  Note = {715LY MOL PHYLOGENET EVOL},
  Abstract = {Research on the molecular systematics of higher taxa, in the butterfly family Nymphalidae (Lepidoptera) is only just beginning. Outgroup selection is difficult at the moment due to the lack of consensus on the basal relationships of the major groups in Nymphalidae. We identify four major clades in the Nymphalidae based on a cladistic analysis of one mitochondrial gene sequence (COI, I450 bp) and two nuclear gene sequences (EF-1alpha, 1064 bp, and wingless, 412-415 bp) from 54 exemplar species sampled from all currently recognized subfamilies. The COI data set was found to be highly incongruent with the nuclear data sets and a Partitioned Bremer Support analysis shows that the COI data set largely undermines support for most clades. Transitions at the third codon positions of the COI data set were highly saturated, but analyzing the combined data set with the COI third positions removed did not change the results. The major clades we found are termed the danaine clade (including Danainae), the satyrine clade (including Charaxinae, Satyrinae, Calinaginae, and Morphinae), the heliconiine clade (including Heliconiinae and Limenitidinae excluding Biblidini, Cyrestini, Pseudergolini, and Coeini) and the nymphaline clade (including Nymphalinae, Apaturinae, and Coeini, Cyrestini, Pseudergolini, and Biblidini from Limenitidinae). The heliconiine and nymphaline clades are sister groups, while the most parsimonious explanation for the combined data set places the danaine clade as the most basal large group of Nymphalidae. Our results give one of the strongest hypotheses for the subfamilial relationships within Nymphalidae. We were able to resolve the polyphyletic nature of Limenitidinae, which we recommend to be split into three subfamilies: Limenitidinae, Biblidinae, and Cyrestinae. The tribe Coeini belongs in Nymphalinae. (C) 2003 Elsevier Science (USA). All rights reserved.},
  Year = {2003}
}
• [2003,article] bibtex
  R. E. Naisbit, C. D. Jiggins, and J. Mallet, "Mimicry: developmental genes that contribute to speciation," Evolution & Development, vol. 5, iss. 3, pp. 269-280, 2003.
  @article{ Author = {Naisbit, R. E. and Jiggins, C. D. and Mallet, J.},
  Title = {Mimicry: developmental genes that contribute to speciation},
  Journal = {Evolution & Development},
  Volume = {5},
  Number = {3},
  Pages = {269-280},
  Note = {672TT Times Cited:33 Cited References Count:82},
  Abstract = {Despite renewed interest in the role of natural selection as a catalyst for the origin of species, the developmental and genetic basis of speciation remains poorly understood. Here we describe the genetics of Mullerian mimicry in Heliconius cydno and H. melpomene (Lepidoptera: Nymphalidae), sister species that recently diverged to mimic other Heliconius. This mimetic shift was a key step in their speciation, leading to pre- and postmating isolation. We identify 10 autosomal loci, half of which have major effects. At least eight appear to be homologous with genes known to control pattern differences within each species. Dominance has evolved under the influence of identifiable "modifier" loci rather than being a fixed characteristic of each locus. Epistasis is found at many levels: phenotypic interaction between specific pairs of genes, developmental canalization due to polygenic modifiers so that patterns are less sharply defined in hybrids, and overall fitness through ecological selection against nonmimetic hybrid genotypes. Most of the loci are clustered into two genomic regions or "supergenes," suggesting color pattern evolution is constrained by preexisting linked elements that may have arisen via tandem duplication rather than having been assembled by natural selection. Linkage, modifiers, and epistasis affect the strength of mimicry as a barrier to gene flow between these naturally hybridizing species and may permit introgression in genomic regions unlinked to those under disruptive selection. Mullerian mimics in Heliconius use different genetic architectures to achieve the same mimetic patterns, implying few developmental constraints. Therefore, although developmental and genomic constraints undoubtedly influence the evolutionary process, their effects are probably not strong in comparison with natural selection.},
  Keywords = {warning-color heliconius butterflies reproductive isolation natural-selection evolutionary genetics postzygotic isolation adaptive radiation pattern-formation hybrid zones body-plan},
  Year = {2003}
}
• [2003,incollection] bibtex
  L. E. Gilbert, "Adaptive novelty through introgression in Heliconius wing patterns: Evidence for shared genetic ‘tool box’ from synthetic hybrid zones and a theory of diversification," , Boggs, C. L., Watt, W. B., and Ehrlich, P. R., Eds., Chicago: Univ. of Chicago Press, 2003.
  @incollection{ Author = {Gilbert, L. E.},
  Title = {Adaptive novelty through introgression in Heliconius wing patterns: Evidence for shared genetic 'tool box' from synthetic hybrid zones and a theory of diversification},
  BookTitle = {Ecology and Evolution Taking Flight: Butterflies as Model Systems},
  Editor = {Boggs, C. L. and Watt, W. B. and Ehrlich, P.R.},
  Publisher = {Univ. of Chicago Press},
  Address = {Chicago},
  Keywords = {HELICONIUS; WING PATTERN; INSECT DEVELOPMENT},
  Year = {2003}
}
• [2003,article] bibtex
  R. D. Reed, "Gregarious oviposition and clutch size adjustment by a Heliconius butterfly," Biotropica, vol. 35, iss. 4, pp. 555-559, 2003.
  @article{ Author = {Reed, R. D.},
  Title = {Gregarious oviposition and clutch size adjustment by a Heliconius butterfly},
  Journal = {Biotropica},
  Volume = {35},
  Number = {4},
  Pages = {555-559},
  Note = {762VG Times Cited:1 Cited References Count:17},
  Abstract = {Female Heliconius hewitsoni butterflies were found to aggregate during oviposition, producing multi-parent egg clutches. This behavior occurred when host plants were locally plentiful, indicating that females chose to oviposit gregariously. Collective clutch size correlated with host growth rate and with the number of females contributing to a clutch. Eggs did not positively bias host plant growth. Collective clutch size adjustment may represent a mechanism for facilitating larval aggregation while reducing intraspecific competition.},
  Keywords = {communal oviposition intraspecific competition larval aggregation passiflora social oviposition behavior eggs},
  Year = {2003}
}
• [2003,article] bibtex
  S. Andersson and H. E. M. Dobson, "Antennal responses to floral scents in the butterfly Heliconius melpomene," Journal of Chemical Ecology, vol. 29, iss. 10, pp. 2319-2330, 2003.
  @article{ Author = {Andersson, S. and Dobson, H. E. M.},
  Title = {Antennal responses to floral scents in the butterfly Heliconius melpomene},
  Journal = {Journal of Chemical Ecology},
  Volume = {29},
  Number = {10},
  Pages = {2319-2330},
  Note = {735RR J CHEM ECOL},
  Abstract = {Floral scent, together with visual floral cues, are important signals to adult butterflies searching for food-rewarding plants. To identify which compounds in a floral scent are more attractive and, thus, of biological importance to foraging butterflies, we applied electrophysiological methods. Antennal responses of male and female adults of the tropical butterfly Heliconius melpomene L. ( Lepidoptera: Nymphalidae: Heliconiinae) to individual compounds of natural floral scents and synthetic floral scent mixtures were investigated using gas chromatography - electroantennographic detection (GC-EAD). The natural floral scents included those of two tropical plant species, Lantana camara L. ( Verbenaceae) andWarszewiczia coccinea (Vahl) Kl. (Rubiaceae), and two temperate species, Buddleja davidii Franchet (Loganiaceae) and Cirsium arvense (L.) Scop. (Asteraceae). The two synthetic floral scent mixtures contained many of the compounds found in the natural scents, but all in equal quantities. Compounds both present in relatively high abundance in the floral scents and detected exclusively in the floral parts of the plant, such as linalool, linalool oxide (furanoid) I and II, oxoisophoroneoxide, and phenylacetaldehyde, elicited the strongest antennal responses, suggesting that they may reflect adaptations by the plant to attract butterfly pollinators. However, other compounds also present in high abundance in the floral scent, but detected in the vegetative as well as floral plant parts, either elicited strong antennal responses, such as trans-beta-ocimene and benzaldehyde, or failed to elicit antennal responses, such as the sesquiterpenes beta-caryophyllene and alpha-humulene from L. camara. The most volatile monoterpene alkenes in the synthetic scent mixtures elicited only low or no responses. Furthermore, the overall antennal responses were stronger in females than in males. The findings suggest that several floral scent volatiles, especially those of exclusively floral origin, are of high biological significance to H. melpomene butterflies. These include compounds of different biosynthetic origins belonging to the benzenoids, monoterpenoids, and irregular terpenoids.},
  Year = {2003}
}
• [2003,article] bibtex
  S. Andersson and H. E. M. Dobson, "Behavioral foraging responses by the butterfly Heliconius melpomene to Lantana camara floral scent," Journal of Chemical Ecology, vol. 29, iss. 10, pp. 2303-2318, 2003.
  @article{ Author = {Andersson, S. and Dobson, H. E. M.},
  Title = {Behavioral foraging responses by the butterfly Heliconius melpomene to Lantana camara floral scent},
  Journal = {Journal of Chemical Ecology},
  Volume = {29},
  Number = {10},
  Pages = {2303-2318},
  Note = {735RR J CHEM ECOL},
  Abstract = {Floral color has been shown to influence flower selection by butterflies, but few studies have investigated the role of floral scent. In this study, adults of Heliconius melpomene L. ( Lepidoptera: Nymphalidae: Heliconiinae) were tested in two- choice bioassays to investigate their ability to distinguish floral scent of the butterfly pollinated plant Lantana camara L. ( Verbenaceae) from other plant scents. The relative importance of floral scent vs. color was also studied. Butterfly foraging behavior was measured as probing with proboscis. This probing, on floral models varying in scent and color, was timed. When given a choice of floral and vegetative scents of L. camara, newly emerged butterflies preferred floral scent, indicating an innate response to floral scents. When butterflies were conditioned to L. camara floral scent by offering the scent with yellow color and sugar water, yellow color elicited stronger feeding responses than did the floral scent. However, the floral scent of L. camara was preferred to that of the novel species Philadelphus coronarius L. (Hydrangiaceae). The floral scent of L. camara was dominated by tepenoid compounds, while that of P. coronarius by fatty acid derivatives, thus demonstrating totally different compositions. It is concluded that, while H. melpomene butterflies often use visual floral traits when selecting which flowers to visit, floral scents elicit behavioral responses that initiate and maintain foraging on flowers.},
  Year = {2003}
}
• [2003,article] bibtex
  D. M. O’Brien, C. L. Boggs, and M. L. Fogel, "Pollen feeding in the butterfly Heliconius charitonia: isotopic evidence for essential amino acid transfer from pollen to eggs," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 270, iss. 1533, pp. 2631-2636, 2003.
  @article{ Author = {O'Brien, D. M. and Boggs, C. L. and Fogel, M. L.},
  Title = {Pollen feeding in the butterfly Heliconius charitonia: isotopic evidence for essential amino acid transfer from pollen to eggs},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {270},
  Number = {1533},
  Pages = {2631-2636},
  Note = {762NW Times Cited:19 Cited References Count:36},
  Abstract = {Heliconius and Laparus butterflies exhibit a unique pollen-collecting behaviour that enhances lifespan and fecundity. The specific nutritional contribution of pollen, however, had not been previously demonstrated. We used stable isotope variation to trace the carbon flow into eggs from corn pollen provided experimentally to ovipositing female Heliconius charitonia, and to evaluate the use of isotopically contrasting nectar sugars in egg amino acids. The delta(13)C of individual amino acids from pollen, larval host plant and the eggs from experimental butterflies was measured with gas chromatography/combustion/isotope ratio mass spectrometry (GC/C/IRMS), to evaluate amino acid transfer. The delta(13)C of egg essential amino acids indicated a transfer of essential amino acids from pollen to butterfly eggs. However, the delta(13)C of non-essential amino acids reflected the isotopic composition of the artificial nectar, indicating that H. charitonia synthesizes non-essential amino acids from dietary sugars. This, to our knowledge, is the first direct demonstration of amino acid transfer from pollen to butterfly eggs, and suggests that essential amino acids in particular are a key resource for extending lifespan and fecundity in Heliconius butterflies.},
  Keywords = {heliconius pollen feeding reproductive resource allocation essential amino acids compound-specific stable isotope analysis nectar allocation patterns reproduction lepidoptera behavior adaptations nymphalidae preference resources},
  Year = {2003}
}
• [2002,article] bibtex
  R. B. Simmons and S. J. Weller, "What kind of signals do mimetic tiger moths send? A phylogenetic test of wasp mimicry systems (Lepidoptera : Arctiidae : Euchromiini)," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 269, iss. 1495, pp. 983-990, 2002.
  @article{ Author = {Simmons, R. B. and Weller, S. J.},
  Title = {What kind of signals do mimetic tiger moths send? A phylogenetic test of wasp mimicry systems (Lepidoptera : Arctiidae : Euchromiini)},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {269},
  Number = {1495},
  Pages = {983-990},
  Note = {556AW Times Cited:7 Cited References Count:53},
  Abstract = {Mimicry has been examined in field and laboratory studies of butterflies and its evolutionary dynamics have been explored in computer simulations. Phylogenetic studies examining the evolution of mimicry, however, are rare. Here, the phylogeny of wasp-mimicking tiger moths, the Sphecosoma group, was used to test evolutionary predictions of computer simulations of conventional Mullerian mimicry and quasi-Batesian mimicry dynamics. We examined whether mimetic traits evolved individually, or as suites of characters, using concentrated change tests. The phylogeny of these moth mimics revealed that individual mimetic characters were conserved, as are the three mimetic wasp forms: yellow Polybia, black Polybia and Parachartergus mimetic types. This finding was consistent with a 'supergene' control of linked loci and the Nicholson two-step model of mimicry evolution. We also used a modified permutation-tail probability approach to examine the rate of mimetic-type evolution. The observed topology, hypothetical Mullerian and Batesian scenarios, and 1000 random trees were compared using Kishino-Hasegawa tests. The observed phylogeny was more consistent with the predicted Mullerian distribution of mimetic traits than with that of a quasi-Batesian scenario. We suggest that the range of discriminatory abilities of the predator community plays a key role in shaping mimicry dynamics.},
  Keywords = {comparative biology arctiinae mullerian mimicry evolution of mimicry mullerian mimicry heliconius butterflies muellerian mimicry sequence data evolution permutation adaptation diversity predator dynamics},
  Year = {2002}
}
• [2002,article] bibtex
  P. Beldade and P. M. Brakefield, "The genetics and evo-devo of butterfly wing patterns," Nature Reviews Genetics, vol. 3, iss. 6, pp. 442-452, 2002.
  @article{ Author = {Beldade, P. and Brakefield, P. M.},
  Title = {The genetics and evo-devo of butterfly wing patterns},
  Journal = {Nature Reviews Genetics},
  Volume = {3},
  Number = {6},
  Pages = {442-452},
  Note = {558AR Times Cited:66 Cited References Count:101},
  Abstract = {Understanding how the spectacular diversity of colour patterns on butterfly wings is shaped by natural selection, and how particular pattern elements are generated, has been the focus of both evolutionary and developmental biologists. The growing field of evolutionary developmental biology has now begun to provide a link between genetic variation and the phenotypes that are produced by developmental processes and that are sorted by natural selection. Butterfly wing patterns are set to become one of the few examples of morphological diversity to be studied successfully at many levels of biological organization, and thus to yield a more complete picture of adaptive morphological evolution.},
  Keywords = {precis-coenia lepidoptera evolutionary developmental biology color pattern bicyclus-anynana seasonal polyphenism eyespot patterns dopa decarboxylase warning-color swallowtail butterflies heliconius butterflies},
  Year = {2002}
}
• [2002,article] bibtex
  W. O. McMillan, A. Monteiro, and D. D. Kapan, "Development and evolution on the wing," Trends in Ecology & Evolution, vol. 17, iss. 3, pp. 125-133, 2002.
  @article{ Author = {McMillan, W. O. and Monteiro, A. and Kapan, D. D.},
  Title = {Development and evolution on the wing},
  Journal = {Trends in Ecology & Evolution},
  Volume = {17},
  Number = {3},
  Pages = {125-133},
  Note = {527ZB Times Cited:44 Cited References Count:51},
  Abstract = {Butterfly wing patterns are more than just visually stunning examples of the evolutionary process. They are also emerging as exceptional model systems with which to link the developmental and genetic processes that generate morphological variation with the ecological and evolutionary processes that mould variation in natural populations. Work over the past few years has proceeded simultaneously on key developmental and evolutionary aspects of patterns on butterfly wings. Important clues into molecular and developmental events occurring during wing development are now available that refine our understanding of pattern formation. In addition, recent behavioural, field and molecular phylogenetic work places butterfly wing pattern change in a solid ecological and evolutionary context. There are still large gaps in our understanding, but current research priorities are well laid out and experimental methodologies are in place to address them. The challenge is to synthesize diverse research strategies into a cohesive picture of morphological evolution.},
  Keywords = {butterfly bicyclus-anynana color pattern-formation gradient formation heliconius butterflies quantitative genetics eyespot patterns imaginal disc precis-coenia wild-type mimicry},
  Year = {2002}
}
• [2002,article] bibtex
  V. Clausen, K. Frydenvang, R. Koopmann, L. B. Jorgensen, D. K. Abbiw, P. Ekpe, and J. W. Jaroszewski, "Plant analysis by butterflies: Occurrence of cyclopentenylglycines in Passifloraceae, Flacourtiaceae, and Turneraceae and discovery of the novel nonproteinogenic amino acid 2-(3 ‘-cyclopentenyl)glycine in Rinorea," Journal of Natural Products, vol. 65, iss. 4, pp. 542-547, 2002.
  @article{ Author = {Clausen, V. and Frydenvang, K. and Koopmann, R. and Jorgensen, L. B. and Abbiw, D. K. and Ekpe, P. and Jaroszewski, J. W.},
  Title = {Plant analysis by butterflies: Occurrence of cyclopentenylglycines in Passifloraceae, Flacourtiaceae, and Turneraceae and discovery of the novel nonproteinogenic amino acid 2-(3 '-cyclopentenyl)glycine in Rinorea},
  Journal = {Journal of Natural Products},
  Volume = {65},
  Number = {4},
  Pages = {542-547},
  Note = {547HV Times Cited:6 Cited References Count:53},
  Abstract = {Following records about feeding habits of nymphalid butterflies, a novel nonproteinogenic L-amino acid, (S)-2-(3'-cyclopentenyl)glycine (11), was discovered in Rinorea ilicifolia, a species where the presence of a cyclopentanoid natural product of this kind was neither known nor anticipated from the taxonomic point of view. Another novel amino acid, (2S,1'S,2'S)-2-(2'-hydroxy-3'-cyclopentenyl)glycine (12), the stereochemistry of which was determined by single-crystal X-ray diffraction, was shown to occur in species belonging to Flacourtiaceae, Passifloraceae, and Turneraceae. These species, many of which serve as hosts for nymphalid butterflies (Acraeinae, Heliconiinae, Argynninae), also produce 2-(2'-cyclopentenyl)glycine. Cyclopentenylglycines are proposed to be novel chemical recognition templates for plant-insect interactions. Ratios between the epimers of (2S)-2-(2'-cyclopentenyl)glycine, which co-occur in plants, were determined by H-1 NMR spectroscopy. Contrary to a previous report, the (2S,1'R) epimer always appears to predominate over the (2S,1'S) epimer. Stereochemical aspects of biosynthesis of natural cyclopentanoid cyanogenic glycosides are discussed in relation to these findings.},
  Keywords = {cyclopentenoid cyanohydrin glycosides heliconius species lepidoptera cyanogenic glucosides biosynthesis lotaustralin linamarin gynocardin zygaena deidaclin evolution},
  Year = {2002}
}
• [2002,article] bibtex
  K. J. A. Vanhoutte, B. J. L. Eggen, J. J. M. Janssen, and D. G. Stavenga, "Opsin cDNA sequences of a UV and green rhodopsin of the satyrine butterfly Bicyclus anynana," Insect Biochemistry and Molecular Biology, vol. 32, iss. 11, pp. 1383-1390, 2002.
  @article{ Author = {Vanhoutte, K. J. A. and Eggen, B. J. L. and Janssen, J. J. M. and Stavenga, D. G.},
  Title = {Opsin cDNA sequences of a UV and green rhodopsin of the satyrine butterfly Bicyclus anynana},
  Journal = {Insect Biochemistry and Molecular Biology},
  Volume = {32},
  Number = {11},
  Pages = {1383-1390},
  Note = {608MH Times Cited:3 Cited References Count:27},
  Abstract = {The cDNAs of an ultraviolet (UV) and long-wavelength (LW) (green) absorbing rhodopsin of the bush brown Bicyclus anynana were partially identified. The UV sequence, encoding 377 amino acids, is 76-79% identical to the UV sequences of the papilionids Papilio glaucus and Papilio xuthus and the moth Manduca sexta. A dendrogram derived from aligning the amino acid sequences reveals an equidistant position of Bicyclus between Papilio and Manduca. The sequence of the green opsin cDNA fragment, which encodes 242 amino acids, represents six of the seven transmembrane regions. At the amino acid level, this fragment is more than 80% identical to the corresponding LW opsin sequences of Dryas, Heliconius, Papilio (rhodopsin 2) and Manduca. Whereas three LW absorbing rhodopsins were identified in the papilionid butterflies, only one green opsin was found in B. anynana. (C) 2002 Elsevier Science Ltd. All rights reserved.},
  Keywords = {rt-pcr cdna cloning race ultraviolet and long-wavelength absorbing rhodopsin opsin insect vision bicyclus anynana visual pigments papilio-xuthus color-vision physiological characterization histological localization photoreceptor cell ultraviolet drosophila receptor identification},
  Year = {2002}
}
• [2002,article] bibtex
  M. Beltran, C. D. Jiggins, V. Bull, M. Linares, J. Mallet, W. O. McMillan, and E. Bermingham, "Phylogenetic discordance at the species boundary: Comparative gene genealogies among rapidly radiating Heliconius butterflies," Molecular Biology and Evolution, vol. 19, iss. 12, pp. 2176-2190, 2002.
  @article{ Author = {Beltran, M. and Jiggins, C. D. and Bull, V. and Linares, M. and Mallet, J. and McMillan, W. O. and Bermingham, E.},
  Title = {Phylogenetic discordance at the species boundary: Comparative gene genealogies among rapidly radiating Heliconius butterflies},
  Journal = {Molecular Biology and Evolution},
  Volume = {19},
  Number = {12},
  Pages = {2176-2190},
  Note = {625AT Times Cited:49 Cited References Count:53},
  Abstract = {Recent adaptive radiations provide excellent model systems for understanding speciation, but rapid diversification can cause problems for phylogenetic inference. Here we use gene genealogies to investigate the phylogeny of recent speciation in the heliconiine butterflies. We sequenced three gene regions, intron 3 (approximate to550 bp) of sex-linked triosephosphate isomerase (Tpi), intron 3 (approximate to450 bp) of autosomal mannose-phosphate isomerase (Mpi), and 1,603 bp of mitochondrial cytochrome oxidase subunits I and II (COI and COII), for 37 individuals from 25 species of Heliconius and related genera. The nuclear intron sequences evolved at rates similar to those of mitochondrial coding sequences, but the phylogenetic utility of introns was restricted to closely related geographic populations and species due to high levels of indel variation. For two sister species pairs, Heliconius erato-Heliconius himera and Heliconius melpoinene-Heliconius cydno, there was highly significant discordance between the three genes. At mtDNA and Tpi, the hypotheses of reciprocal monophyly and paraphyly of at least one species with respect to its sister could not be distinguished. In contrast alleles sampled from the third locus, Mpi, showed polyphyletic relationships between both species pairs. In all cases, recent coalescence of mtDNA lineages within species suggests that polyphyly of nuclear genes is not unexpected. In addition, very similar alleles were shared between melpomene and cydno, implying recent gene flow. Our finding of discordant genealogies between genes is consistent with models of adaptive speciation with ongoing gene How and highlights the need for multiple locus comparisons to resolve phylogeny among closely related species.},
  Keywords = {coalescence hybridization phylogeny mitochondrial DNA nuclear genes taq error mitochondrial-DNA sequences passion-vine butterflies ribosomal-rna genes hybrid zone population-genetics cladistic-analysis natural-selection nuclear gene lepidoptera nymphalidae},
  Year = {2002}
}
• [2002,article] bibtex
  M. J. Blum, "Rapid movement of a Heliconius hybrid zone: Evidence for phase III of Wright’s shifting balance theory?," Evolution, vol. 56, iss. 10, pp. 1992-1998, 2002.
  @article{ Author = {Blum, M. J.},
  Title = {Rapid movement of a Heliconius hybrid zone: Evidence for phase III of Wright's shifting balance theory?},
  Journal = {Evolution},
  Volume = {56},
  Number = {10},
  Pages = {1992-1998},
  Note = {615JH Times Cited:14 Cited References Count:39},
  Abstract = {It has been proposed that a moving hybrid zone can be a mechanism for the spread of adaptive traits in phase III of Wright's shifting balance model of evolution. Here I present an example of a moving hybrid zone in warningly colored Heliconius butterflies, a system which is considered to be a possible case of shifting balance evolution. Having moved approximately 47 km in 17 years, the hybrid zone shift has led to the H. erato hydara color pattern rapidly displacing the adjacent H. erato petiverana pattern. The movement is potentially due to dominance drive augmenting a slight selective advantage of H. erato hydara over H. erato petiverana, which is largely consistent with theoretical conditions favoring the success of phase III.},
  Keywords = {dominance drive heliconius moving hybrid zone shifting balance warning-color gene flow evolution mimicry perspective butterflies speciation selection dispersal diversity},
  Year = {2002}
}
• [2002,article] bibtex
  R. E. Naisbit, C. D. Jiggins, M. Linares, C. Salazar, and J. Mallet, "Hybrid sterility, Haldane’s rule and speciation in Heliconius cydno and H-melpomene," Genetics, vol. 161, iss. 4, pp. 1517-1526, 2002.
  @article{ Author = {Naisbit, R. E. and Jiggins, C. D. and Linares, M. and Salazar, C. and Mallet, J.},
  Title = {Hybrid sterility, Haldane's rule and speciation in Heliconius cydno and H-melpomene},
  Journal = {Genetics},
  Volume = {161},
  Number = {4},
  Pages = {1517-1526},
  Note = {588WT Times Cited:20 Cited References Count:52},
  Abstract = {Most genetic studies of Haldane's rule, in which hybrid sterility or inviability affects the heterogametic sex preferentially, have focused on Drosophila. It therefore remains unclear to what extent the conclusions of that work apply more generally, particularly in female-heterogametic taxa such as birds and Lepidoptera. Here we present a genetic analysis of Haldane's rule in Heliconius butterflies. Female F, hybrids between Heliconius melpomene and H. cydno are completely sterile, while males have normal to mildly reduced fertility. In backcrosses of male F-1 hybrids, female offspring range from completely sterile to fully fertile. Linkage analysis using the Z-linked triose-phosphate isomerase locus demonstrates a "large X" (Z) effect on sterility. Expression of female sterility varies among crosses in this and a previous study of Heliconius. Sterility may result from the production of normal but infertile eggs, production of small infertile eggs, or from a complete failure to develop ovarioles, which suggests multiple routes to the evolution of hybrid sterility in these Heliconius species. These results conform to the expectations of the "dominance" rather than "faster male" theories of Haldane's rule and suggest that relatively few loci are responsible. The two species are broadly sympatric and hybridize in the wild, so that female hybrid sterility forms one of several strong but incomplete barriers to gene flow in nature. The effect of female sterility is comparable to that of selection against non-mimetic hybrids, while mate choice forms a much stronger barrier to gene transfer.},
  Keywords = {passion-vine butterflies reproductive isolation drosophila-simulans gene flow evolution mimicry zones sex chromosomes selection},
  Year = {2002}
}
• [2002,article] bibtex
  M. J. Blum, C. D. Jiggins, and E. Bermingham, "Contrasting selected and neutral genetic markers across a stable Heliconius erato hybrid zone," , vol. in prep., 2002.
  @article{ Author = {Blum, M. J. and Jiggins, C. D. and Bermingham, E.},
  Title = {Contrasting selected and neutral genetic markers across a stable Heliconius erato hybrid zone},
  Volume = {in prep.},
  Year = {2002}
}
• [2002,article] bibtex
  N. Davies and E. Bermingham, "The historical biogeography of two Caribbean butterflies (Lepidoptera : Heliconiidae) as inferred from genetic variation at multiple loci," Evolution, vol. 56, iss. 3, pp. 573-589, 2002.
  @article{ Author = {Davies, N. and Bermingham, E.},
  Title = {The historical biogeography of two Caribbean butterflies (Lepidoptera : Heliconiidae) as inferred from genetic variation at multiple loci},
  Journal = {Evolution},
  Volume = {56},
  Number = {3},
  Pages = {573-589},
  Note = {542JV Times Cited:16 Cited References Count:39},
  Abstract = {Mitochondrial DNA and allozyme variation was examined in populations of two Neotropical butterflies, Heliconius charithonia and Dryas iulia. On the mainland, both species showed evidence of considerable gene flow over huge distances, The island populations, however, revealed significant genetic divergence across some, but not all, ocean passages. Despite the phylogenetic relatedness and broadly similar ecologies of these two butterflies, their intraspecific biogeography clearly differed. Phylogenetic analyses of mitochondrial DNA sequences revealed that populations of D iulia north of St. Vincent are monophyletic and were probably derived from South America. By contrast, the Jamaican subspecies of H. charithonia rendered West Indian H. charithonia polyphyletic with respect to the mainland populations thus, H. charithonia seems to have colonized the Greater Antilles on at least two separate occasions from Central America. Colonization velocity does not correlate with subsequent levels of gene flow in either species. Even where range expansion seems to have been instantaneous on a geological timescale, significant allele frequency differences at allozyme loci demonstrate that gene flow is severely curtailed across narrow ocean passages. Stochastic extinction, rapid (re)colonization, but low gene flow probably explain why, in the same species, some islands support genetically distinct and nonexpanding populations, while nearby a single lineage is distributed across several islands. Despite the differences, some common biogeographic patterns were evident between these butterflies and other West Indian taxa; such congruence suggests that intraspecific evolution in the West Indies has been somewhat constrained by earth history events, such as changes in sea level.},
  Keywords = {allozymes island mitochondrial DNA phylogeography population genetics west indies west-indies DNA phylogeography sequence},
  Year = {2002}
}
• [2002,article] bibtex
  N. S. Flanagan, M. J. Blum, A. Davison, M. Alamo, R. Albarran, K. Faulhaber, E. Peterson, and W. O. McMillan, "Characterization of microsatellite loci in neotropical Heliconius butterflies," Molecular Ecology Notes, vol. 2, iss. 4, pp. 398-401, 2002.
  @article{ Author = {Flanagan, N. S. and Blum, M. J. and Davison, A. and Alamo, M. and Albarran, R. and Faulhaber, K. and Peterson, E. and McMillan, W. O.},
  Title = {Characterization of microsatellite loci in neotropical Heliconius butterflies},
  Journal = {Molecular Ecology Notes},
  Volume = {2},
  Number = {4},
  Pages = {398-401},
  Note = {622ZM Times Cited:16 Cited References Count:9},
  Abstract = {The Heliconius butterflies offer exceptional opportunities for the study of the ecology and evolution of mimicry. Despite previous reports of difficulties in the development of microsatellite loci in Lepidoptera, we characterize 15 polymorphic loci in H. erato that show promise for genetic mapping and population studies in this and other species. Levels of variation were high, in both numbers and size ranges of alleles. The loci showed broad amplification success across the genus and in two other genera. All loci that amplified in a population of H. melpomene were polymorphic.},
  Keywords = {lepidoptera linkage mapping mimicry nymphalidae},
  Year = {2002}
}
• [2002,article] bibtex
  C. Estrada and C. D. Jiggins, "Patterns of pollen feeding and habitat preference among Heliconius species," Ecological Entomology, vol. 27, iss. 4, pp. 448-456, 2002.
  @article{ Author = {Estrada, C. and Jiggins, C. D.},
  Title = {Patterns of pollen feeding and habitat preference among Heliconius species},
  Journal = {Ecological Entomology},
  Volume = {27},
  Number = {4},
  Pages = {448-456},
  Note = {568YJ Times Cited:13 Cited References Count:36},
  Abstract = {1. The ecological circumstances that precipitate speciation remain poorly understood. Here, a community of Heliconius butterflies in lowland Panama was studied to investigate patterns of pollen use, and more specifically the ecological changes associated with the recent divergence of Heliconius melpomene (Linnaeus) and H. cydno (Doubleday). 2. Considering the seven commonest Heliconius species in the community, 32 types of pollen or spore were encountered in pollen loads but only five pollen species were common. Systematic exploitation of pollen was therefore confined to a small proportion of the flowers visited. 3. Most of the variation in pollen load composition between individuals was explained by differences in collecting locality. The exception was Psiguria, which was used in all habitats by the melpomene/hecale clade far more than by the erato/sapho clade. This may suggest an ancestral switch within Heliconius towards increased reliance on Psiguria pollen. 4. Heliconius cydno and H. melpomene differed significantly in pollen load composition for three of the five most commonly collected pollen species. This is most probably explained by differences in habitat preference; H. melpomene and its co-mimic H. erato are found in open habitat while H. cydno and its co-mimic H. sapho are found in closed-canopy forest. 5. As melpomene and cydno are known to hybridise occasionally, such differences in adult microhabitat contribute to pre-mating isolation. Habitat divergence between H. cydno and H. melpomene, which is associated with changes in mimicry, must have played a role in their recent speciation.},
  Keywords = {co-evolution ecological isolation lepidoptera pollen feeding speciation passion-vine butterflies sympatric speciation mimicry nymphalidae evolution lepidoptera adaptations sequences behavior biology},
  Year = {2002}
}
• [2002,article] bibtex
  D. G. Stavenga, "Reflections on colourful ommatidia of butterfly eyes," Journal of Experimental Biology, vol. 205, iss. 8, pp. 1077-1085, 2002.
  @article{ Author = {Stavenga, D. G.},
  Title = {Reflections on colourful ommatidia of butterfly eyes},
  Journal = {Journal of Experimental Biology},
  Volume = {205},
  Number = {8},
  Pages = {1077-1085},
  Note = {547FX J EXP BIOL},
  Abstract = {The eye shine of butterflies from a large number of ommatidia was observed with a modified epi-illumination apparatus equipped with an objective lens of large numerical aperture. A few representative cases are presented: the satyrine Bicyclus anynana, the heliconian Heliconius melpomene, the small white Pieris rapae and the small copper Lycaena phlaeas. The colour of the eye shine is determined mainly by the reflectance spectrum of the tapetal mirror and the transmittance spectrum of the photoreceptor screening pigments, if present near the light-guiding rhabdom. Reflectance spectra measured from individual ommatidia show that tapetum and screening pigments are co-expressed in fixed combinations, thus determining different ommatidial classes. The classes are distributed in an irregular pattern that can be rapidly assessed with the novel epi-illumination apparatus. Many butterfly species appear to have red-reflecting ommatidia, which is interpreted to indicate the presence of red-sensitive photoreceptors.},
  Year = {2002}
}
• [2002,article] bibtex
  A. Sweeney, C. Jiggins, and S. Johnsen, "Bio-optics and visual ecology of Heliconius color patterns.," Integrative and Comparative Biology, vol. 42, iss. 6, pp. 1321-1321, 2002.
  @article{ Author = {Sweeney, A. and Jiggins, C. and Johnsen, S.},
  Title = {Bio-optics and visual ecology of Heliconius color patterns.},
  Journal = {Integrative and Comparative Biology},
  Volume = {42},
  Number = {6},
  Pages = {1321-1321},
  Note = {669ND Times Cited:0 Cited References Count:0},
  Year = {2002}
}
• [2001,article] bibtex
  R. E. Naisbit, C. D. Jiggins, and J. Mallet, "Disruptive sexual selection against hybrids contributes to speciation between Heliconius cydno and Heliconius melpomene," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 268, iss. 1478, pp. 1849-1854, 2001.
  @article{ Author = {Naisbit, R. E. and Jiggins, C. D. and Mallet, J.},
  Title = {Disruptive sexual selection against hybrids contributes to speciation between Heliconius cydno and Heliconius melpomene},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {268},
  Number = {1478},
  Pages = {1849-1854},
  Note = {469UN Times Cited:35 Cited References Count:43},
  Abstract = {Understanding the fate of hybrids in wild populations is fundamental to understanding speciation. Here we provide evidence for disruptive sexual selection against hybrids between Heliconius cydno and Heliconius melpomene. The two species are sympatric across most of Central and Andean South America, and coexist despite a low level of hybridization. No-choice mating experiments show strong assortative mating between the species. Hybrids mate readily with one another, but both sexes show a reduction in mating success of over 50% with the parental species. Mating preference is associated with a shift in the adult colour pattern, which is involved in predator defence through Mullerian mimicry, but also strongly affects male courtship probability. The hybrids, which lie outside the curve of protection afforded by mimetic resemblance to the parental species, are also largely outside the curves of parental mating preference. Disruptive sexual selection against F-1 hybrids therefore forms an additional post-mating barrier to gene flow, blurring the distinction between pre-mating and post-mating isolation, and helping to maintain the distinctness of these hybridizing species.},
  Keywords = {lepidoptera nymphalidae hybridization mate choice post-mating isolation pre-mating isolation sympatric speciation ecological speciation haldane rule butterflies behavior evolution genetics mimicry traits origin},
  Year = {2001}
}
• [2001,article] bibtex
  C. L. Mazer and A. G. Appel, "Water loss and desiccation tolerances of longwing butterflies (Lepidoptera: Nymphalidae)," Environmental Entomology, vol. 30, iss. 4, pp. 631-636, 2001.
  @article{ Author = {Mazer, C. L. and Appel, A. G.},
  Title = {Water loss and desiccation tolerances of longwing butterflies (Lepidoptera: Nymphalidae)},
  Journal = {Environmental Entomology},
  Volume = {30},
  Number = {4},
  Pages = {631-636},
  Note = {465QU Times Cited:2 Cited References Count:23},
  Abstract = {Water relations and desiccation tolerance of adults of five species of heliconiine butterflies were examined in laboratory studies. Percentage of total body water ranged from 63.25% in female Dryas iulia F. to 67.8% in male Heliconius charitonius L.; water content of females was less than that for males in all five species. Male and female H. hecale Hewitson and D. iulia had lower percentages of total body water than the other species and sexes. Cuticular permeability ranged from 30.71 +/- 2.21 mug cm(-2) h(-1) mmHg(-1) in female D. iulia to 50.1 +/- 15.5 Ag cm(-2) h(-1) mmHg(-1) in female H. charitonius using Meeh's formula to estimate body surface area. Actual surface area measurements of butterflies were 12-15 times greater for a given body mass than surface areas calculated from Meeh's formula. Mortality of butterflies exposed to 30 degreesC and 0-2% RH ranged from 15% (female H. melpoinene rosina Boisduval and H. hecale) to 60% (male H, cydno Bates). Time of death varied between 8.67 +/- 2.0 h in male D. iulia and 11.3 +/- 1.55 h in female H, hecale; percentage of total body water lost at time of death ranged from 30.97% in male H, charitonius to 39.45% in female H, cydno. Percentage of initial mass lost and percentage of total body,,vater lost increased as a power function with desiccation time. The implications of water loss and desiccation tolerance to butterfly exhibits are discussed.},
  Keywords = {heliconius dryas iulia cuticular permeability body water content surface area},
  Year = {2001}
}
• [2001,article] bibtex
  Y. C. Golding, A. R. Ennos, and M. Edmunds, "Similarity in flight behaviour between the honeybee Apis mellifera (Hymenoptera : Apidae) and its presumed mimic, the dronefly Eristalis tenax (Diptera : Syrphidae)," Journal of Experimental Biology, vol. 204, iss. 1, pp. 139-145, 2001.
  @article{ Author = {Golding, Y. C. and Ennos, A. R. and Edmunds, M.},
  Title = {Similarity in flight behaviour between the honeybee Apis mellifera (Hymenoptera : Apidae) and its presumed mimic, the dronefly Eristalis tenax (Diptera : Syrphidae)},
  Journal = {Journal of Experimental Biology},
  Volume = {204},
  Number = {1},
  Pages = {139-145},
  Note = {393FW Times Cited:11 Cited References Count:34},
  Abstract = {It is generally accepted that the dronefly Eristalis tenax is a Batesian mimic of the honeybee Apis mellifera. Previous work has established that the foraging behaviour of droneflies is more similar to that of its model than to that of other more closely related flies, suggesting that behaviour may be important in the mimicry. Locomotor mimicry has been demonstrated in mimetic Heliconius butterflies but not in hoverflies. This study therefore investigated aspects of the flight behaviour of Eristalis tenax, Apis mellifera and two other flies, Syrphus ribesii and a Musca sp, Insects were filmed foraging on Helichrysum bracteum flowers, and flight sequences were analysed to determine flight velocities, flight trajectories and the percentage of time spent hovering. It was found that the flight behaviour of droneflies was more similar to that of honeybees than to that of the other flies. This suggests that the flight behaviour of Eristalis tenax may be mimetic.},
  Keywords = {batesian mimicry eristalis tenax apis mellifera dronefly honeybee foraging flight behaviour locomotor mimicry flight mimicry vine butterflies heliconius locomotor mimicry temperature morphology aerodynamics kinematics selection flies prey mass},
  Year = {2001}
}
• [2001,article] bibtex
  A. Briscoe, "Functional diversification of Lepidopteran Opsins following gene duplication," Molecular Biology and Evolution, vol. 18, pp. 2270-2279, 2001.
  @article{ Author = {Briscoe, A.},
  Title = {Functional diversification of Lepidopteran Opsins following gene duplication},
  Journal = {Molecular Biology and Evolution},
  Volume = {18},
  Pages = {2270-2279},
  Keywords = {HELICONIUS; MOLECULAR EVOLUTION; INSECTS, UV VISION},
  Year = {2001}
}
• [2001,article] bibtex
  R. Hsu, A. D. Briscoe, B. S. W. Chang, and N. E. Pierce, "Molecular evolution of a long wavelength-sensitive opsin in mimetic Heliconius butterflies (Lepidoptera : Nymphalidae)," Biological Journal of the Linnean Society, vol. 72, iss. 3, pp. 435-449, 2001.
  @article{ Author = {Hsu, R. and Briscoe, A. D. and Chang, B. S. W. and Pierce, N. E.},
  Title = {Molecular evolution of a long wavelength-sensitive opsin in mimetic Heliconius butterflies (Lepidoptera : Nymphalidae)},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {72},
  Number = {3},
  Pages = {435-449},
  Note = {BIOL J LINN SOC},
  Abstract = {This study examines the pattern of opsin nucleotide and amino acid substitution among mimetic species 'rings' of Heliconius butterflies that are characterized by divergent wing colour patterns. A long wavelength opsin gene, OPS1, was sequenced from each of seven species of Heliconius and one species of Dryas (Lepidoptera: Nymphalidae). A parsimony analysis of OPS1 nucleotide and amino acid sequences resulted in a phylogeny that was consistent with that presented by Brewer & Egan in 1997, which was based on mitochondrial cytochrome oxidase I and II as well as nuclear wingless genes. Nodes in the OPS1 phylogeny were well supported by bootstrap analysis and decay indices. An analysis of specific sites within the gene indicates that the accumulation of amino acid substitutions has occurred independently of the morphological diversification of Heliconius wing colour patterns. Amino acid substitutions were examined with respect to their location within the opsin protein and their possible interactions with the chromophore and the G-protein. Of the 15 amino acid substitutions identified among the eight species, one nonconservative replacement (A226Q) was identified in a position that may be involved in binding with the G-protein. (C) 2001 The Linnean Society of London.},
  Keywords = {PHYLOGENY; color vision; HELICONIUS},
  Year = {2001}
}
• [2001,article] bibtex
  D. D. Kapan, "Three-butterfly system provides a field test of mullerian mimicry," Nature, vol. 409, iss. 6818, pp. 338-340, 2001.
  @article{ Author = {Kapan, D. D.},
  Title = {Three-butterfly system provides a field test of mullerian mimicry},
  Journal = {Nature},
  Volume = {409},
  Number = {6818},
  Pages = {338-340},
  Note = {392VY Times Cited:55 Cited References Count:30},
  Abstract = {In 1879, Muller proposed that two brightly coloured distasteful butterfly species (co-models) that share a single warning-colour pattern would benefit by spreading the selective burden of educating predators(1-5). The mutual benefit of sharing warning signals among distasteful species, so-called mullerian mimicry, is supported by comparative evidence(2,3), theoretical studies(5,6) and laboratory simulations(7); however, to date, this key exemplar of adaptive evolution has not been experimentally tested in the field. To measure natural selection generated by mullerian mimicry, I exploited the unusual polymorphism of Heliconius cydno (Lepidoptera: Nymphalidae)(8). Here I show increased survival of H. cydno morphs that match locally abundant monomorphic co-model species. This study demonstrates mullerian mimicry in the field. It also shows that mullerian mimicry with several co-models generates geographically divergent selection, which explains the existence of polymorphism in distasteful species with warning coloration(9).},
  Keywords = {natural-selection shifting balance evolution diversity dynamics},
  Year = {2001}
}
• [2001,article] bibtex
  H. W. Krenn, K. P. Zulka, and T. Gatschnegg, "Proboscis morphology and food preferences in nymphalid butterflies (Lepidoptera : Nymphalidae)," Journal of Zoology, vol. 254, pp. 17-26, 2001.
  @article{ Author = {Krenn, H. W. and Zulka, K. P. and Gatschnegg, T.},
  Title = {Proboscis morphology and food preferences in nymphalid butterflies (Lepidoptera : Nymphalidae)},
  Journal = {Journal of Zoology},
  Volume = {254},
  Pages = {17-26},
  Note = {Part 1 433XD Times Cited:9 Cited References Count:50},
  Abstract = {Many species of nymphalid butterflies only exceptionally visit flowers and feed instead on tree sap, juice of rotting fruits and other decaying substances. To investigate whether the proboscis morphology of these non-flower-visiting Nymphalidae differs from that of nectarivorous butterflies, representatives from 64 nymphalid species with known feeding preferences were examined. Morphometric comparison of the proboscis revealed characteristic differences in proboscis length, tip-region length, wall composition, and number and shape of proboscis sensilla between these two feeding guilds. The investigated non-flower-visiting species belonging to Apaturinae, Limenitidinae, Morphinae, Brassolinae, Nymphalinae and Satyrinae, possess a relatively short and light-coloured proboscis which has a long tip-region with a great number of club-shaped sensilla styloconica. Densely arranged, these sensilla form a flat brush located laterally from the openings into the food canal on the dorsal side of the tip-region. Among the non-flower-visiting species, a second type of proboscis was found in fruit-feeding Charaxinae the stout tip-region of which is equipped with more widely spaced sensilla styloconica. The investigated flower-visiting Heliconiinae, Nymphalinae, Satyrinae, Danainae and Ithomiinae are characterized by a slender, darker-coloured proboscis with a rather short tip-region bearing fewer sensilla styloconica in a loose arrangement. Discriminant analysis revealed that the tip-region length, the number of sensilla styloconica and the relative proboscis length are the most important discriminating variables between the flower-visiting and the non-flower-visiting species. The proboscis morphology of nymphalid butterflies corresponds with certain feeding habits and allow us to make predictions on their food preferences. The 'brush-tipped' proboscis seems to have a functional role in the accumulation of fluid and the uptake of liquid from wet surfaces such as rotting fruits or tree sap. We conclude from the phylogeny of the examined taxa that this derived proboscis lip morphology evolved several times independently as an adaptation to the exploitation of new food resources.},
  Keywords = {insect butterfly mouthparts sensilla adaptation heliconius butterflies functional-morphology insecta papilionoidea behavior sensilla search family},
  Year = {2001}
}
• [2001,article] bibtex
  H. A. Orr and A. J. Betancourt, "Haldane’s sieve and adaptation from the standing genetic variation," Genetics, vol. 157, iss. 2, pp. 875-884, 2001.
  @article{ Author = {Orr, H. A. and Betancourt, A. J.},
  Title = {Haldane's sieve and adaptation from the standing genetic variation},
  Journal = {Genetics},
  Volume = {157},
  Number = {2},
  Pages = {875-884},
  Note = {399KL Times Cited:63 Cited References Count:32},
  Abstract = {We consider populations that adapt to a sudden environmental change by fixing alleles found at mutation-selection balance. In particular, we calculate probabilities of fixation for previously deleterious alleles, ignoring the input of new mutations. We find that "Haldane's siere"-the bias against the establishment of recessive beneficial mutations-does not hold under these conditions. Instead probabilities of fixation are generally independent of dominance. We show that this result is robust to patterns of sex expression for both X-Linked and autosomal loci. We further show that adaptive evolution is invariably slower at X-linked than autosomal loci when evolution begins from mutation-selection balance. This result differs from that obtained when adaptation uses new mutations, a finding that may have some bearing on recent attempts to distinguish between hitchhiking and background selection by contrasting the molecular population genetics of X-linked vs. autosomal loci. Last, we suggest a test to determine whether adaptation used new mutations or previously deleterious alleles from the standing genetic variation.},
  Keywords = {cyclodiene insecticide resistance population-genetics evolution heliconius selection trait rates},
  Year = {2001}
}
• [2001,article] bibtex
  N. Z. Cardoso, "Patterns of pollen collection and flower visitation by Heliconius butterflies in southeastern Mexico," Journal of Tropical Ecology, vol. 17, pp. 763-768, 2001.
  @article{ Author = {Cardoso, N. Z.},
  Title = {Patterns of pollen collection and flower visitation by Heliconius butterflies in southeastern Mexico},
  Journal = {Journal of Tropical Ecology},
  Volume = {17},
  Pages = {763-768},
  Note = {Part 5 499QX Times Cited:3 Cited References Count:25},
  Keywords = {disturbed habitats foraging lepidoptera lowland tropical forest pollen feeding pollen load psiguria-warscewiczii biology},
  Year = {2001}
}
• [2001,article] bibtex
  C. D. Jiggins, R. E. Naisbit, R. L. Coe, and J. Mallet, "Reproductive isolation caused by colour pattern mimicry," Nature, vol. 411, iss. 6835, pp. 302-305, 2001.
  @article{ Author = {Jiggins, C. D. and Naisbit, R. E. and Coe, R. L. and Mallet, J.},
  Title = {Reproductive isolation caused by colour pattern mimicry},
  Journal = {Nature},
  Volume = {411},
  Number = {6835},
  Pages = {302-305},
  Note = {432RT Times Cited:116 Cited References Count:23},
  Abstract = {Speciation is facilitated if ecological adaptation directly causes assortative mating(1), but few natural examples are known. Here we show that a shift in colour pattern mimicry was crucial in the origin of two butterfly species. The sister species Heliconius melpomene and Heliconius cydno recently diverged to mimic different model taxa, and our experiments show that their mimetic coloration is also important in choosing mates. Assortative mating between the sister species means that hybridization is rare in nature, and the few hybrids that are produced are nonmimetic, poorly adapted intermediates. Thus, the mimetic shift has caused both pre-mating and post-mating isolation. In addition, individuals from a population of H. melpomene allopatric to H. cydno court and mate with H. cydno more readily than those from a sympatric population. This suggests that assortative mating has been enhanced in sympatry.},
  Keywords = {heliconius butterflies natural-selection race formation warning-color speciation reinforcement evolution},
  Year = {2001}
}
• [2001,article] bibtex
  M. R. Kronforst and T. H. Fleming, "Lack of genetic differentiation among widely spaced subpopulations of a butterfly with home range behaviour," Heredity, vol. 86, pp. 243-250, 2001.
  @article{ Author = {Kronforst, M. R. and Fleming, T. H.},
  Title = {Lack of genetic differentiation among widely spaced subpopulations of a butterfly with home range behaviour},
  Journal = {Heredity},
  Volume = {86},
  Pages = {243-250},
  Note = {Part 2 443QK Times Cited:7 Cited References Count:37},
  Abstract = {We examine seven geographically separate subpopulations of Heliconius charithonia, a butterfly with well-documented home range behaviour, in Miami-Dade County, Florida, for genetic differentiation using cellulose acetate electrophoresis. These subpopulations exhibit little genetic variation (percent polymorphic loci = 27, average heterozygosity = 0.103) especially in comparison to populations of the same and related species from mainland South America. Allele frequencies do not differ among the subpopulations in south Florida and estimates of Wright's fixation index (F-ST) support that there is no detectable genetic differentiation among them. This result supports an earlier finding that the dispersal ability of Heliconius butterflies may be underestimated. However, it is unlikely that increased dispersal ability alone could account for the lack of genetic differentiation observed among subpopulations separated by almost 80 km. Given the likely effective population size of these subpopulations (N-e = 205) and the average generation lime of this species in the subtropics (in the range of 30-90 days), this lack of genetic differentiation is best explained by current or very recent gene flow following a stepping-stone model. Furthermore, this result provides evidence that the current extensive degree of habitat fragmentation surrounding the city of Miami does not limit gene flow among urban subpopulations of Heliconius charithonia.},
  Keywords = {allozyme electrophoresis gene flow genetic differentiation heliconius charithonia stepping-stone model urban fragmentation population-structure heliconius butterflies lepidoptera flow biology nymphalidae lycaenidae allozymes},
  Year = {2001}
}
• [2001,article] bibtex
  A. D. Briscoe and L. Chittka, "The evolution of color vision in insects," Annual Review of Entomology, vol. 46, pp. 471-510, 2001.
  @article{ Author = {Briscoe, A. D. and Chittka, L.},
  Title = {The evolution of color vision in insects},
  Journal = {Annual Review of Entomology},
  Volume = {46},
  Pages = {471-510},
  Note = {ANNU REV ENTOMOL},
  Abstract = {We review the physiological, molecular, and neural mechanisms of insect color vision. Phylogenetic and molecular analyses reveal that the basic bauplan, UV-blue-green-trichromacy, appears to date back to the Devonian ancestor of all pterygote insects. There are variations on this theme, however. These concern the number of color receptor types, their differential expression across the retina, and their fine tuning along the wavelength scale. In a few cases (but not in many others), these differences can be linked to visual ecology. Other insects have virtually identical sets of color receptors despite strong differences in lifestyle. Instead of the adaptionism that has dominated visual ecology in the past, we propose that chance evolutionary processes, history, and constraints should be considered. In addition to phylogenetic analyses designed to explore these factors, we suggest quantifying variance between individuals and populations and using fitness measurements to test the adaptive value of traits identified in insect color vision systems.},
  Keywords = {PHYLOGENY; color vision; HELICONIUS},
  Year = {2001}
}
• [2001,article] bibtex
  C. D. Jiggins, M. Linares, R. E. Naisbit, C. Salazar, Z. H. Yang, and J. Mallet, "Sex-linked hybrid sterility in a butterfly," Evolution, vol. 55, iss. 8, pp. 1631-1638, 2001.
  @article{ Author = {Jiggins, C. D. and Linares, M. and Naisbit, R. E. and Salazar, C. and Yang, Z. H. and Mallet, J.},
  Title = {Sex-linked hybrid sterility in a butterfly},
  Journal = {Evolution},
  Volume = {55},
  Number = {8},
  Pages = {1631-1638},
  Note = {475ET Times Cited:21 Cited References Count:50},
  Abstract = {Recent studies, primarily in Drosophila. have greatly advanced our understanding of Haldane's rule, the tendency for hybrid sterility or inviability to affect primarily the heterogametic sex (Haldane 1922). Although dominance theory (Turelli and Orr 1995) has been proposed as a general explanation of Haldane's rule, this remains to be tested in female-heterogametic taxa, such as the Lepidoptera. Here we describe a novel example of Haldane's rule in Heliconius melpomene (Lepidoptera; Nymphalidae). Female F-1 offspring are sterile when a male from French Guiana is crossed to a female from Panama, but fertile in the reciprocal cross. Male F(1)s are fertile in both directions. Similar female F-1 sterility occurs in crosses between French Guiana and eastern Colombian populations, Backcrosses and linkage analysis show that sterility results from an interaction between gene(s) on the Z chromosome of the Guiana race with autosomal factors in the Panama genome. Large X (or Z) effects are commonly observed in Drosophila, but to our knowledge have not been previously demonstrated for hybrid sterility in Lepidoptera. Differences in the abundance of male versus female or Z-linked versus autosomal sterility factors cannot be ruled out in our crosses as causes of Haldane's rule. Nonetheless, the demonstration that recessive Z-linked loci cause hybrid sterility in a female heterogametic species supports the contention that dominance theory provides a general explanation of Haldane's rule (Turelli and Orr 2000).},
  Keywords = {haldane's rule heliconius hybrid sterility lepidoptera speciation drosophila-simulans clade passion-vine butterflies haldane-rule reproductive isolation heliconius butterflies species-differences evolution genetics speciation lepidoptera},
  Year = {2001}
}
• [2001,article] bibtex
  J. M. Janssen, A. Monteiro, and P. M. Brakefield, "Correlations between scale structure and pigmentation in butterfly wings," Evolution & Development, vol. 3, iss. 6, pp. 415-423, 2001.
  @article{ Author = {Janssen, J. M. and Monteiro, A. and Brakefield, P. M.},
  Title = {Correlations between scale structure and pigmentation in butterfly wings},
  Journal = {Evolution & Development},
  Volume = {3},
  Number = {6},
  Pages = {415-423},
  Note = {512GD Times Cited:11 Cited References Count:43},
  Abstract = {We examined the correlation between color and structure of wing scales in the nymphalid butterflies Bicyclus, anynana and Heliconius melpomene. All scales in B. anynana are rather similar in comparison to the clear structural differences of differently pigmented scales in H. melpomene. Where scale structural differences in H. melpomene are qualitative, they seem to be quantitative in B. anynana. There is a "gradient" in the density of some structural elements, the cross ribs, in the scales of B. anynana: black, gold, and brown scales show progressively lower cross rib density within an individual. There is, however, high individual variation in the absolute cross rib densities (i.e., scales with a particular color and cross rib density in one individual may have a different color but similar density in another individual). By ectopically inducing color pattern during early pupal development, we examined whether a scale's color and its microstructure could be uncoupled. The effect of these manipulations appears to be different in B. anynana and H. melpomene. In Bicyclus, "black" scales induced by wing damage at an ectopic location normally containing brown scales acquire both an intermediate structure and color between that of brown and normal black scales. In Heliconius, however, intermediate colors or scale structure were never observed, and scales with an altered color (due to damage) always have the same structure as normal scales with that color. The results are discussed on the basis of gene expression patterns, variability in rates of scale development and pigment, and scale sclerotization pathways.},
  Keywords = {precis-coenia lepidoptera color pattern-formation dopa decarboxylase seasonal polyphenism melanin synthesis pieris-rapae nymphalidae papilionidae selection biology},
  Year = {2001}
}
• [2000,article] bibtex
  W. M. Neukirchen, "Zwei neue Heliconiinae aus Ecuador," Entomologische Zeitschrift, vol. 5, pp. 141-143, 2000.
  @article{ Author = {Neukirchen, W. M.},
  Title = {Zwei neue Heliconiinae aus Ecuador},
  Journal = {Entomologische Zeitschrift},
  Volume = {5},
  Pages = {141-143},
  Abstract = {New race of pardalinus 'julia' (after Julia Robinson) - Paratype from Yasuni collected by McMillan and Jiggins and new race of N. aoede, 'alicia' (after Alice Nield) from Morona-Santiago},
  Keywords = {HELICONIUS; SYSTEMATICS, TAXONOMY},
  Year = {2000}
}
• [2000,article] bibtex
  R. B. Srygley, "Locomotor mimicry among passion-vine butterflies Heliconius.," American Zoologist, vol. 40, iss. 6, pp. 1219-1219, 2000.
  @article{ Author = {Srygley, R. B.},
  Title = {Locomotor mimicry among passion-vine butterflies Heliconius.},
  Journal = {American Zoologist},
  Volume = {40},
  Number = {6},
  Pages = {1219-1219},
  Note = {422RA Times Cited:0 Cited References Count:0 Abstract},
  Year = {2000}
}
• [2000,misc] bibtex
  J. Mallet, Specious species arguments, 2000.
  @misc{ Author = {Mallet, J.},
  Title = {Specious species arguments},
  Month = {10 Jan 2000},
  Abstract = {email to Neal Smith from Jim Dear Neal, This is my response to your reponse to the article by Martin Brookes in the New Scientist. After your letter about David Lees, I remembered your comments that Chris forwarded it to me. Feel free to circulate this, but if you do please circulate it complete. Any comments by you should come afterwards. I doubt I shall reply again, so you have a free rein to vent your spleen! Jim ____________________________________________________________________________ 1) Your comments were about a New Scientist article about hybridization in general, which doesn't necessarily represent my own views. In my view Martin went over the top on this, and you should read my original papers rather than getting incensed about Martin's article. 2) Genetic continua can have humps in their distribution; we call these humps species, but viewing speciation as a complete cessation of reproductive continuity is clearly wrong, since many species hybridize regularly in nature (the blue whale and the fin whale, virtually all the birds of paradise and the ducks, about 25% of American warblers, 10% of all birds, about 12% of European butterflies, around 25% of Heliconius butterflies, and about 30% or more of plants). 3) So clearly, the "biological species concept" can't mean "complete reproductive isolation". Personally, I like to think that the mallard and the teal have speciated, and I would like also to think that Heliconius melpomene and Heliconius cydno have. They haven't completely closed down the reproductive channels, however, and it is very likely that some genes are still flowing between these closely related species. Increasingly, even in Drosophila, there is molecular evidence of actual interspecific gene flow. The point is, the production of phenomena like Drosophila pseudoobscura and D. persimilis, or blue and fin whale, or Heliconius melpomene and Heliconius cydno, i.e. the production of diversity in general, is more interesting than the final loss of reproductive compatibility, which they haven't yet achieved. Incidentally, phylogenetic or lineage definitions will do no better, since gene flow will mean there aren't necessarily any fixed differences, and also different genes under different levels of disruptive selection will have have different phylogenies. 4) The question is, therefore, HOW does the "biological species concept" "deal very well" (as you put it) with hybridization if reproductive isolation is not complete? The only thing I can think of, given that genes do "get through" is that we have to decide whether the genes that get through "matter" or not. If the flowing genes do "matter", then the two species fuse. If they don't "matter" then the two species stay distinct, at multiple loci. So in fact we use the genotypic distribution to tell us whether the gene flow "matters". Therefore, we can dispense with the knowledge about gene flow, and whether it "matters", and just look at the genotypes to define species. As I suggest. 5) We have to introduce a qualification of sympatry here. If two populations are abutting, but not completely overlapping, selection may produce a narrow cline (or hybrid zone) between two forms with fixed differences at multiple loci. Both the BSC and I agree on this: if hybrids are common in the overlap zone, such forms should be seen as separate races or subspecies, rather than as full species; if we don't make this qualification, all kinds of chaos gets loose, like defining geographical populations such as the melanic form of the peppered moth, with one difference, as separate species. [Actually, this chaos is precisely what Cracraft's version of the so-called "Phylogenetic Species Concept", which depends on fixed differences, proposes, in particular applied to geographic populations.] [However, there will always be people who persist in splitting. They will regard the toad Bombina bombina as a distinct species from Bombina variegata, or the bonobo from the chimpanzee, or the American black duck from the American mallard. This doesn't much matter, as it happens, because the same name can be applied equally to well-differentiated subspecies and species. What does matter more to me is if every trivial local variant gets named as a separate species, as is possible under the Cracraft idea; the utility of the species to classify continental faunas becomes very limited]. 5) Therefore, the only way of getting the biological species to work, and by this I mean in PRACTICE, is by referring to the state of multilocus genotypes in sympatry, rather than to the processes that produce this state. Theoretical appeals to vague concepts like "natural evolutionary units" will not help. So, if you are interested in the practical application of the biological species concept, then you have to do it via the existence or non-existence of distinguishable multilocus genotypic clusters in sympatry or zones of overlap. This practical application of the species and the invention of the subspecies concept for its converse was introduced formally for morphology by Rothschild and Karl Jordan in butterflies, and by people like DS Jordan in the US and Rothschild and Hartert in the 1880s-1910s, long before the "biological species concept" came along. All the mechanisms of cohesion, mate choice, hybrid inviability, ecological differences are of course evolutionarily important and indeed very interesting; but all you need for a species definition is to decide whether the sum of all these factors lead to stable separation of the genotypes into recognizably different (in genes, phenotype, or behaviour) populations when they overlap. 6) The production of a pair of observably different genotypic clusters is called speciation, and by admitting that you must use genetic markers or inherited traits to identify species, you now have complete freedom to test different mechanisms of the production of these trait differences, rather than incorporating these processes as "reproductive isolating mechanisms" into your definition of the problem in the first place. 7) Seems logical to me, but people educated in the Mayr tradition find it hard to get their head around it! However, I am certainly not the first to have made this argument. Read Darwin's chapter on hybridization (1859), for instance, which has been widely misinterpreted since the 1930s from the Mayr/Dobzhansky point of view. If you read it carefully (and not just the Mayr extracts!), you will see that Darwin agrees that reproductive isolation (specifically hybrid sterility and inviability) is CORRELATED with species status, but states firmly that it cannot be the "essence" of species that the BSC demands. There are many other more recent works whom I cite in my 1995 article. And if you don't think that defining what species is a practical matter, then I am quite happy to leave you swimming in the theoretical slime! But for me, science is practical. 9) We should be especially suspicious of philosophical arguments of the form: "Only under my view are species natural evolutionary units". Like christianity, this essentialist and basically religious stance can be and has been used to justify completely opposing points of view. In this case, every single one of the various phylogenetic as well as reproductive isolation-related ideas, and the evolutionary species concept, the recognition concept, the cohesion concept, and the general lineage concept all claim to have sorted out the true "reality" of species, rather than making the more modest and in my view more sensible argument that their method is actually useful. The idea that a single true species "reality" exists is part of the problem (although I do of course believe that species are completely "real" in the more limited "identifiable" sense). Jim Mallet 10 Jan 2000 At 09:55 14/09/1999 -0400, you wrote: >Jim, I think Neil Smith remains unconverted. See forwarded >message. ChrisDate: Mon, 13 Sep 1999 17:57:33 -0400 >From: "Neal Smith" >To: JIGGINSC@gamboa.si.edu >Subject: This came from BIRDCHAT >Mime-Version: 1.0 >Content-Type: text/plain; charset=US-ASCII >Content-Disposition: inline > >>But for those of you interested in reading some >>thought-provoking comments on (reasonably) modern species >concepts, I >>think you'll find it worth seeking out. >>"Mallet believes that biodiversity must be liberated from >its human straitjacket by abandoning the BSC and accepting >hybrids as an integral part of the biological picture. > >The Biological Species Concept (BSC) deals very well with >the >phenomenon of hybridization. It has nothing to do with a >"human >straitjacket." It simply recognized the clusters of >interbreeding >individuals as natural evolutionary units. > >>"This not to say that terms like speciesd, genus and family >do not have a place in our efforts to classify and catalogue >the biological world, but they also muddy the waters and get >in the way of our understanding." > >My understanding is just fine. It is Mr. Mallet who appears >to be >attempting to muddy the waters. > >>"Natural selection works on individuals or genes, not on >species." > >The unit of selection is the individual, not genes. The >consequences >of natural selection affect the composition of the gene pool >which is >the unit which evolves. Species are natural gene pools. > >>"Hybrids may face handicaps as a result of genetic >incompatibility, but essentially they are playing the same >evolutionary game as any other individual." > >Of course. They are individuals, and thus subject to >selection. Who >says different? > >>"The fittest survive, regarless of species boundaries." > >Survival has nothing to do with it. Fitness is a measure of >reproductive success. And whether or not you are a member >of a >species which offers potential mates, makes a lot of >difference in >your fitness. > >>"In Mallet's view, nature is full of different kinds of >things, and there are degrees of difference between these." > >And Mr. Mallet's point is...? > >>"In other words, life is a genetic continuum." > >If life were a genetic continuum, there is no way we could >identify >any of the birds we see. They would all be part of a >continuum of >life. We couldn't even diagnose a plant from an animal >under Mr. >Mallet's vision as described here; not to mention a Hawk >from a >Handsaw. >},
  Keywords = {SPECIES CONCEPTS},
  Year = {2000}
}
• [2000,article] bibtex
  C. M. Penz and H. W. Krenn, "Behavioral adaptations to pollen-feeding in Heliconius butterflies (Nymphalidae, Heliconiinae): An experiment using Lantana flowers," Journal of Insect Behavior, vol. 13, iss. 6, pp. 865-880, 2000.
  @article{ Author = {Penz, C. M. and Krenn, H. W.},
  Title = {Behavioral adaptations to pollen-feeding in Heliconius butterflies (Nymphalidae, Heliconiinae): An experiment using Lantana flowers},
  Journal = {Journal of Insect Behavior},
  Volume = {13},
  Number = {6},
  Pages = {865-880},
  Note = {381AW Times Cited:9 Cited References Count:34},
  Abstract = {Butterflies in the genera Heliconius and Laparus obtain fitness-related benefits from using amino acids derived from pollen. These butterflies have morphological features of the proboscis that facilitate pollen-feeding. Here we investigate behavioral characteristics potentially involved in pollen-gathering. Analysis of four behavioral characters showed that pollen-feeding species manipulate Lantana flowers faster and more thoroughly compared to non-pollen-feeding relatives. Although this suggests that pollen-feeding species are potentially more efficient in harvesting pollen, every butterfly tested successfully removed pollen from Lantana and non-pollen-feeding butterflies generally extracted larger amounts of pollen than Heliconius and Laparus. Morphological characteristics of the proboscis and the production of abundant fluid exudate help keep pollen attached in the proboscis for long periods of time-possibly key to Heliconius' and Laparus' ability to obtain amino acids from pollen. Our results, in concert with those of previous morphological analysis, indicate that behavioral and structural attributes associated with pollen-feeding in Heliconius and Laparus are subtle modifications of widespread butterfly characteristics and raise the question why other butterflies do not use pollen in their diet.},
  Keywords = {heliconius laparus lantana pollen-feeding flower-handling amino-acids cyanogenic glucosides egg-production lepidoptera nectar insecta exploitation biosynthesis lotaustralin morphology},
  Year = {2000}
}
• [2000,article] bibtex
  A. V. Z. Brower, "On the validity of Heliconius tristero Brower and Heliconius melpomene mocoa Brower, with notes on species concepts in Heliconius Kluk (Lepidoptera : Nymphalidae)," Proceedings of the Entomological Society of Washington, vol. 102, iss. 3, pp. 678-+, 2000.
  @article{ Author = {Brower, A. V. Z.},
  Title = {On the validity of Heliconius tristero Brower and Heliconius melpomene mocoa Brower, with notes on species concepts in Heliconius Kluk (Lepidoptera : Nymphalidae)},
  Journal = {Proceedings of the Entomological Society of Washington},
  Volume = {102},
  Number = {3},
  Pages = {678-+},
  Note = {335UY Times Cited:0 Cited References Count:37},
  Abstract = {Lamas' (1998) criticisms of Brewer's (1996a) taxonomy are shown to be based on misinterpretations of the evidence and of the rules of nomenclature. The names H. amaryllis amaryllis f. bellula Stichel and H. melpomene bellula Turner are unavailable. The name H. bellula Brewer is available but invalid. The name H. melpomene mocoa Brewer is the valid name for the subspecies of H. melpomene from the Putumayo region of southeastern Colombia. The Genotypic Cluster Species Concept is contrasted unfavorably to the Phylogenetic Species Concept with respect to the aims of systematics in general, and the resolution of relationships among geographically differentiated Heliconius taxa in particular.},
  Keywords = {heliconius melpomene bellula heliconius tristero nomenclature circumscription subspecies mitochondrial-DNA sequences cladistic-analysis race formation butterflies delimitation evolution mimicry},
  Year = {2000}
}
• [2000,article] bibtex
  E. Garcia-Barros, "Body size, egg size, and their interspecific relationships with ecological and life history traits in butterflies (Lepidoptera : Papilionoidea, Hesperioidea)," Biological Journal of the Linnean Society, vol. 70, iss. 2, pp. 251-284, 2000.
  @article{ Author = {Garcia-Barros, E.},
  Title = {Body size, egg size, and their interspecific relationships with ecological and life history traits in butterflies (Lepidoptera : Papilionoidea, Hesperioidea)},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {70},
  Number = {2},
  Pages = {251-284},
  Note = {326WG Times Cited:38 Cited References Count:157},
  Abstract = {The interspecific relationships between egg size and body size in butterflies (Papilionoidea and Hesperiidae), and between size and egg and larval development rime, larval trophic specificity, foodplant structure, climate, and phenology were investigated based on a sample of more than 1180 species. The independent contrasts method was used to avoid taxonomy-dependent results. Egg size is allometrically related to adult wing length by a slope of 0.43. Based on a subset of species, fecundity is correlated to adult body size, and there is evidence for a compromise between egg number and egg size (relative to adult size) across species. Butterfly size increases in correlation to the mean annual temperature of the species geographic range, but decreases in relation to increased aridity (or the length of the dry season). Larger butterflies tend to have longer larval development times, use large or structurally complex host plants, and are more likely to lay their eggs in batches, irrespective of climate. Larger eggs tend to develop more slowly, and give rise to larvae with longer developmental periods that will result in larger adults. No evidence was found to support a relationship between butterfly body size and polyphagy. A complex pattern of interrelationships links body size (and egg size) to other traits, although correlations other than that between egg size and body size are generally low. The results suggest the necessity of separating climate and seasonality into components that are relevant to insect life histories in comparative studies. (C) 2000 The Linnean Society of London.},
  Keywords = {allometry climate clutch size comparative methods fecundity host plant phenology polyphagy resource-allocation parasitoid hymenoptera heliconius butterflies independent contrasts latitudinal patterns reproductive effort offspring fitness weight variation fecundity evolution},
  Year = {2000}
}
• [2000,article] bibtex
  H. S. Engler, K. C. Spencer, and L. E. Gilbert, "Preventing cyanide release from leaves," Nature (London), vol. 406, pp. 144-145, 2000.
  @article{ Author = {Engler, H. S. and Spencer, K. C. and Gilbert, L. E.},
  Title = {Preventing cyanide release from leaves},
  Journal = {Nature (London)},
  Volume = {406},
  Pages = {144-145},
  Keywords = {HELICONIUS; INSECT/PLANT RELS (CHEM);},
  Year = {2000}
}
• [2000,article] bibtex
  A. D. Briscoe, "Six opsins from the butterfly Papilio glaucus: Molecular phylogenetic evidence for paralogous origins of red-sensitive visual pigments in insects," Journal of Molecular Evolution, vol. 51, iss. 2, pp. 110-121, 2000.
  @article{ Author = {Briscoe, A. D.},
  Title = {Six opsins from the butterfly Papilio glaucus: Molecular phylogenetic evidence for paralogous origins of red-sensitive visual pigments in insects},
  Journal = {Journal of Molecular Evolution},
  Volume = {51},
  Number = {2},
  Pages = {110-121},
  Note = {J MOL EVOL},
  Abstract = {It has been hypothesized that the UV-, blue-, and green- sensitive visual pigments of insects were present in the common ancestor of crustaceans and insects, whereas red-sensitive visual pigments evolved later as a result of convergent evolution. This hypothesis is examined with respect to the placement of six opsins from the swallowtail butterfly Papilio glaucus (PglRhl-6) in relationship to 46 other insect, crustacean, and chelicerate opsin sequences. All basal relationships established with maximum parsimony analysis except two are present in the distance and maximum likelihood analyses. In all analyses, the six P. glaucus opsins fall into three well-supported clades, comprised, respectively, of ultraviolet (UV), blue, and long-wavelength (LW) pigments, which appear to predate the radiation of the insects. Lepidopteran green- and red-sensitive visual pigments form a monophyletic clade, which lends support to the hypothesis from comparative physiological studies that red-sensitive visual pigments in insects have paralogous origins. Polymorphic amino acid sites (180, 197, 277, 285, 308), which are essential for generating the spectral diversity among the vertebrate red- and green-sensitive pigments are notably invariant in the Papilio red- and green-sensitive pigments. Other major tuning sites must be sought to explain the spectral diversification among these and other insect visual pigments.},
  Keywords = {PHYLOGENY; color vision; HELICONIUS},
  Year = {2000}
}
• [2000,phdthesis] bibtex
  R. Das, "Theoretical basis of likelihood methods in molecular phylogenetic inference," PhD Thesis , 2000.
  @phdthesis{ Author = {Das, R.},
  Title = {Theoretical basis of likelihood methods in molecular phylogenetic inference},
  School = {University College London},
  Type = {Masters},
  Keywords = {PHYLOGENY; HELICONIUS; LIKELIHOOD},
  Year = {2000}
}
• [2000,article] bibtex
  P. J. Wijngaarden and P. M. Brakefield, "The genetic basis of eyespot size in the butterfly Bicyclus anynana: an analysis of line crosses," Heredity, vol. 85, iss. 5, pp. 471-479, 2000.
  @article{ Author = {Wijngaarden, P. J. and Brakefield, P. M.},
  Title = {The genetic basis of eyespot size in the butterfly Bicyclus anynana: an analysis of line crosses},
  Journal = {Heredity},
  Volume = {85},
  Number = {5},
  Pages = {471-479},
  Note = {394KK Times Cited:10 Cited References Count:25},
  Abstract = {The tropical butterfly Bicyclus anynana shows wide variation in the size of the eyespots on the ventral side of its wings. Differences in the environmental temperature during late larval and early pupal stages are a major source of this variation, but variation also exists within temperatures. Using lines selected at a single temperature for large and small eyespots, and a number of crosses derived from these lines, we studied the genetic basis of eyespot size in B. anynana. We applied Lande's modification of the Castle-Wright (C-W) estimator to estimate the minimum number of genes contributing to the difference between the two lines. Estimates indicated that at least five genes are involved. As the C-W estimator is based on a number of simplifying assumptions, we tested to what extent one of these assumptions (additive gene action) is actually met. Joint-scaling tests indicated that the assumption of additivity is not satisfied and that dominance and probably epistasis play a role. Because reciprocal crosses were available we looked for evidence for sex-linkage and cytoplasmic effects. No evidence for cytoplasmic effects was found, but the data were consistent with the presence of one or more loci on the X-chromosome. The results are discussed in the context of the current model of eyespot formation.},
  Keywords = {bicyclus anynana butterfly eyespots castle-wright estimator cytoplasmic effects sex-linkage quantitative character evolution patterns number wings populations plasticity heliconius absence},
  Year = {2000}
}
• [1999,article] bibtex
  J. R. G. Turner and M. Speed, "Learning and memory in mimicry. I. Simulations of laboratory experiments," Philosophical Transactions of the Royal Society of London (B), vol. 351, pp. 1157-1170, 1999.
  @article{ Author = {Turner, J. R. G. and Speed, M.},
  Title = {Learning and memory in mimicry. I. Simulations of laboratory experiments},
  Journal = {Philosophical Transactions of the Royal Society of London (B)},
  Volume = {351},
  Pages = {1157-1170},
  Year = {1999}
}
• [1999,article] bibtex
  A. V. Z. Brower, "Delimitation of phylogenetic species with DNA sequences: A critique of Davis and Nixon’s population aggregation analysis," Systematic Biology, vol. 48, iss. 1, pp. 199-213, 1999.
  @article{ Author = {Brower, A. V. Z.},
  Title = {Delimitation of phylogenetic species with DNA sequences: A critique of Davis and Nixon's population aggregation analysis},
  Journal = {Systematic Biology},
  Volume = {48},
  Number = {1},
  Pages = {199-213},
  Note = {187VN Times Cited:40 Cited References Count:44},
  Keywords = {mitochondrial-DNA gene trees heliconius-erato lepidoptera nymphalidae systematics hypothesis evolution homology biology},
  Year = {1999}
}
• [1999,article] bibtex
  P. M. Brakefield and V. French, "Butterfly wings: the evolution of development of colour patterns," Bioessays, vol. 21, iss. 5, pp. 391-401, 1999.
  @article{ Author = {Brakefield, P. M. and French, V.},
  Title = {Butterfly wings: the evolution of development of colour patterns},
  Journal = {Bioessays},
  Volume = {21},
  Number = {5},
  Pages = {391-401},
  Note = {194RL Times Cited:37 Cited References Count:63},
  Abstract = {The diversity in colour patterns on butterfly wings provides great potential for understanding how developmental mechanisms may be modulated in the evolution of adaptive traits. In particular, we discuss concentric eyespot patterns, which have been shown by surgical experiments to be formed in response to signals from a central focus. Seasonal polyphenism shows how alternate phenotypes can develop through environmental sensitivity mediated by ecdysteroid hormones, whereas artificial selection and single gene mutants demonstrate genetic variation influencing the number, shape, size, position, and colour composition of the eyespots. The expression patterns of the regulatory gene Distal-less reveal that these changes can arise at several different developmental stages, and the phenotypes indicate that some forms of changed pattern may occur much more readily than others. Further study of the genes, of the developmental mechanisms, and of the functions of the patterns will provide novel insights about the evolution of morphological diversity. (C) 1999 John Wiley & Sons, Inc.},
  Keywords = {western white butterflies precis-coenia lepidoptera long-range action less gene activity bicyclus-anynana seasonal polyphenism phenotypic plasticity eyespot patterns heliconius lepidoptera quantitative genetics},
  Year = {1999}
}
• [1999,article] bibtex
  R. B. Srygley, "Incorporating motion into investigations of mimicry," Evolutionary Ecology, vol. 13, iss. 7-8, pp. 691-708, 1999.
  @article{ Author = {Srygley, R. B.},
  Title = {Incorporating motion into investigations of mimicry},
  Journal = {Evolutionary Ecology},
  Volume = {13},
  Number = {7-8},
  Pages = {691-708},
  Note = {418ZA Times Cited:7 Cited References Count:74},
  Abstract = {During the past thirty years, natural selection due to predation has been investigated with regard to prey motion in three areas that are relevant to the evolution of mimicry: (1) anti-apostatic selection, (2) locomotor mimicry, and (3) escape mimicry. Anti-apostatic selection, or selection against the odd individuals, arises when prey are at very high densities or when prey are Mullerian mimics. When prey are at high densities, motion of the prey increases selection against odd individuals. When the prey are Mullerian mimics, motion may also play an important role in strengthening selection against odd individuals. This may explain locomotor mimicry between Mullerian mimics. Locomotor mimicry arises when two distantly-related prey species appear alike in behaviour, and there is a corresponding suite of morphological, physiological, and biomechanical traits that the prey have in common. Locomotor mimicry has been demonstrated in Mullerian mimics. It is also predicted to occur in Batesian mimics but with important limitations due to selection by the predator for the prey to maintain the ability to escape if detected. Locomotor mimicry may also occur between palatable species that are alike as a result of unprofitable prey (or escape) mimicry. Escape mimicry arises when prey are difficult to capture. By frustration learning, the predator associates the colour of the prey with unprofitability. In all three instances, dis-similarity in colour or motion probably increases selection against the odd individual. In addition, the interaction of colour and motion gives rise to greater reliability of the signals to a specialist predator. However for a generalist predator, multiple component signals of the prey lead to errors in signal perception and greater risk of cheating.},
  Keywords = {anti-apostatic selection escape mimicry locomotor mimicry perception predator-prey selection vine butterflies heliconius locomotor mimicry neotropical butterflies apostatic selection receiver psychology aggressive mimicry pierid butterflies flight performance natural-selection body-mass},
  Year = {1999}
}
• [1999,article] bibtex
  J. R. G. Turner and M. P. Speed, "How weird can mimicry get?," Evolutionary Ecology, vol. 13, pp. 807-827, 1999.
  @article{ Author = {Turner, J. R. G. and Speed, M. P.},
  Title = {How weird can mimicry get?},
  Journal = {Evolutionary Ecology},
  Volume = {13},
  Pages = {807-827},
  Year = {1999}
}
• [1999,article] bibtex
  A. D. Briscoe, "Intron splice sites of Papilio glaucus PglRh3 corroborate insect opsin phylogeny," Gene, vol. 230, iss. 1, pp. 101-109, 1999.
  @article{ Author = {Briscoe, A. D.},
  Title = {Intron splice sites of Papilio glaucus PglRh3 corroborate insect opsin phylogeny},
  Journal = {Gene},
  Volume = {230},
  Number = {1},
  Pages = {101-109},
  Note = {GENE},
  Abstract = {Full-length cDNA clones encoding the PglRh3 opsin from the tiger swallowtail butterfly Papilio glaucus were isolated from cDNA synthesized from adult head tissue total RNA. This cDNA consists of 1679 nucleotides and contains a single open reading frame predicted to be 379 amino acids in length. PCR amplification of genomic DNA with primers spanning the coding region yielded a single 2760 bp fragment which was sequenced. The PglRh3 gene has nine exons and eight introns, four of which are in unique locations relative to the positions of introns in other known insect opsin sequences. Phylogenetic analyses of amino acid and nucleotide sequence data places PglRh3 within a clade of insect visual pigments thought to be sensitive to long wavelengths of light. The genomic structure of PglRh3 is the first characterized from a member of this opsin clade. Three PglRh3 intron positions are shared with Drosophila Rhl, and one of these is also shared with Drosophila Rh2. By contrast, none of the known intron locations in a clade of anciently diverged ultraviolet- and blue-sensitive visual pigments are shared by P. glaucus PglRh3, Drosophila Rhl or Rh2, The placement of introns within opsin genes therefore independently supports the clustering of a putatively long-wavelength-sensitive clade with a clade of blue-green-sensitive visual pigments. (C) 1999 Elsevier Science B.V. All rights reserved.},
  Keywords = {PHYLOGENY; color vision; HELICONIUS},
  Year = {1999}
}
• [1999,article] bibtex
  F. Pichaud, A. Briscoe, and C. Desplan, "Evolution of color vision," Current Opinion in Neurobiology, vol. 9, iss. 5, pp. 622-627, 1999.
  @article{ Author = {Pichaud, F. and Briscoe, A. and Desplan, C.},
  Title = {Evolution of color vision},
  Journal = {Current Opinion in Neurobiology},
  Volume = {9},
  Number = {5},
  Pages = {622-627},
  Note = {CURR OPIN NEUROBIOL},
  Abstract = {Color vision is achieved by comparing the inputs from retinal photoreceptor neurons that differ in their wavelength seansitivity. Recent studies have elucidated the distribution and phytogeny of opsins, the family of light-sensitive molecules involved in this process. interesting new findings suggest that animals have evolved a strategy to achieve specific sensitivity through the mutually exclusive expression of different opsin genes in photoreceptors.},
  Keywords = {PHYLOGENY; color vision; HELICONIUS},
  Year = {1999}
}
• [1999,article] bibtex
  J. Mallet and M. Joron, "Evolution of diversity in warning color and mimicry: Polymorphisms, shifting balance, and speciation," Annual Review of Ecology and Systematics, vol. 30, pp. 201-233, 1999.
  @article{ Author = {Mallet, J. and Joron, M.},
  Title = {Evolution of diversity in warning color and mimicry: Polymorphisms, shifting balance, and speciation},
  Journal = {Annual Review of Ecology and Systematics},
  Volume = {30},
  Pages = {201-233},
  Note = {276MX Times Cited:114 Cited References Count:183},
  Abstract = {Mimicry and warning color are highly paradoxical adaptations. Color patterns in both Mullerian and Batesian mimicry are often determined by relatively few pattern-regulating loci with major effects. Many of these loci are "supergenes," consisting of multiple, tightly linked epistatic elements. On the one hand, strong purifying selection on these genes must explain accurate resemblance (a reduction of morphological diversity between species), as well as monomorphic color patterns within species. On the other hand, mimicry has diversified at every taxonomic level; warning color has evolved from cryptic patterns, and there are mimetic polymorphisms within species, multiple color patterns in different geographic races of the same species, mimetic differences between sister species, and multiple mimicry rings within local communities. These contrasting patterns can be explained, in part, by the shape of a "number-dependent" selection function first modeled by Fritz Muller in 1879: Purifying selection against any warning-colored morph is very strong when that morph is rare, but becomes weak in a broad basin of intermediate frequencies, allowing opportunities for polymorphisms and genetic drift. This Mullerian explanation, however, makes unstated assumptions about predator learning and forgetting which have recently been challenged. Today's "receiver psychology" models predict that classical Mullerian mimicry could be much rarer than believed previously, and that "quasi-Batesian mimicry," a new type of mimicry intermediate between Mullerian and Batesian, could be common. However, the new receiver psychology theory is untested, and indeed it seems to us unlikely; alternative assumptions could easily lead to a more traditional Mullerian/Batesian mimicry divide.},
  Keywords = {aposematism batesian mimicry mullerian mimicry defensive coloration predator behavior aposematic coloration mullerian mimicry heliconius butterflies individual selection natural-selection hybrid zones danaus-chrysippus neotropical butterflies cyanogenic glucosides receiver psychology},
  Year = {1999}
}
• [1999,phdthesis] bibtex
  M. Beltrán, "Evidencia genética (Alozimas) para evaluar el posible origen híbrido de Heliconius heurippa (Lepidoptera: Nymphalidae)," PhD Thesis , 1999.
  @phdthesis{ Author = {Beltrán, M.},
  Title = {Evidencia genética (Alozimas) para evaluar el posible origen híbrido de Heliconius heurippa (Lepidoptera: Nymphalidae)},
  School = {Universidad de los Andes},
  Type = {Masters},
  Keywords = {ALLOZYMES; HELICONIUS;},
  Year = {1999}
}
• [1999,article] bibtex
  C. M. Penz, "Higher level phylogeny for the passion-vine butterflies (Nymphalidae; Heliconiinae) based on early stage and adult morphology," Zoological Journal of the Linnean Society, vol. 127, pp. 277-344, 1999.
  @article{ Author = {Penz, C. M.},
  Title = {Higher level phylogeny for the passion-vine butterflies (Nymphalidae; Heliconiinae) based on early stage and adult morphology},
  Journal = {Zoological Journal of the Linnean Society},
  Volume = {127},
  Pages = {277-344},
  Keywords = {HELICONIUS; PHYLOGENY; CLASSIFICATION/TAXONOMY},
  Year = {1999}
}
• [1999,article] bibtex
  R. B. Srygley, "Locomotor mimicry in Heliconius butterflies: contrast analyses of flight morphology and kinematics," Philosophical Transactions of the Royal Society of London Series B-Biological Sciences, vol. 354, iss. 1380, pp. 203-214, 1999.
  @article{ Author = {Srygley, R. B.},
  Title = {Locomotor mimicry in Heliconius butterflies: contrast analyses of flight morphology and kinematics},
  Journal = {Philosophical Transactions of the Royal Society of London Series B-Biological Sciences},
  Volume = {354},
  Number = {1380},
  Pages = {203-214},
  Note = {164EZ Times Cited:16 Cited References Count:40},
  Abstract = {Mullerian mimicry is a mutualism involving the evolutionary convergence of colour patterns of prey on a warning signal to predators. Behavioural mimicry presumably adds complexity to the signal and makes it more difficult for Batesian mimics to parasitize it. To date, no one has quantified behavioural mimicry in Mullerian mimicry groups. However, morphological similarities among members of mimicry groups suggested that pitching oscillations of the body and wing-beat frequency (WBF) might converge with colour pattern. I compared the morphology and kinematics of four Heliconius species, which comprised two mimicry pairs. Because the mimics arose from two distinct lineages, the relative contributions of mimicry and phylogeny to variation in the species' morphologies and kinematics were examined. The positions of the centre of body mass and centre of wing mass and wing shape diverged among species within lineages, and converged among species within mimicry groups. WBF converged within mimicry groups, and it was coupled with body pitching frequency However, body-pitching frequency was too variable to distinguish mimicry groups. Convergence in WBF may be due, at least in part, to biomechanical consequences of similarities in wing length, wing shape or the centre of wing mass among co-mimics. Nevertheless, convergence in WBF among passion-vine butterflies serves as the first evidence of behavioural mimicry in a mutualistic context.},
  Keywords = {mimicry heliconius comparative analysis flight mutualism hovering insect flight aerodynamics evolution predation mass temperature},
  Year = {1999}
}
• [1999,article] bibtex
  P. J. DeVries, R. Lande, and D. Murray, "Associations of co-mimetic ithomiine butterflies on small spatial and temporal scales in a neotropical rainforest," Biological Journal of the Linnean Society, vol. 67, iss. 1, pp. 73-85, 1999.
  @article{ Author = {DeVries, P. J. and Lande, R. and Murray, D.},
  Title = {Associations of co-mimetic ithomiine butterflies on small spatial and temporal scales in a neotropical rainforest},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {67},
  Number = {1},
  Pages = {73-85},
  Note = {197JR Times Cited:12 Cited References Count:47},
  Abstract = {To test whether ithomiine butterfly species within Mullerian mimetic classes are associated in space and time, we sampled a community of ithomiine butterflies at monthly intervals with traps in the canopy and the understory of four forest habitats: primary, higrade, secondary and edge. A species accumulation curve reached an asymptote at 22 species, suggesting that these species have a greater preference for feeding on fruit juices than other ithomiines known to occur at the study site. Species richness and individual abundance showed marked temporal variation, and there were slight differences in the distribution of species richness and individual abundance among the four habitats. The 22 species sampled in this study were not stratified vertically. The five mimetic colour classes of these butterflies were unequally distributed among the four habitats and over the course of the twelve months. There is suggestive evidence that co-mimic species occurred in the same habitats, and strong evidence that they occurred at the same times. Habitat and temporal effects each contributed approximately 10% to the total mimetic class diversity, with the temporal effect being slightly larger than that of habitat. This study demonstrates that Mullerian co-mimic associations can be measured on a much smaller scale than has been done previously. (C) 1999 The Linnean Society of London.},
  Keywords = {ithomiinae nymphalidae mullerian mimicry habitat association vertical stratification species abundance distribution species randomization spatial distribution temporal distribution habitat destruction heliconius butterflies muellerian mimicry species-diversity shifting balance evolution forest selection lepidoptera nymphalidae behavior},
  Year = {1999}
}
• [1999,article] bibtex
  N. M. A. Holmgren and M. Enquist, "Dynamics of mimicry evolution," Biological Journal of the Linnean Society, vol. 66, iss. 2, pp. 145-158, 1999.
  @article{ Author = {Holmgren, N. M. A. and Enquist, M.},
  Title = {Dynamics of mimicry evolution},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {66},
  Number = {2},
  Pages = {145-158},
  Note = {170QZ Times Cited:21 Cited References Count:36},
  Abstract = {We simulated mimicry evolution by allowing three populations to coevolve: two populations of senders and one of receivers. Artificial neural networks were used to model receivers, and it was assumed that recognition was inherited. The senders' signals consisted of nine dimensions. Changes to receivers and senders were caused by random mutations during the course of the simulation. Whereas it paid both types of senders to elicit the same response from the receiver, it benefited the receiver to respond in this way only towards one of the sender types. The receiver was thus in conflict with one of the senders, e.g. as in Batesian mimicry. Monotonically increasing response gradients caused the appearance of the model and the mimic to move in the same direction. Mimicry evolved because the mimic approached the model faster than the model moved away. Even after mimicry was established the model and the mimic were constantly changing in appearance. Our results conform with what is known in comparative psychology and ethology about how animals respond to stimuli. Several of our results are a direct consequence of recognition and have not, to our knowledge, been reported before, showing the importance of considering the recognition mechanism in detail when studying mimicry. (C) 1999 The Linnean Society of London.},
  Keywords = {artificial neural network batesian mimicry communication imitation signalling muellerian mimicry heliconius selection},
  Year = {1999}
}
• [1999,article] bibtex
  A. Davison, W. O. McMillan, A. S. Griffin, C. D. Jiggins, and J. L. B. Mallet, "Behavioral and physiological differences between two parapatric Heliconius species," Biotropica, vol. 31, iss. 4, pp. 661-668, 1999.
  @article{ Author = {Davison, A. and McMillan, W. O. and Griffin, A. S. and Jiggins, C. D. and Mallet, J. L. B.},
  Title = {Behavioral and physiological differences between two parapatric Heliconius species},
  Journal = {Biotropica},
  Volume = {31},
  Number = {4},
  Pages = {661-668},
  Note = {274QM Times Cited:2 Cited References Count:36},
  Abstract = {The behavior and physiology of two parapatric sibling species, Heliconius erato cyrbia Godt. and H. himera Hew., were investigated to assess if environmental adaptation enabled stable morphological, generic, and ecological differences to exist in the face of hybridization. Morning and evening activity, egg production, and Larval development time of H. himera and H. erato in insectaries were recorded; individuals were collected in allopatry and in sympatry from a hybrid zone in which the species overlapped. Studies were performed at ambient conditions within the natural range of H; himera. H. himera was considerably more active than H. erato, flying earlier in the morning and later in the evening, even when both species were collected in sympatry. Similarly, H. himera laid more eggs, and the hatched larvae developed more rapidly The results suggest that physiological constraints are an important selective force that may have been important in speciation and counteracts hybridization in the maintenance of the H. himera/H. erato contact zone. Ecological selection, arising from adaptation to low temperatures, may help explain the competitive exclusion of H. erato by H. himera in the drier, cooler montane habitat favored by the latter species.},
  Keywords = {competitive exclusion heliconius hybridization hybrid zone lepidoptera parapatry speciation fire-bellied toads hybrid zones bombina-bombina butterflies speciation erato adaptation radiation evolution variegata},
  Year = {1999}
}
• [1999,article] bibtex
  M. P. Speed and J. R. H. Turner, "Learning and memory in mimicry: II. Do we understand the mimicry spectrum?," Biological Journal of the Linnean Society, vol. 67, pp. 281-312, 1999.
  @article{ Author = {Speed, M. P. and Turner, J. R. H.},
  Title = {Learning and memory in mimicry: II. Do we understand the mimicry spectrum?},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {67},
  Pages = {281-312},
  Year = {1999}
}
• [1999,article] bibtex
  M. P. Speed, "Robot predators in virtual ecologies: the importance of memory in mimicry studies," Animal Behaviour, vol. 57, pp. 203-213, 1999.
  @article{ Author = {Speed, M. P.},
  Title = {Robot predators in virtual ecologies: the importance of memory in mimicry studies},
  Journal = {Animal Behaviour},
  Volume = {57},
  Pages = {203-213},
  Note = {Part 1 162JD Times Cited:18 Cited References Count:56},
  Abstract = {An important means of investigating gains and losses to prey caused by mimicry is through mathematical or computer constructs which represent and explore limited aspects of mimicry situations. Such studies use virtual predators which are usually simple automata, 'robots' that, through simple rules, vary virtual attack rates on virtual insect prey. In this paper I consider the effect of variations in predator memory and learning on mimicry dynamics. When there is mimicry between unequally noxious prey, the way that memory is modelled is shown to be crucial. If forgetting rates are fixed, an increase in the density of the least defended prey produces monotonic gains or losses in protection. However, if forgetting rate is inversely related in some way to degree of noxiousness of the prey then attack rates initially rise with the density of the least defended prey, reach a cusp and then fall. I show that the generation of this highly unconventional up-down result appears to be independent of variations in learning rate. This work shows how sensitive the predictions of virtual predators may be to relatively small changes in behavioural rules. (C) 1999 The Association for the Study of Animal Behaviour.},
  Keywords = {heliconius butterflies muellerian mimicry mullerian mimicry hybrid zones evolution psychology coloration paradigms stability spectrum},
  Year = {1999}
}
• [1999,article] bibtex
  M. Joron, I. R. Wynne, G. Lamas, and J. Mallet, "Variable selection and the coexistence of multiple mimetic forms of the butterfly Heliconius numata," Evolutionary Ecology, vol. 13, iss. 7-8, pp. 721-754, 1999.
  @article{ Author = {Joron, M. and Wynne, I. R. and Lamas, G. and Mallet, J.},
  Title = {Variable selection and the coexistence of multiple mimetic forms of the butterfly Heliconius numata},
  Journal = {Evolutionary Ecology},
  Volume = {13},
  Number = {7-8},
  Pages = {721-754},
  Note = {418ZA Times Cited:13 Cited References Count:51},
  Abstract = {Polymorphism in aposematic animals and coexistence of multiple mimicry rings within a habitat are not predicted by classical Mullerian mimicry. The butterfly Heliconius numata Cramer (Lepidoptera: Nymphalidae; Heliconiinae) is both polymorphic and aposematic. The polymorphism is due to variation at a single locus (or 'supergene') which determines colour patterns involved in Mullerian mimicry. We sampled 11 sites in a small area (approx. 60x30km) of North-eastern Peru for H. numata and its co-mimics in the genus Melinaea and Athyrtis (Ithomiinae), and examined the role of temporal and spatial heterogeneity in the maintenance of polymorphism. Colour-patterns of Melinaea communities, which constitute the likely 'mimetic environment' for H. numata, are differentiated on a more local scale than morphs of H. numata, but the latter do show a strong and significant response to local selection for colour-pattern. In contrast, analysis of enzyme polymorphism in H. numata across the region revealed no spatial structure, which is consistent with a high mobility of this species. Differences in spatial variability in the two taxa may have caused H. numata to become polymorphic, while temporal variability, not significant in this study, probably has a lesser effect. The mimetic polymorphism is therefore explained by means of multiple selection-migration clines at a single locus, a similar process to that which explains narrow hybrid zones between geographic races of other Heliconius butterflies.},
  Keywords = {heliconius numata melinaea ithomiinae aposematism mullerian mimicry polymorphism spatial heterogeneity frequency-dependent selection population genetic structure danaus-chrysippus l hybrid zone warning-color mullerian mimicry home range gene flow lepidoptera polymorphism nymphalidae spectrum},
  Year = {1999}
}
• [1999,article] bibtex
  D. Rodrigues and G. R. P. Moreira, "Feeding preference of Heliconius erato (Lep.: Nymphalidae) in relation to leaf age and consequences for larval performance," Journal of the Lepidopterists’ Society, vol. 53, pp. 108-113, 1999.
  @article{ Author = {Rodrigues, D. and Moreira, G. R. P.},
  Title = {Feeding preference of Heliconius erato (Lep.: Nymphalidae) in relation to leaf age and consequences for larval performance},
  Journal = {Journal of the Lepidopterists' Society},
  Volume = {53},
  Pages = {108-113},
  Keywords = {HELICONIUS; LEPIDOPTERA LIFE HISTORIES},
  Year = {1999}
}
• [1999,article] bibtex
  R. B. Srygley and C. P. Ellington, "Estimating the relative fitness of local adaptive peaks: the aerodynamic costs of flight in mimetic passion-vine butterflies Heliconius," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 266, iss. 1435, pp. 2239-2245, 1999.
  @article{ Author = {Srygley, R. B. and Ellington, C. P.},
  Title = {Estimating the relative fitness of local adaptive peaks: the aerodynamic costs of flight in mimetic passion-vine butterflies Heliconius},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {266},
  Number = {1435},
  Pages = {2239-2245},
  Note = {263DZ Times Cited:7 Cited References Count:21},
  Abstract = {In a related paper, we demonstrated that mimetic Heliconius butterflies have converged in wing-beat frequency and degree of asymmetry in the wing motion, whereas sister species are dissimilar in these same traits. Warning signals of sympatric, distasteful species converge in evolutionary models in order to educate their predators more efficiently that the signal is associated with unprofitable prey. Barring other constraints, the behaviours of the different co-mimetic pairs should ultimately converge on that behaviour which minimizes the energetic cost of flight. We estimated the energetic cost of each mimic's flight behaviour in order to predict the difference in height of each fitness peak and the direction of convergent selection qualitatively. Following adjustments for body mass, mimetic Heliconius melpomene and Heliconius erato required more aerodynamic power than Heliconius cydno and Heliconius sapho. This difference was attributed to the slower flight speeds and higher wing-beat frequencies of H. melpomene and H. erato. Consequently, H. melpomene and H, erato expended more energy per unit distance per unit body mass than H. cydno and H. saaho. However, differences in body mass may equalize energy budgets and stabilize the sympatric coexistence of the two pairs of co-mimics.},
  Keywords = {locomotor mimicry behaviour biomechanics coexistence mutualism energetics hovering insect flight neotropical butterflies power requirements natural-selection mimicry kinematics mass},
  Year = {1999}
}
• [1999,article] bibtex
  J. Mallet, "Causes and consequences of a lack of coevolution in mullerian mimicry," Evolutionary Ecology, vol. 13, iss. 7-8, pp. 777-806, 1999.
  @article{ Author = {Mallet, J.},
  Title = {Causes and consequences of a lack of coevolution in mullerian mimicry},
  Journal = {Evolutionary Ecology},
  Volume = {13},
  Number = {7-8},
  Pages = {777-806},
  Note = {418ZA Times Cited:28 Cited References Count:73},
  Abstract = {Mullerian mimicry, in which both partners are unpalatable to predators, is often used as an example of a coevolved mutualism. However, it is theoretically possible that some Mullerian mimics are parasitic if a weakly defended mimic benefits at the expense of a more highly defended model, a phenomenon known as 'quasi-Batesian mimicry'. The theory expounded by Muller and extended here for unequal unpalatability, on the other hand, suggests that quasi-Batesian mimicry should be rare in comparison with classical, or mutualistic Mullerian mimicry. Evolutionarily, quasi-Batesian mimicry has consequences similar to classical Batesian mimicry, including unilateral 'advergence' of the mimic to the model, and diversifying frequency-dependent selection on the mimic which may lead to mimetic polymorphism. In this paper, theory and empirical evidence for mutual benefit and coevolution in Mullerian mimicry are reviewed. I use examples from well-known insect Mullerian mimicry complexes: the Limenitis-Danaus (Nymphalidae) system in North America, the Bombus-Psithyrus (Apidae) system in the north temperate zone, and the Heliconius-Laparus (Nymphalidae) system in tropical America. These give abundant evidence for unilateral advergence, and no convincing evidence, to my knowledge, for coevolved mutual convergence. Furthermore, mimetic polymorphisms are not uncommon. Yet classical mutualistic Mullerian mimicry, coupled with spatial (and possibly temporal) variation in model abundances convincingly explain these apparent anomalies without recourse to a quasi-Batesian explanation. Nevertheless, the case against classical Mullerian mimicry is not totally disproved, and should be investigated further. I hope that this tentative analysis of actual mimicry rings may encourage others to look for evidence of coevolution and quasi-Batesian effects in a variety of other Mullerian mimicry systems.},
  Keywords = {advertising coloration aposematism batesian mimicry mimicry mutualism signalling warning colour heliconius butterflies natural-selection warning-color hybrid zones muellerian mimicry shifting balance race formation evolution diversity wild},
  Year = {1999}
}
• [1999,article] bibtex
  R. B. Srygley and C. P. Ellington, "Discrimination of flying mimetic, passion-vine butterflies Heliconius," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 266, iss. 1434, pp. 2137-2140, 1999.
  @article{ Author = {Srygley, R. B. and Ellington, C. P.},
  Title = {Discrimination of flying mimetic, passion-vine butterflies Heliconius},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {266},
  Number = {1434},
  Pages = {2137-2140},
  Note = {255YD Times Cited:11 Cited References Count:25},
  Abstract = {Wing-beat frequency and the degree of asymmetry in wing motion were more similar among mimics than among sister species of passion-vine butterflies in the genus Heliconius. Asymmetry in wing motion is not attributed to lift production, and serves as the first clear example of a mimetic behavioural signal for a flying organism. Because the similarities in wing motion are too subtle for humans to observe with the naked eye, they serve as a previously unexplored mimetic signal.},
  Keywords = {locomotor mimicry behaviour mutualism kinematics tropical rainforest signal hawkmoth manduca-sexta mimicry flight evolution temperature morphology kinematics predation mechanics},
  Year = {1999}
}
• [1998,article] bibtex
  D. A. S. Smith, I. J. Gordon, L. A. Depew, and D. F. Owen, "Genetics of the butterfly Danaus chrysippus (L.) in a broad hybrid zone, with special reference to sex ratio, polymorphism and intragenomic conflict," Biological Journal of the Linnean Society, vol. 65, iss. 1, pp. 1-40, 1998.
  @article{ Author = {Smith, D. A. S. and Gordon, I. J. and Depew, L. A. and Owen, D. F.},
  Title = {Genetics of the butterfly Danaus chrysippus (L.) in a broad hybrid zone, with special reference to sex ratio, polymorphism and intragenomic conflict},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {65},
  Number = {1},
  Pages = {1-40},
  Note = {124GE Times Cited:22 Cited References Count:54},
  Abstract = {The aposematic savanna butterfly Danaus chrysippus seems to be an exception to the rule that Batesian models and Mullerian mimics are not polymorphic. Throughout Asia and in much of Africa the species is in fact monomorphic and polytypic, as expected. It is, however, polymorphic for 3-4 colour genes over a large area of central and eastern Africa, where deviant sex ratios and non-Mendelian segregations also occur. All-female broods are widespread and frequent, often outnumbering bisexual progenies and giving rise to heavily female-biased populations. Full and partial sex linkage, which is not controlled from the X or Y chromosomes, and male-biased broods also occur. Genetical analysis for the BC-autosome carrying colour genes suggests there are two: probably mitochondrial, cytotypes (microbe-induced early male death syndrome is considered unlikely) and an autosomal, incompatibility (I) gene, two alleles of which are male-specific killers. F2 and backcross matings by females heterozygous at the I-locus give progenies which are either thelygenic, all males dying at or soon after hatching, or bisexual but showing full or partial sex linkage. Male death is attributed to nuclear-cytoplasmic incompatibility (NCI). Females achieve reversion from a thelygenic to a bisexual line by mating with males of compatible (maternal) cytotype. A second NCI system causes meiotic drive in female oogenesis for a cytoplasmically compatible A-autosome, with no effect on SR. The array or, non-Mendelian segregations for ses and two autosomes, with switching between bisexual and unisexual lineages, indicates autosomal intragenomic conflict with the outcome that one autosome is preserved from elimination in females,and the other condemned in dead males. The geographical distribution of the colour marker genes suggests, that the polymorphisms hale a hybrid origin involving several once allopatric subspecies: some of which hale divergent cytotypes. Secondary contact probably followed range expansion induced by thr periodic spread of savanna during the Pleistocene and massive deforestation by man in recent times. The unusual extent of, the hybrid zone is attributed to the large number of subspecies involved in its origin, and to introgression promoted hy the butterfly's extensive migratory activity. (C) 1998 The Linnean Society of London.},
  Keywords = {allopatric evolution meiotic drive non-mendelian inheritance nuclear cytoplasmic incompatibility pleistocene polytypism sperm displacement thelygeny all-female broods acraea-encedon l lepidoptera heliconius evolution sterility selection africa clines rule},
  Year = {1998}
}
• [1998,techreport] bibtex
  W. O. McMillan and D. G. Heckel, "The genetic architecture of color pattern evolution in Heliconius erato," University of Puerto Rico, N.S.F. Grant Proposal (funded) , 1998.
  @techreport{ Author = {McMillan, W. O. and Heckel, D. G.},
  Title = {The genetic architecture of color pattern evolution in Heliconius erato},
  Institution = {University of Puerto Rico},
  Type = {N.S.F. Grant Proposal (funded)},
  Year = {1998}
}
• [1998,article] bibtex
  J. Mallet, W. O. McMillan, and C. D. Jiggins, "Estimating the mating behavior of a pair of hybridizing Heliconius species in the wild," Evolution, vol. 52, iss. 2, pp. 503-510, 1998.
  @article{ Author = {Mallet, J. and McMillan, W. O. and Jiggins, C. D.},
  Title = {Estimating the mating behavior of a pair of hybridizing Heliconius species in the wild},
  Journal = {Evolution},
  Volume = {52},
  Number = {2},
  Pages = {503-510},
  Note = {Zm270 Times Cited:18 Cited References Count:40},
  Abstract = {Premating isolation between incipient species is rarely studied in nature, even though mating tests in captivity may give an inaccurate picture of natural hybridization. We studied premating barriers between the warningly colored butterflies Heliconius erato and H. himera (Lepidoptera) in a narrow contact zone in Ecuador, where hybrids are found at low frequency. Eggs obtained from wild-mated females, supplemented with eggs and young larvae collected from the wild, were reared to adulthood. Adult color patterns of these progeny were then used to infer how their parents must have mated. Likelihood was used to estimate both the frequencies of potential parental genotypes from adult phenotypes collected in the wild, and the degree of assortative mating from the inferred parents. The frequencies of parental genotypes varied across the hybrid zone, but our statistical method allowed estimates of hybrid deficit and assortative mating to be integrated across all sites sampled. The best estimate of the frequency of F-1 and backcross hybrid adults in the center of the hybrid zone was 10%, with support limits (7.1%, 13.0%; support limits are asymptotically equivalent to 95% confidence limits). Mating was highly assortative: in the center of the hybrid zone the cross-mating probability between H. erato and H. himera was only 5% (0.3%, 21.4%). Wild hybrids themselves mated with both pure forms, and the probabilities that they mated in any direction were not significantly lower than those among conspecifics. These results are consistent with earlier laboratory studies on mate choice, and suggest that selection against hybrids must be strong to prevent formation of a hybrid swarm. Unfortunately, the wide support limits on mating behavior precluded a measure of the strength of selection from these data alone. Our statistical approach provides a useful general method for estimating mate choice in the wild.},
  Keywords = {generalized linear model hybrid zones interspecific hybridization likelihood mate choice reproductive isolation speciation grasshopper podisma-pedestris natural-selection gene flow butterflies mimicry zone evolution erato lepidoptera nymphalidae},
  Year = {1998}
}
• [1998,article] bibtex
  S. Knapp and J. Mallet, "A new species of Passiflora (Passifloraceae) from Ecuador with notes on the natural history of its herbivore, Heliconius (Lepidoptera : Nymphalidae : Heliconiiti)," Novon, vol. 8, iss. 2, pp. 162-166, 1998.
  @article{ Author = {Knapp, S. and Mallet, J.},
  Title = {A new species of Passiflora (Passifloraceae) from Ecuador with notes on the natural history of its herbivore, Heliconius (Lepidoptera : Nymphalidae : Heliconiiti)},
  Journal = {Novon},
  Volume = {8},
  Number = {2},
  Pages = {162-166},
  Note = {104NG Times Cited:1 Cited References Count:3},
  Abstract = {A new species of Passiflora, P. telesiphe, is described in subgenus Decaloba. This species was discovered as a result of observations on the natural history and foodplants of a butterfly (Heliconius telesiphe-Lepidoptera: Nymphalidae: Heliconiiti), whose young stages were previously unknown. The new species and its herbivore are illustrated. The relationships of P. telesiphe to other similar Ecuadorean species of Passiflora are briefly discussed.},
  Year = {1998}
}
• [1998,article] bibtex
  M. Joron and J. L. B. Mallet, "Diversity in mimicry: paradox or paradigm?," Trends in Ecology & Evolution, vol. 13, iss. 11, pp. 461-466, 1998.
  @article{ Author = {Joron, M. and Mallet, J. L. B.},
  Title = {Diversity in mimicry: paradox or paradigm?},
  Journal = {Trends in Ecology & Evolution},
  Volume = {13},
  Number = {11},
  Pages = {461-466},
  Note = {132HZ Times Cited:80 Cited References Count:46},
  Abstract = {Visual mimicry is a textbook case of natural selection because it is both intuitively understandable and has repeatedly evolved in a range of organisms: it is the ultimate example of parallel evolution. In many mimetic groups, particularly butterflies, a huge variety of colour patterns has arisen, even in closely related species. There has been much recent controversy over explanations of this variety. Mimicry is today a broad field of evolutionary study; here we discuss the evolution of its diversity in predator-prey systems.},
  Keywords = {papilio-polyxenes fabr mullerian mimicry batesian mimicry heliconius butterflies limited variability danaus-chrysippus color patterns swallowtail evolution spectrum},
  Year = {1998}
}
• [1998,article] bibtex
  A. V. Z. Brower and R. DeSalle, "Patterns of mitochondrial versus nuclear DNA sequence divergence among nymphalid butterflies: the utility of wingless as a source of characters for phylogenetic inference," Insect Molecular Biology, vol. 7, pp. 73-82, 1998.
  @article{ 6554,
  author = {Brower, A. V. Z. and DeSalle, R.},
  Title = {Patterns of mitochondrial versus nuclear DNA sequence divergence among nymphalid butterflies: the utility of wingless as a source of characters for phylogenetic inference},
  Journal = {Insect Molecular Biology},
  Volume = {7},
  Pages = {73-82},
  Keywords = {HELICONIUS; MTDNA; PHYLOGENY},
  Year = {1998}
}
• [1998,article] bibtex
  C. Grases and N. Ramirez, "Reproduction in five ornythophilous plants of a secondary deciduous forest fragment in Venezuela," Revista De Biologia Tropical, vol. 46, iss. 4, pp. 1095-1108, 1998.
  @article{ Author = {Grases, C. and Ramirez, N.},
  Title = {Reproduction in five ornythophilous plants of a secondary deciduous forest fragment in Venezuela},
  Journal = {Revista De Biologia Tropical},
  Volume = {46},
  Number = {4},
  Pages = {1095-1108},
  Note = {256CJ Times Cited:5 Cited References Count:32},
  Abstract = {The floral biology, the pollination biology and the mating system were studied in two native and three introduced species of an arboretum in Venezuela. All the species presented thin, long, tubular corollas; predominantly red. The antheses were diurnal (between 04:00 and 11:00 hr). Nectar production was high, except for Salvia coccinea (Lamiaceae) (native). Nicoliana glauca (Solanaceae) (native) and Erythrina mitis (Fabaceae) (introduced) shared their hummingbird pollinators, Chlorostilbon mellisugus and Amazilia tobaci (Apodiformes, Throchilidae). S. coccinea was pollinated by Heliconius erato (Lepidoptera). Kalanchoe pinnata and K. daigremontiana (Crassulaceae) (exotic) did not receive any visits. All the species were hermaphrodite. K. daigremontiana and K. pinnata were obligate outcrossers, and S. coccinea, N. glauca and E. mitis were partialy outcrossers and non-autogamus. The species presented high total biomass allocation, compared to other polination systems. The reproductive biomass allocation were allways smaller than the attraction and vegetative biomass allocation. The male biomass allocation were greater than the female biomass allocation in N. glauca, S. coccinea and E. mitis; and smaller in K. daigremontiana and K. pinnata. The introduced species did not seem to affect the reproductive efficiencies of the native species during our observation period. The lowest reproductive efficiencies in introduced species resulted as a consequence of the absence of their natural pollinators.},
  Keywords = {hummingbirds hummingbird pollination deciduous forest biomass allocation reproductive system reproductive eficiency introduced species tropical cloud forest pollination success angiosperms biology predictability interference disturbance dichogamy avoidance patterns},
  Year = {1998}
}
• [1998,article] bibtex
  H. W. Krenn and C. M. Penz, "Mouthparts of Heliconius butterflies (Lepidoptera : Nymphalidae): A search for anatomical adaptations to pollen-feeding behavior," International Journal of Insect Morphology & Embryology, vol. 27, iss. 4, pp. 301-309, 1998.
  @article{ Author = {Krenn, H. W. and Penz, C. M.},
  Title = {Mouthparts of Heliconius butterflies (Lepidoptera : Nymphalidae): A search for anatomical adaptations to pollen-feeding behavior},
  Journal = {International Journal of Insect Morphology & Embryology},
  Volume = {27},
  Number = {4},
  Pages = {301-309},
  Note = {155FH Times Cited:17 Cited References Count:31},
  Abstract = {Proboscis length, the length of the tip, the number and length of the various sensilla throughout the proboscis, and the size and shape of the labial palpi were compared iri 25 species of pollen-feeding and non-pollen-feeding Heliconiinae (Lepidoptera, Nymphalidae). The mouthparts of pollen-feeding species tall belonging ts the genera Heliconius and Laparus) do not have structures exclusive to them. However, in comparison with non-pollen-feeding Heliconiiti, the pollen-feeding species have a significantly longer proboscis without elongation of the tip-region; the bristle-shaped sensilla trichodea were found to be significantly more numerous and longer on the proximal and mid-region of the proboscis, while the sensilla of the tip-region are significantly shorter. In addition to these proboscis features, the labial palpi were shorter in the pollen-feeding species, which is likewise possibly associated with pollen-feeding behavior. The biological role of these features is discussed and the evolution bf this unique feeding behavior among Lepidoptera is considered in the context of the phylogenetic relationships among genera of Heliconiini. (C) 1998 Elsevier Science Ltd. All rights reserved.},
  Keywords = {proboscis galea labial palpus function in food uptake morphology proboscis insecta morphology sensilla biology},
  Year = {1998}
}
• [1998,article] bibtex
  G. Lamas, "Comentarios taxonómicos y nomenclaturales sobre Heliconiini neotropicales, con designación de lectotipos y descripción de cuatro subespecies nuevas (Lepidoptera: Nymphalidae: Heliconiinae)," Revista Peruana de Entomología, vol. 24, pp. 111-125, 1998.
  @article{ Author = {Lamas, G.},
  Title = {Comentarios taxonómicos y nomenclaturales sobre Heliconiini neotropicales, con designación de lectotipos y descripción de cuatro subespecies nuevas (Lepidoptera: Nymphalidae: Heliconiinae)},
  Journal = {Revista Peruana de Entomología},
  Volume = {24},
  Pages = {111-125},
  Abstract = {Description of new form of timareta from Cordillera de Condor and rebuttal of tristero as new species - heurippa, tristero and timareta should be same species},
  Year = {1998}
}
• [1998,incollection] bibtex
  J. Mallet, W. O. McMillan, and C. D. Jiggins, "Mimicry and warning colour at the boundary between races and species," , Howard, D. J. and Berlocher, S. H., Eds., New York: Oxford University Press, 1998, pp. 390-403.
  @incollection{ 6493,
  author = {Mallet, J. and McMillan, W. O. and Jiggins, C. D.},
  Title = {Mimicry and warning colour at the boundary between races and species},
  BookTitle = {Endless Forms: Species and Speciation},
  Editor = {Howard, D. J. and Berlocher, S. H.},
  Publisher = {Oxford University Press},
  Address = {New York},
  Pages = {390-403},
  Keywords = {HELICONIUS; MIMICRY, APOSEMATISM, PREDATOR DEFENCE; SPECIATION, GENERAL},
  Year = {1998}
}
• [1998,article] bibtex
  P. M. Brakefield, "The evolution-development interface and advances with the eyespot patterns of Bicyclus butterflies," Heredity, vol. 80, pp. 265-272, 1998.
  @article{ Author = {Brakefield, P. M.},
  Title = {The evolution-development interface and advances with the eyespot patterns of Bicyclus butterflies},
  Journal = {Heredity},
  Volume = {80},
  Pages = {265-272},
  Note = {Part 3 Ze347 Times Cited:32 Cited References Count:41},
  Abstract = {Studies of the evolution of diversity in colour pattern or morphology have seldom included any analysis of the developmental processes involved in translating genetic variation into the phenotypes influenced by natural selection. This gap is being filled by research on eyespot patterns in the butterfly Bicyclus anynana. This review discusses how data on the developmental genetics of eyespot traits can help in the description and analysis of evolutionary constraints.},
  Keywords = {butterfly development evolution eyespot pattern heritability mutants color pattern heliconius lepidoptera quantitative genetics precis-coenia wings nymphalidae plasticity elements anynana mimicry},
  Year = {1998}
}
• [1998,article] bibtex
  M. I. M. Hernandez and W. W. Benson, "Small-male advantage in the territorial tropical butterfly Heliconius sara (Nymphalidae): a paradoxical strategy?," Animal Behaviour, vol. 56, pp. 533-540, 1998.
  @article{ Author = {Hernandez, M. I. M. and Benson, W. W.},
  Title = {Small-male advantage in the territorial tropical butterfly Heliconius sara (Nymphalidae): a paradoxical strategy?},
  Journal = {Animal Behaviour},
  Volume = {56},
  Pages = {533-540},
  Note = {Part 3 126HG Times Cited:34 Cited References Count:73},
  Abstract = {Large size is often decisive to victory in territorial disputes. Here we report for the first time a natural territorial advantage associated with small size, possibly exemplifying a 'paradoxical strategy' (Maynard Smith Sr Parker 1976, Animal Behaviour, 24, 159-175) in which small individuals with inferior resource holding power win war-of-attrition contests against superior adversaries because resource value/rate of cost accrual (V/K) is greater for small contestants. Males of the aposematic nymphalid butterfly Heliconius sara that defend scattered mating arenas in subtropical Brazilian forest have wings that are on average 3% shorter than males caught away from territories during any part of the year. Smaller residents tend to return to territories over longer periods, and field experiments show that intruders retreat faster when confronting smaller than average territory owners. Heliconius sara has a second, seemingly much more important mating system in which female pupae attract males pheromonally, and in which large males may be more successful in winning mates. Because it is unlikely that small H. sara have intrinsically superior resource holding power, and because territories should be about equally valuable to all males, we propose that large males, supposedly favoured in pupal mating, may risk losing more in terms of future reproductive success through chance injury in territorial fights (large K) and thus avoid combats with small opponents which risk little from injury because of their diminished mating prospects (small K). (C) 1998 The Association for the Study of Animal Behaviour.},
  Keywords = {pamphilus l lepidoptera resource holding power male mating success coenonympha-pamphilus pumpkinseed sunfish sexual dimorphism pararge-aegeria body-size behavior asymmetries},
  Year = {1998}
}
• [1998,article] bibtex
  C. D. Jiggins and N. Davies, "Genetic evidence for a sibling species of Heliconius charithonia (Lepidoptera; Nymphalidae)," Biological Journal of the Linnean Society, vol. 64, iss. 1, pp. 57-67, 1998.
  @article{ Author = {Jiggins, C. D. and Davies, N.},
  Title = {Genetic evidence for a sibling species of Heliconius charithonia (Lepidoptera; Nymphalidae)},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {64},
  Number = {1},
  Pages = {57-67},
  Note = {Zv641 Times Cited:13 Cited References Count:34},
  Abstract = {Heliconius charithonia is a widespread species which, unlike many Heliconius, is non-mimetic and shows little racial differentiation. Only one form, "peruvianus', which occurs in the dry forest habitats of western Ecuador and Peru: has a distinct and clearly mimetic colour pattern. Here it was shown that H. peruvianus was distinct from H. charithonia bassleri at allozyme loci (D = 0.25 over 22 loci). This differentiation was ten limes greater than that between H. charithonia sampled from Ecuador and the Caribbean (D = 0.027) and was consistent with analysis of mitochondrial sequence data (3.4-4% sequence divergence between H. peruvianus and H. charithonia). One individual with a H. peruvianus colour pattern and allozyme genotype was collected in an area where H. charithonia was known to be common, demonstrating that contact between the taxa occurs in western Ecuador Furthermore, the allozyme genotype of another individual was heterozygous for four of fi;I e diagnostic loci and was most likely an F1 hybrid between H. charithonia and H. peruvianus. These data imply that H. charithonia and H. pe,peruvianus are distinct species which hybridize occasionally. This species pair show many similarities with W. erato and H. himera, which are similarly differentiated genetically and also show ecological and colour pattern differences. These species fulfil some of the predictions of both allopatric refugium and parapatric adaptationist models of speciation in the neotropics, suggesting that elements of both hypotheses may be true. (C) 1998 The Linnean Society of London.},
  Keywords = {allopatry butterfly ecuador hybridization hybrid zone parapatry peru speciation butterflies evolution coevolution mimicry races erato},
  Year = {1998}
}
• [1997,article] bibtex
  D. A. S. Smith, D. F. Owen, I. J. Gordon, and N. K. Lowis, "The butterfly Danaus chrysippus (L) in East Africa: Polymorphism and morph-ratio clines within a complex, extensive and dynamic hybrid zone," Zoological Journal of the Linnean Society, vol. 120, iss. 1, pp. 51-78, 1997.
  @article{ Author = {Smith, D. A. S. and Owen, D. F. and Gordon, I. J. and Lowis, N. K.},
  Title = {The butterfly Danaus chrysippus (L) in East Africa: Polymorphism and morph-ratio clines within a complex, extensive and dynamic hybrid zone},
  Journal = {Zoological Journal of the Linnean Society},
  Volume = {120},
  Number = {1},
  Pages = {51-78},
  Note = {Xb934 Times Cited:13 Cited References Count:73},
  Abstract = {Samples of the polymorphic butterfly Danaus chrysippus are analysed from six well separated sites in East Africa. Morph-ratio dines are described for four diallelic genes A, B, C and L, each of which influences the visual phenotype. Each of the four dines has a different orientation, consistent with an hypothesis thai the polymorphism originated from hybridization between a number of polytypic demes which have at Various times undergone range expansion. Allopatric subspeciation in isolated Pleistocene refugia is postulated. The phenotype of each geographical race is shared with one of the morphs within the hybrid zone; other sympatrically maintained polymorphic forms are normally confined to the hybrid zone. Wright's isolation-by-distance model best explains the present distribution of gene frequencies. Morph-ratios differ significantly between the sexes and are sometimes associated with heterozygote excess; gametic and genotypic disequilibria are general throughout the region and suggest the dines are maintained by strong natural selection. Seasonal cycling of phenotype frequency is believed to result from extensive migratory movements rather than natural selection. Female-biased sex-ratio, which is also seasonal, and Haldane rule effects, result from hybrid breakdown when genetically distinct demes meet and interbreed. Oscillating sex-ratios and frequency of colour genes are functionally linked by negative feedback. The polymorphism owes its origin to allopatric evolution but is now maintained sympatrically. (C) 1997 The Linnean Society of London.},
  Keywords = {allopatry heterozygote excess linkage disequilibrium migration mimicry pleistocene polytypism sex-ratio subspeciation all-female broods acraea-encedon l lepidoptera heliconius butterflies natural-selection nymphalidae evolution mimicry},
  Year = {1997}
}
• [1997,incollection] bibtex
  M. Linares, "Origin of Neotropical mimetic biodiversity from a three-way hybrid zone of Heliconius butterflies," , Ulrich, H., Ed., Bonn: Zoologishunginstitut und Museum Alexander Koenig, 1997.
  @incollection{ Author = {Linares, M.},
  Title = {Origin of Neotropical mimetic biodiversity from a three-way hybrid zone of Heliconius butterflies},
  BookTitle = {Proceedings of the International Symposium on Biodiversity and Systematics in Tropical Ecosystems, Bonn 1994},
  Editor = {Ulrich, H.},
  Publisher = {Zoologishunginstitut und Museum Alexander Koenig},
  Address = {Bonn},
  Keywords = {MIMICRY, APOSEMATISM, PREDATOR DEFENCE; HELICONIUS; HYBRID ZONES, CLINES (FACT)},
  Year = {1997}
}
• [1997,article] bibtex
  G. Beccaloni, "Vertical stratification of ithomiine butterfly (Nymphalidae: Ithomiinae) mimicry complexes: The relationship between adult flight height and larval host-plant height," Biological Journal of the Linnean Society, vol. 62, pp. 313-341, 1997.
  @article{ 5804,
  author = {Beccaloni, G.},
  Title = {Vertical stratification of ithomiine butterfly (Nymphalidae: Ithomiinae) mimicry complexes: The relationship between adult flight height and larval host-plant height},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {62},
  Pages = {313-341},
  Keywords = {HELICONIUS; ITHOMIINAE; LEPIDOPTERA LIFE HISTORIES; MIMICRY, APOSEMATISM, PREDATOR DEFENCE},
  Year = {1997}
}
• [1997,article] bibtex
  M. Linares, "The ghost of mimicry past: Laboratory reconstitution of an extinct butterfly ‘race’," Heredity, vol. 78, pp. 628-635, 1997.
  @article{ Author = {Linares, M.},
  Title = {The ghost of mimicry past: Laboratory reconstitution of an extinct butterfly 'race'},
  Journal = {Heredity},
  Volume = {78},
  Pages = {628-635},
  Note = {Part 6 Xe873 Times Cited:6 Cited References Count:39},
  Abstract = {Four variable traits that determine mimetic colour patterns in the butterfly, Heliconius cydno, evolved between 1908 and 1984-91. There was a decline in the frequencies of alleles and phenotypes that confer resemblance to the co-mimic, Elzunia humboldt regalis, and an increase in the frequencies of alleles and phenotypes that confer resemblance to the alternative co-mimic, Heliconius erato chestertonii. Elzunia humbolt regalis was formerly common but is now restricted to forest fragments, whereas H. e. chestertonii occurs principally in disturbed habitats, where it is now common. Human disturbance of habitats is thought to have changed the relative abundances of the two co-mimics, and hence the selection operating on H. cydno. The form of H. cydno that is presumed to have been a near-perfect mimic of H. h. regalis is no longer found in the wild: this form would have been homozygous for the rarer alleles at all four loci. It has been possible to reconstruct this form on two occasions in the laboratory, breeding from partially heterozygous wild-caught female butterflies.},
  Keywords = {butterfly mimicry heliconius microevolution mimicry evolution mullerian mimicry population genetics rapid microevolution host range evolution heliconius butterflies muellerian mimicry warning color hybrid zones genetics erato melpomene},
  Year = {1997}
}
• [1997,article] bibtex
  C. D. Jiggins, W. O. McMillan, P. King, and J. Mallet, "The maintenance of species differences across a Heliconius hybrid zone," Heredity, vol. 79, pp. 495-505, 1997.
  @article{ Author = {Jiggins, C. D. and McMillan, W. O. and King, P. and Mallet, J.},
  Title = {The maintenance of species differences across a Heliconius hybrid zone},
  Journal = {Heredity},
  Volume = {79},
  Pages = {495-505},
  Note = {Part 5 Yf509 Times Cited:34 Cited References Count:48},
  Abstract = {A contact zone between Heliconius erato and H. himera in southern Ecuador provides an opportunity to study the transition from races to species in Heliconiine butterflies. Genetic differentiation at 30 allozyme loci (D = 0.28) is five times greater between himera and erato than among races of erato (D<0.062). Analysis of restriction fragment length polymorphisms in an 800-bp region of the mitochondrial genome shows fixed differences between the species. Despite 5-10 per cent hybridization per generation, these mtDNA, allozyme and colour pattern differences are in almost complete linkage disequilibrium throughout the contact zone. In mixed populations, there was no consistent evidence for convergence of himera and erato allozyme allele frequencies, and only four individuals out of 383 examined showed evidence for interspecific mtDNA gene flow. Linkage analysis of backcross broods showed that the 11 allozyme and two colour pattern loci, which are markedly divergent between the species, map to eight of 21 chromosomal linkage groups. Therefore, barriers to gene flow are not restricted to just a few strongly selected loci. Although analysis of population structure shows little evidence of interspecific gene flow, strong differences between allozyme loci in levels of divergence suggest that selection and gene flow may affect loci in different ways. Hybrid zones such as this, in which divergent genotypes coexist, should provide good model systems for the study of speciation.},
  Keywords = {allozymes interspecific hybridization mitochondrial DNA parapatry population structure speciation fire-bellied toads mitochondrial-DNA biochemical genetics warning-color chorthippus-parallelus natural-selection bombina-bombina evolution butterflies lepidoptera},
  Year = {1997}
}
• [1997,article] bibtex
  V. French, "Pattern formation in colour on butterfly wings," Current Opinion in Genetics & Development, vol. 7, iss. 4, pp. 524-529, 1997.
  @article{ Author = {French, V},
  Title = {Pattern formation in colour on butterfly wings},
  Journal = {Current Opinion in Genetics & Development},
  Volume = {7},
  Number = {4},
  Pages = {524-529},
  Abstract = { Butterflies have uniquely vivid and varied wing-colour patterns. Recent studies show that their formation involves novel expression of homologues of Drosophila appendage-patterning genes, notably Distal-less. Pattern diversity can arise at several developmental stages, either through mutation or in response to environmental conditions. Many loci are known (e.g. in Heliconius) that have major effects on the colour pattern and these invite molecular analysis.},
  Year = {1997}
}
• [1997,article] bibtex
  A. V. Z. Brower and M. G. Egan, "Cladistic analysis of Heliconius butterflies and relatives (Nymphalidae: Heliconiiti): a revised phylogenetic position for Eueides based on sequences from mtDNA and a nuclear gene," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 264, iss. 1384, pp. 969-977, 1997.
  @article{ Author = {Brower, A. V. Z. and Egan, M. G.},
  Title = {Cladistic analysis of Heliconius butterflies and relatives (Nymphalidae: Heliconiiti): a revised phylogenetic position for Eueides based on sequences from mtDNA and a nuclear gene},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {264},
  Number = {1384},
  Pages = {969-977},
  Note = {Xp131 Times Cited:50 Cited References Count:39},
  Abstract = {A new phylogenetic hypothesis for Heliconius and related genera is presented, based on DNA sequence data from mtDNA combined with a region of the wingless gene. This study also adds eight new taxa to a previous cladistic hypothesis based on the mtDNA alone. Simultaneous phylogenetic analysis of the two gene regions together supports a topology largely in agreement with traditional views of heliconiine relationships based on morphology and suggests that the mtDNA support for the sister relationship between Eueides and H. charithonia is due to convergent evolution of homoplasious mtDNA sites.},
  Keywords = {support},
  Year = {1997}
}
• [1997,book] bibtex
  P. J. DeVries, The Butterflies of Costa Rica and Their Natural History, Volume I: Papilionidae, Pieridae, Nymphalidae, Baskerville, USA: Princeton University Press, 1997, vol. I.
  @book{ Author = {DeVries, P. J.},
  Title = {The Butterflies of Costa Rica and Their Natural History, Volume I: Papilionidae, Pieridae, Nymphalidae},
  Publisher = {Princeton University Press},
  Address = {Baskerville, USA},
  Volume = {I},
  Year = {1997}
}
• [1997,article] bibtex
  C. D. Jiggins and W. O. McMillan, "The genetic basis of an adaptive radiation: warning colour in two Heliconius species," Proceedings of the Royal Society of London Series B-Biological Sciences, vol. 264, iss. 1385, pp. 1167-1175, 1997.
  @article{ Author = {Jiggins, C. D. and McMillan, W. O.},
  Title = {The genetic basis of an adaptive radiation: warning colour in two Heliconius species},
  Journal = {Proceedings of the Royal Society of London Series B-Biological Sciences},
  Volume = {264},
  Number = {1385},
  Pages = {1167-1175},
  Note = {Xu828 Times Cited:20 Cited References Count:19},
  Abstract = {Mimetic colour pattern races of Heliconius butterflies provide a striking example of adaptive radiation and numerous crossing experiments have investigated the genetics of these racial differences. However, colour pattern differentiation between closely related Heliconius species has not been previously studied. Here we present data from crosses between H. erato cyrbia and its sister species, H. himera. The genetic architecture underlying colour pattern divergence between these species is identical to that observed between races of H. erato. As in inter-racial crosses, colour pattern differences resulted from segregation at a few major loci. Evidence from 1321 offspring in 4 F-1; 17 backcross, 7 F-2 and 21 further crosses showed that two major loci controlled most of the colour pattern differences between H. erato and H. himera. There were strong interactions between these loci in their patterns of expression and evidence for other loci with relatively minor phenotypic effects. More importantly, based on patterns of expression within broods and linkage with Aconitase, we conclude that these major loci were homologous with those known to be responsible for colour pattern differences within H. erato. Our crosses also permit a re-evaluation of the relationships between colour pattern races of H. erato. This suggests that H. e. hydara, which occurs across a major mtDNA break, is the ancestral phenotype from which other races have evolved. Based on this assumption, we find no evidence to support the recent suggestion that apparently homologous colour pattern alleles have arisen multiple times.},
  Keywords = {hybrid zones evolution butterflies mimicry patterns erato lepidoptera nymphalidae},
  Year = {1997}
}
• [1997,article] bibtex
  C. L. Boggs, "Reproductive allocation from reserves and income in butterfly species with differing adult diets," Ecology, vol. 78, iss. 1, pp. 181-191, 1997.
  @article{ Author = {Boggs, C. L.},
  Title = {Reproductive allocation from reserves and income in butterfly species with differing adult diets},
  Journal = {Ecology},
  Volume = {78},
  Number = {1},
  Pages = {181-191},
  Note = {Wd345 Times Cited:62 Cited References Count:30},
  Abstract = {Allocation of stored and incoming nutrients to reproduction determines an organism's age-specific fecundity curve. In holometabolous insects, differences among species in the shape of the curve are correlated with differences in the potential importance of adult food to reproduction. I examined allocation patterns underlying this association. Specific changes throughout life in body mass and reproductive effort were predicted to result from use of stored vs. incoming nutrients for reproduction and other metabolic needs at each age. Data for three nymphalid butterfly species, Euphydryas editha, Speyeria mormonia, and Heliconius charitonius, were compared with the predictions. These three species differ in adult diet and fraction of oocytes mature at adult emergence (hence, potential for adult nutrients to be used to make eggs), with E. editha showing the least potential for use of adult nutrients in egg production and H. charitonius showing the greatest potential. For all three species, body mass declined with age, although nonlinearly for E. editha. This indicated that metabolic expenditures were greater than intake at all ages, and that a constant fraction of stored nutrients was allocated to reproduction and other metabolic uses at each age for E. editha. Reproductive effort also declined with age for all three species. The specific patterns seen suggested that incoming nutrients may be stored, to some extent, early in life and then used late in life by both S. mormonia and H. charitonius. The similarity between S. mormonia and H. charitonius is rather surprising, given the qualitative differences in adult diet and suggests either that qualitative age-specific allocation patterns for incoming vs. stored nutrients may be independent of adult diet quality, or that the observed patterns are constrained by phylogenetic relatedness of these two species.},
  Keywords = {age-specific body mass age-specific fecundity nutrient reserves nymphalidae ovarian dynamics reproductive allocation reproductive effort euphydryas-editha life-history model dynamics},
  Year = {1997}
}
• [1997,article] bibtex
  W. O. McMillan, C. D. Jiggins, and J. Mallet, "What initiates speciation in passion-vine butterflies?," Proceedings of the National Academy of Sciences of the United States of America, vol. 94, iss. 16, pp. 8628-8633, 1997.
  @article{ Author = {McMillan, W. O. and Jiggins, C. D. and Mallet, J.},
  Title = {What initiates speciation in passion-vine butterflies?},
  Journal = {Proceedings of the National Academy of Sciences of the United States of America},
  Volume = {94},
  Number = {16},
  Pages = {8628-8633},
  Note = {Xq124 Times Cited:55 Cited References Count:39},
  Abstract = {Studies of the continuum between geographic races and species provide the clearest insights into the causes of speciation, Here we report on mate choice and hybrid viability experiments in a pair of warningly colored butterflies, Heliconius erato and Heliconius himera, that maintain their genetic integrity in the face of hybridization. Hybrid sterility and inviability have been unimportant in the early stages of speciation of these two Heliconius. We find no evidence of reduced fecundity, egg hatch, or larval survival nor increases in developmental time in three generations of hybrid crosses, Instead, speciation in this pair appears to have been catalyzed by the association of strong mating preferences with divergence in warning coloration and ecology, In mate choice experiments, matings between the two species are a tenth as likely as matings within species. FI hybrids of both sexes mate frequently with both pure forms, However, male F1 progeny from crosses between H. himera mothers and N. erato fathers have somewhat reduced mating success, The strong barrier to gene flow provided by divergence in mate preference is probably enhanced by frequency-dependent predation against hybrids similar to the type known to occur across interracial hybrid zones of H. erato, In addition, the transition between this pair falls at the boundary between wet and dry forest, and I are hybrids may also be selected against because they are poorly adapted to either biotope, These results add to a growing body of evidence that challenge the importance of genomic incompatibilities in the earliest stages of speciation.},
  Keywords = {heliconius speciation mating behavior postmating isolation interspecific hybridization heliconius butterflies hybrid zones gene flow drosophila evolution selection hybridization adaptation radiation patterns},
  Year = {1997}
}
• [1997,article] bibtex
  A. V. Z. Brower, "The evolution of ecologically important characters in Heliconius butterflies (Lepidoptera: Nymphalidae): A cladistic review," Zoological Journal of the Linnean Society, vol. 119, iss. 4, pp. 457-472, 1997.
  @article{ Author = {Brower, A. V. Z.},
  Title = {The evolution of ecologically important characters in Heliconius butterflies (Lepidoptera: Nymphalidae): A cladistic review},
  Journal = {Zoological Journal of the Linnean Society},
  Volume = {119},
  Number = {4},
  Pages = {457-472},
  Note = {Wz698 Times Cited:9 Cited References Count:79},
  Abstract = {The biology of Heliconius butterflies has provided a rich source of data to test theories of ecological genetics, coevolution and community ecology. Many putatively adaptive characters have been discussed with reference to a phylogenetic hypothesis based on a variety of morphological and life-history traits interpreted from an evolutionary taxonomic perspective. Here, alternate interpretations of characters on the traditional tree and a more recent mitochondrial DNA cladogram with a substantially different topology are compared and contrasted. It is shown that many characters ostensibly providing support for the traditional phylogenetic hypothesis are almost equally parsimoniously distributed and in some cases more parsimoniously distributed on the mtDNA tree than on the tree inferred from those characters. Discussion of alternate evolutionary scenarios based on the mDNA-based topology is presented for pupal mating, pollen feeding, foodplant coevolution, and other ecologically significant features. (C) 1997 The Linnean Society of London.},
  Keywords = {coevolution passiflora mullerian mimicry mitochondrial DNA sequence patterns biology convergence morphology radiation mimicry trees},
  Year = {1997}
}
• [1997,article] bibtex
  C. D. Jiggins, W. O. McMillan, and J. Mallet, "Host plant adaptation has not played a role in the recent speciation of Heliconius himera and Heliconius erato," Ecological Entomology, vol. 22, iss. 3, pp. 361-365, 1997.
  @article{ Author = {Jiggins, C. D. and McMillan, W. O. and Mallet, J.},
  Title = {Host plant adaptation has not played a role in the recent speciation of Heliconius himera and Heliconius erato},
  Journal = {Ecological Entomology},
  Volume = {22},
  Number = {3},
  Pages = {361-365},
  Note = {Xw011 Times Cited:7 Cited References Count:33},
  Keywords = {coevolution heliconius host plant shifts passiflora speciation apple maggot fly rhagoletis-pomonella butterflies evolution races hypothesis},
  Year = {1997}
}
• [1997,phdthesis] bibtex
  C. D. Jiggins, "The ecology and genetics of speciation in Heliconius butterflies," PhD Thesis , 1997.
  @phdthesis{ Author = {Jiggins, C. D.},
  Title = {The ecology and genetics of speciation in Heliconius butterflies},
  School = {University College London},
  Type = {PhD},
  Keywords = {SPECIATION, GENERAL; ECOLOGICAL SPECIATION; ALLOZYMES;},
  Year = {1997}
}
• [1997,article] bibtex
  I. Kajanto, "Emmanuel Swedenborg, ‘Ludus Heliconius’ and other Latin poems. Edited, with introduction, translation and commentary.," Latomus, vol. 56, iss. 4, pp. 905-907, 1997.
  @article{ Author = {Kajanto, I.},
  Title = {Emmanuel Swedenborg, 'Ludus Heliconius' and other Latin poems. Edited, with introduction, translation and commentary.},
  Journal = {Latomus},
  Volume = {56},
  Number = {4},
  Pages = {905-907},
  Note = {Zl642 Times Cited:0 Cited References Count:2 },
  Year = {1997}
}
• [1996,article] bibtex
  C. D. Jiggins, W. O. McMillan, W. Neukirchen, and J. Mallet, "What can hybrid zones tell us about speciation? The case of Heliconius erato and H-himera (Lepidoptera: Nymphalidae)," Biological Journal of the Linnean Society, vol. 59, iss. 3, pp. 221-242, 1996.
  @article{ Author = {Jiggins, C. D. and McMillan, W. O. and Neukirchen, W. and Mallet, J.},
  Title = {What can hybrid zones tell us about speciation? The case of Heliconius erato and H-himera (Lepidoptera: Nymphalidae)},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {59},
  Number = {3},
  Pages = {221-242},
  Note = {Vu424 Times Cited:37 Cited References Count:62},
  Abstract = {To understand speciation we need to study the genetics and ecology of intermediate cases where interspecific hybridization still occurs. Two closely related species of Heliconius butterflies meet this criterion: Heliconius himera is endemic to dry forest and thorn scrub in southern Ecuador and northern Peru, while its sister species, H. erato, is ubiquitous in wet forest throughout south and central America. In three known zones of contact, the two species remain distinct, while hybrids are found at low frequency. Collections in southern Ecuador show that the contact zone is about 5 km wide, half the width of the narrowest dines between colour pattern races of H. erato. The narrowness of this dine argues that very strong selection (s approximate to 1) is maintaining the parapatric distributions of these two species. The zone is closely related with a habitat transition from wet to dry forest, which suggests that the narrow zone of parapatry is maintained primarily by ecological adaptation. Selection on colour pattern loci, assortative mating and hybrid inviability may also be important. The genetics of hybrids between the two species shows that the major gene control of pattern elements is similar to that found in previous studies of H. erato races, and some of the loci are homologous. This suggests that similar generic processes are involved in the morphological divergence of species and races. Evidence from related Heliconius supports a hypothesis that ecological adaptation is the driving force for speciation in the group. (C) 1996 The Linnean Society of London},
  Keywords = {parapatry passiflora refugium biogeography hybridization fire-bellied toads warning-color bombina-bombina gene flow butterflies evolution selection mitochondrial hybridization convergence},
  Year = {1996}
}
• [1996,article] bibtex
  D. L. Grodnitsky and R. Dudley, "Vortex visualization during free flight of heliconiine butterflies (Lepidoptera: Nymphalidae)," Journal of the Kansas Entomological Society, vol. 69, iss. 2, pp. 199-203, 1996.
  @article{ Author = {Grodnitsky, D. L. and Dudley, R.},
  Title = {Vortex visualization during free flight of heliconiine butterflies (Lepidoptera: Nymphalidae)},
  Journal = {Journal of the Kansas Entomological Society},
  Volume = {69},
  Number = {2},
  Pages = {199-203},
  Note = {Ve557 Times Cited:0 Cited References Count:13},
  Abstract = {An experimental procedure was developed to visualize the vortex wake of free-flying insects. Neotropical Heliconius erato butterflies were filmed hovering inside a transparent plexiglas cube. A dispersed cloud of Sephadex particles was used as the visualization medium. The flow field around the flying insect was illuminated with a planar laser beam, thereby generating a cross-sectional perspective of the three-dimensional vortex wake. Thirty-seven flight sequences from five individuals were filmed for analysis. A variety of vortex structures was observed, although the most likely form of the wake in slow forward flight is a series of distinct vortex rings.},
  Keywords = {insect flight tethered flight wake},
  Year = {1996}
}
• [1996,article] bibtex
  J. B. Losos, "Phylogenetic perspectives on community ecology," Ecology, vol. 77, pp. 1344-1354, 1996.
  @article{ Author = {Losos, J. B.},
  Title = {Phylogenetic perspectives on community ecology},
  Journal = {Ecology},
  Volume = {77},
  Pages = {1344-1354},
  Year = {1996}
}
• [1996,article] bibtex
  M. Linares, "The genetics of the mimetic coloration in the butterfly Heliconius cydno weymeri," Journal of Heredity, vol. 87, iss. 2, pp. 142-149, 1996.
  @article{ Author = {Linares, M.},
  Title = {The genetics of the mimetic coloration in the butterfly Heliconius cydno weymeri},
  Journal = {Journal of Heredity},
  Volume = {87},
  Number = {2},
  Pages = {142-149},
  Note = {Ug536 Times Cited:7 Cited References Count:33},
  Keywords = {warning-color hybrid zones muellerian mimicry natural-selection lepidoptera erato nymphalidae adaptation melpomene evolution},
  Year = {1996}
}
• [1996,article] bibtex
  J. Mallet, C. D. Jiggins, and W. McMillan, "Evolution: Mimicry meets the mitochondrion," Current Biology, vol. 6, iss. 8, pp. 937-940, 1996.
  @article{ Author = {Mallet, J. and Jiggins, C. D. and McMillan, W.},
  Title = {Evolution: Mimicry meets the mitochondrion},
  Journal = {Current Biology},
  Volume = {6},
  Number = {8},
  Pages = {937-940},
  Note = {Vb225 Times Cited:6 Cited References Count:15},
  Abstract = {A recent molecular study of the evolution of mimicry in tropical butterflies of the genus Heliconius proves that the mimics adapted to previously diverged 'model' species, but does not clearly distinguish between opposing views of how the model species diverged.},
  Year = {1996}
}
• [1996,article] bibtex
  A. V. Z. Brower, "Parallel race formation and the evolution of mimicry in Heliconius butterflies: A phylogenetic hypothesis from mitochondrial DNA sequences," Evolution, vol. 50, iss. 1, pp. 195-221, 1996.
  @article{ Author = {Brower, A. V. Z.},
  Title = {Parallel race formation and the evolution of mimicry in Heliconius butterflies: A phylogenetic hypothesis from mitochondrial DNA sequences},
  Journal = {Evolution},
  Volume = {50},
  Number = {1},
  Pages = {195-221},
  Note = {Tx891 Times Cited:90 Cited References Count:109},
  Abstract = {Mimicry has been a fundamental focus of research since the birth of evolutionary biology yet rarely has been studied from a phylogenetic perspective beyond the simple recognition that mimics are not similar due to common descent. The difficulty of finding characters to discern relationships among closely related and convergent taxa has challenged systematists for more than a century. The phenotypic diversity of wing pattens among mimetic Heliconius adds an additional twist to the problem, because single species contain more than a dozen radically different-looking geographical races even though the mimetic advantage is theoretically highest when all individuals within and between species appear the same. Mitochondrial DNA (mtDNA) offers an independent way to address these issues. In this study, Cytochrome Oxidase I and II sequences from multiple, parallel races of Heliconius erato and Heliconius melpomene are examined, to estimate intraspecific phylogeny and gauge sequence divergence and ages of clades among races within each species. Although phenotypes of sympatric races exhibit remarkable concordance between the two species, the mitochondrial cladograms show that the species have not shared a common evolutionary history. H. erato exhibits a basal split between trans- and cis-Andean groups of races, whereas H. melpomene originates in the Guiana Shield. Diverse races in either species appear to have evolved within the last 200,000 yr, and convergent phenotypes have evolved independently within as well as between species. These results contradict prior theories of the evolution of mimicry based on analysis of wing-pattern genetics.},
  Keywords = {butterfly wing patterns heliconius erato heliconius melpomene mimicry phylogeny mtdna pleistocene refugium vicariance biogeography warning-color hybrid zones biogeographic relationships historical biogeography hawaiian drosophila natural-populations gene flow speciation patterns birds},
  Year = {1996}
}
• [1996,incollection] bibtex
  J. Mallet, "The genetics of diversity at and below the species level," , Gaston, K. J., Ed., Oxford: Blackwell, 1996, p. 000.
  @incollection{ 5821,
  author = {Mallet, J.},
  Title = {The genetics of diversity at and below the species level},
  BookTitle = {Biodiversity: Biology of Numbers and Difference},
  Editor = {Gaston, K. J.},
  Publisher = {Blackwell},
  Address = {Oxford},
  Pages = {000},
  Keywords = {HELICONIUS; POPULATION STRUCTURE (GENES); SPECIES CONCEPTS},
  Year = {1996}
}
• [1996,article] bibtex
  A. V. L. Freitas and P. S. Oliveira, "Ants as selective agents on herbivore biology: Effects on the behaviour of a non-myrmecophilous butterfly," Journal of Animal Ecology, vol. 65, iss. 2, pp. 205-210, 1996.
  @article{ Author = {Freitas, A. V. L. and Oliveira, P. S.},
  Title = {Ants as selective agents on herbivore biology: Effects on the behaviour of a non-myrmecophilous butterfly},
  Journal = {Journal of Animal Ecology},
  Volume = {65},
  Number = {2},
  Pages = {205-210},
  Note = {Ub621 Times Cited:34 Cited References Count:40},
  Abstract = {1. Larvae of the nymphalid butterfly Eunica bechina feed on young leaves of Caryocar brasiliense, a shrub of the Brazilian savannah that is frequently visited by nectar-gathering ants. 2. Eggs are not removed by ants, but ant occupation on plants affects the oviposition behaviour of the butterflies. Adult females avoid laying eggs on highly visited plant locations and visual cues were demonstrated to mediate oviposition by the butterflies. Plant branches with artificial rubber ants were significantly less infested than control branches with rubber circles. This is the first demonstration that ant presence per se can be enough to produce an avoidance response by ovipositing females in a non-myrmecophilous butterfly. 3. Larval mortality was strongly affected by the level of ant visitation to the host plants, and vulnerability to ant predation decreased with larval size. 4. Stick-like frass chains constructed by the larvae at leaf margins were demonstrated to be a safe refuge against ant attacks on the host plant. Live termites placed on leaves were attacked by foraging ants in significantly greater numbers than those placed on the frass chains. 5. It is concluded that the behavioural biology of both immature and adult Eunica bechina is finely linked with the utilization of a host plant where ant visitation patterns strongly affect larval survival.},
  Keywords = {ant-butterfly interaction ant predation extrafloral nectaries oviposition behaviour plant-herbivore relationships extrafloral nectaries membracid mutualism protection consequence heliconius plants evolution predation defense},
  Year = {1996}
}
• [1996,article] bibtex
  J. R. G. Turner and J. L. B. Mallet, "Did forest islands drive the diversity of warningly coloured butterflies? Biotic drift and the shifting balance," Philosophical Transactions of the Royal Society of London Series B-Biological Sciences, vol. 351, iss. 1341, pp. 835-845, 1996.
  @article{ Author = {Turner, J. R. G. and Mallet, J. L. B.},
  Title = {Did forest islands drive the diversity of warningly coloured butterflies? Biotic drift and the shifting balance},
  Journal = {Philosophical Transactions of the Royal Society of London Series B-Biological Sciences},
  Volume = {351},
  Number = {1341},
  Pages = {835-845},
  Note = {Uy365 Times Cited:32 Cited References Count:98},
  Abstract = {Species of the South American butterfly genus Heliconius have undergone remarkably wide racial divergence in their patterns, and most of the resulting races are muellerian mimics. As warning coloration normally imposes stabilizing selection on the pattern, this divergence is much in need of explanation. Two models have been suggested. Brown, Sheppard and Turner proposed that the divergence results from 'mimetic capture', the switching of patterns between adaptive peaks generated by changes in the overall composition of the local biota ('biotic drift') and hence of the mimicry rings to which each species belongs; these changes have in turn been generated by long term patterns of species extinction in island refuges as biota became progressively isolated and contiguous during contraction and expansion of the rain forest during the Pleistocene. An alternative model, proposed by Mallet, is that truly novel colour patterns became established by mutation and random drift, then spreading to become predominant in local areas; subsequently the novel patterns spread over wide areas by the migration of dines. Under this application of Wright's shifting balance model, refuges are not necessary for divergence, and muellerian mimicry evolves after divergence rather than being the driving force for race formation. Although our respective models appear diametrically opposed, the hypotheses are difficult to distinguish and there are broad areas of agreement; in both models there is an initial stochastic event, followed by natural selection for mimicry, and both will operate either in parapatry or allopatry. The diversity of warning patterns is better explained by the shifting balance model, but there are alternative selectionist explanations such as sexual selection.},
  Keywords = {hybrid zones heliconius butterflies gene flow aposematic coloration muellerian mimicry home range evolution selection population erato},
  Year = {1996}
}
• [1996,article] bibtex
  J. L. Stone, "Components of pollination effectiveness in Psychotria suerrensis, a tropical distylous shrub," Oecologia, vol. 107, iss. 4, pp. 504-512, 1996.
  @article{ Author = {Stone, J. L.},
  Title = {Components of pollination effectiveness in Psychotria suerrensis, a tropical distylous shrub},
  Journal = {Oecologia},
  Volume = {107},
  Number = {4},
  Pages = {504-512},
  Note = {Vh672 Times Cited:22 Cited References Count:56},
  Abstract = {In this paper I report components of effectiveness for pollinators of a tropical distylous shrub, Psychotria suerrensis (Rubiaceae), which is visited by a variety of bees, wasps, and butterflies, and by two species of hummingbirds. In the field, I measured the following components of effectiveness: frequency of visits, evenness of visits across plants, and diurnal pattern of visits. I also used flight-cage experiments to compare pollen-transfer abilities of euglossine bees and heliconiid butterflies. Euglossine bees visited more frequently, visited earlier in the day, and visited a higher proportion of plants in the population than did other taxa. In flight cage experiments, bees and butterflies transferred similar amounts of pollen overall, but bees transferred significantly more inter-morph (compatible) pollen. For each component measured, euglossine bees appeared to be the most effective pollinators.},
  Keywords = {distyly euglossine bee heliconiid butterfly pollinator effectiveness psychotria tristylous-pontederia-cordata pollen removal self-incompatibility floral visitors seed set heliconius butterflies reproductive-biology polemonium-viscosum populations rubiaceae},
  Year = {1996}
}
• [1996,article] bibtex
  A. V. Z. Brower, "A new mimetic species of Heliconius (Lepidoptera:Nymphalidae), from southeastern Colombia, revealed by cladistic analysis of mitochondrial DNA sequences," Zoological Journal of the Linnean Society, vol. 116, iss. 3, pp. 317-332, 1996.
  @article{ Author = {Brower, A. V. Z.},
  Title = {A new mimetic species of Heliconius (Lepidoptera:Nymphalidae), from southeastern Colombia, revealed by cladistic analysis of mitochondrial DNA sequences},
  Journal = {Zoological Journal of the Linnean Society},
  Volume = {116},
  Number = {3},
  Pages = {317-332},
  Note = {50},
  Abstract = {A new species of Heliconius and a new geographical race of Heliconius melpomene are described from the vicinity of Mocoa, Dpto. Putumayo, Colombia, based on molecular and morphological characters. The new species, H. tristero, is a close relative of H. cydno, a geographically differentiated species which lacks red coloration and engages in Mullerian mimicry with other blue and yellow Heliconius species in Central and northwestern South America. H. tristero has switched mimetic associations, instead mimicking the local, sympatric forms of two widespread mimetic species, H. erato and H. melpomene. This discovery provides evidence that the splinter species H. heurippa, H. tristero and H. timareta represent phenotypically divergent members of the H. cydno group that are endemic to successive river valleys on the eastern slope of the northern Andean Cordillera. The nominal taxon Heliconius amaryllis bellula Stichel, currently misapplied to both H. tristero and H. melpomene populations from the Mocoa region of Colombia, is considered here to represent a hybrid between H. heurippa and H. tristero. The Mocoa melpomene race is formally named Heliconius melpomene mocaa, new subspecies. (C) 1996 The Linnean Society of London.},
  Keywords = {morphology; Mullerian mimicry; phenotypic variation; mtDNA COIL WARNING COLOR; HYBRID ZONES; BUTTERFLIES; EVOLUTION; GENETICS},
  Year = {1996}
}
• [1996,article] bibtex
  C. E. G. Pinheiro, "Palatability and escaping ability in neotropical butterflies: Tests with wild kingbirds (Tyrannus melancholicus, Tyrannidae)," Biological Journal of the Linnean Society, vol. 59, iss. 4, pp. 351-365, 1996.
  @article{ Author = {Pinheiro, C. E. G.},
  Title = {Palatability and escaping ability in neotropical butterflies: Tests with wild kingbirds (Tyrannus melancholicus, Tyrannidae)},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {59},
  Number = {4},
  Pages = {351-365},
  Note = {Vx816 Times Cited:31 Cited References Count:64},
  Abstract = {The palatability and the ability of neotropicaI butterflies to escape after being detected, attacked and captured by wild kingbirds (Tyrannus melancholicus Vieillot), was investigated by the release of 668 individuals of 98 butterfly species close to the birds, during their usual feeding activities. Most of the butterflies were attacked and eaten. Only the troidine swallowtails (Parides and Battus; Papilionidae) were consistently rejected on taste and elicited aversive behaviours in birds. Most other aposematic and/or mimetic species in the genera Danaus and Lycorea (Danainae), Dione, Eueides and Helicornicus (Heliconiinae), Hgpothyris, Mechanitis and Melinaea (Ithomiinae), Biblis, Callicore and Diaethria (Limenitidinae) were generally eaten. Cryptic and non-mimetic species were always attacked and, if captured, they were also eaten. AU Apaturinae, Charaxinae, Nymphalinae, Hesperidae, most Limenitidinae, Heliconiinae (Agraulis, Dryas, Dryadula and Philaethria) and Papilionidae (Eurytides, Heraclides and Protesilaus) were in this group. Results indicate that the learning process in kingbirds may demand a large mortality in prey populations, even among species generally accepted as unpalatable and aposematic. They also support the assertion that escaping ability and unpalatability evolved in butterflies as alternative strategies to avoid predation by birds. Mimetic relationships among several species are discussed. Evidence for the evolution of aposematism not related to unpalatability, but to escaping ability, was found for two hard-to-catch Morpho species. (C) 1996 The Linnean Society of London},
  Keywords = {tyrant-flycatchers predation butterfly defences aposematism crypsis mimicry overwintering monarch butterflies aposematic coloration individual selection field observations natural-selection evolution mimicry predation flycatchers heliconius},
  Year = {1996}
}
• [1995,article] bibtex
  J. Mallet and L. E. Gilbert, "Why Are There So Many Mimicry Rings – Correlations between Habitat, Behavior and Mimicry in Heliconius Butterflies," Biological Journal of the Linnean Society, vol. 55, iss. 2, pp. 159-180, 1995.
  @article{ Author = {Mallet, J. and Gilbert, L. E.},
  Title = {Why Are There So Many Mimicry Rings - Correlations between Habitat, Behavior and Mimicry in Heliconius Butterflies},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {55},
  Number = {2},
  Pages = {159-180},
  Note = {Rf925 Times Cited:51 Cited References Count:62},
  Abstract = {In the new world tropics there is an extravagant array of sympatric butterfly mimicry rings. This is puzzling under strictly coevolutionary (Mullerian) mimicry: all unpalatable species should converge as Ice-mimics' to the same pattern. If mimicry has usually evolved in unpalatable species by one-sided (Batesian) evolution, however, it is easy to see that mimicry rings centred on different models could remain distinct. If mimicry rings were also segregated by habitat, a diversity of mimicry rings could be stabilized. In this paper we report correlations between behaviour and mimicry of nine unpalatable Heliconius species. It is already known that co-mimics fly in similar habitats, and non-mimics fly in different habitats, although there is much overlap. Contrary to a previous report, we find little difference in flight heights of heliconiine mimicry rings; all species fly from ground level to the canopy. However, co-mimics roost at night in similar habitats and at similar heights above the ground, but in different habitats and at different heights from species in other mimicry rings. Heliconius (especially the erato taxonomic group) are renowned for roosting gregariously; and co-mimics roost gregariously with each other more often than with non-mimics. Gregarious roosting is therefore common between species, as well as within species. There are thus strong links between mimicry and behavioural ecology in Heliconius. The paradoxical correlation between nocturnal roosting and visual mimicry is presumably explained by bird predation at dusk when roosts are forming, or at dawn before they have disbanded. Direct evidence of predation is lacking, but there are high rates of disturbance by birds at these times. These results, together with knowledge of the phylogeny of Heliconius, suggest that species from the melpomene-group of Heliconius have radiated to occupy mimetic niches protected by model species in the Ithomiinae and the erato-group of Heliconius. A variety of sympatric mimicry rings is apparently maintained because key models fail to converge, while more rapidly-evolving unpalatable mimics evolve towards the colour patterns of the models. The maintenance of mimetic diversity would be aided by the habitat and behavioural differences between mimicry rings revealed here, provided that different predators are found in different habitats. This explanation for the maintenance of multiple mimicry rings is more plausible for Heliconius mimicry than alternatives based on visual mating constraints, thermal ecology, or camouflage.},
  Keywords = {behavioral ecology lepidoptera nymphalidae warning coloration cryptic coloration predation sexual behavior gregarious roosting natural-selection lepidoptera nymphalidae biology range flow},
  Year = {1995}
}
• [1995,article] bibtex
  B. Karlsson, "Resource-Allocation and Mating Systems in Butterflies," Evolution, vol. 49, iss. 5, pp. 955-961, 1995.
  @article{ Author = {Karlsson, B.},
  Title = {Resource-Allocation and Mating Systems in Butterflies},
  Journal = {Evolution},
  Volume = {49},
  Number = {5},
  Pages = {955-961},
  Note = {Te990 Times Cited:36 Cited References Count:60},
  Abstract = {A cross-taxonomic comparison of resources allocated to reproductive reserves at adult eclosion reveals that females belonging to polyandrous species receive more ejaculate material and allocate proportionally less of their total reserves to potential reproduction compared to females belonging to monandrous species. These results suggest that adult females of polyandrous species have a higher expected nutrient income and are consistent with the idea that females can benefit from male nutrient donations transferred during mating. Males show the opposite pattern: males of polyandrous species allocate proportionally more to reproduction. This is expected since males in polyandrous species have both proportionally heavier ejaculates and have a higher ejaculative production capacity than do males in monandrous species. Interestingly, adults of the genus Heliconius which can obtain nutrients crucial to reproduction by pollen feeding do not seem to follow these patterns as strong as only nectar-feeding butterflies. Instead, the association between degree of polyandry and resources allocated to reproduction is relaxed.},
  Keywords = {comparative method heliconius lepidoptera mating system nuptial gifts phylogeny polyandry resource allocation sexual size dimorphism male nutrient investment egg weight variation heliconiine butterflies pararge-aegeria paternal investment lepidoptera colias size mass reproduction},
  Year = {1995}
}
• [1995,article] bibtex
  C. J. Bissoondath and C. Wiklund, "Protein-Content of Spermatophores in Relation to Monandry Polyandry in Butterflies," Behavioral Ecology and Sociobiology, vol. 37, iss. 6, pp. 365-371, 1995.
  @article{ Author = {Bissoondath, C. J. and Wiklund, C.},
  Title = {Protein-Content of Spermatophores in Relation to Monandry Polyandry in Butterflies},
  Journal = {Behavioral Ecology and Sociobiology},
  Volume = {37},
  Number = {6},
  Pages = {365-371},
  Note = {Th369 Times Cited:32 Cited References Count:44 Full text not available},
  Abstract = {The evolution of a mating system, and specifically mating frequency, is dependent on the costs and benefits to both sexes of mating once or several times. In butterflies, males transfer a spermatophore that contains both sperm and accessory gland products. Accessory gland-substances contain nutrients which, in some species, females use to increase their reproductive output and longevity. Nutrients contained in these packaged ejaculates represent investment by males in reproduction. Consequently, the nutritional composition of spermatophores may vary depending on the mating system. There are two lines of arguments concerning the evolution of the nutrient content of ejaculates. One hypothesis argues that male nuptial gifts evolved in the context of certainty of paternity and ease of finding mates; thus spermatophores of polyandrous males (with lower certainty of paternity and greater ease of finding mates) should contain less protein than those of monandrous males, since more spermatophores are produced on average. The other hypothesis argues that polyandry evolved in the context of maximization of male transfer of nutrients to females, and hence spermatophores of polyandrous males should contain more protein than those of monandrous males. In an attempt to distinguish between these two hypotheses, we determined how protein content of ejaculates varied with the degree of polyandry in nine species of pierid and two species of satyrid butterflies. We found that both relative ejaculate mass and protein content increased with the degree of polyandry. Hence our results are consistent with the view that polyandry has evolved in the context of male transfer of nutrients to females, and provides another example of a male adaptation to multiple mating in butterflies.},
  Keywords = {lepidoptera spermatophore protein mating system male nutrient investment papilio-machaon l paternal investment heliconius butterflies reproductive output lepidoptera longevity insects matings size},
  Year = {1995}
}
• [1995,article] bibtex
  D. M. Heredia, "Gran irrupci¢n de Eueides vibilia en el campus de la Universidad del Valle, Cali," , 1995.
  @article{ 6357,
  author = {Heredia, M. Dolores},
  Title = {Gran irrupci¢n de Eueides vibilia en el campus de la Universidad del Valle, Cali},
  Note = {on file; ;,},
  Abstract = {feeding on Passiflora arborea and another Passiflora not yet identified, 3 trees sown close together on the campus. Polymorphism in pupal colour, white and melanic. It depends on the substrate.},
  Keywords = {HELICONIUS; LEPIDOPTERA LIFE HISTORIES; PASSIFLORA},
  Year = {1995}
}
• [1995,article] bibtex
  A. M. Scofield, P. Witham, R. J. Nash, G. C. Kite, and L. E. Fellows, "Differentiation of Glycosidase Activity in Some Hemiptera and Lepidoptera by Means of Castanospermine and Other Polyhydroxy Alkaloids," Comparative Biochemistry and Physiology a-Physiology, vol. 112, iss. 1, pp. 197-205, 1995.
  @article{ Author = {Scofield, A. M. and Witham, P. and Nash, R. J. and Kite, G. C. and Fellows, L. E.},
  Title = {Differentiation of Glycosidase Activity in Some Hemiptera and Lepidoptera by Means of Castanospermine and Other Polyhydroxy Alkaloids},
  Journal = {Comparative Biochemistry and Physiology a-Physiology},
  Volume = {112},
  Number = {1},
  Pages = {197-205},
  Note = {Rt286 Times Cited:7 Cited References Count:14},
  Abstract = {Inhibition of glycosidase activity by a ranger of polyhydroxy alkaloids was studied in a number of insect species. Castanosperimine, usually inactive against maltose and sucrose hydrolysis in insects, was an active inhibitor of maltose hydrolysis in aphids and Heliconius melpomone at pH 5.1 but inactive at pH 6.0. Conversely, 2R,5R-dihydroxymethyl-3R,4R-dihydroxypyrrolidine, normally a potent inhibitor of insect sucrose and maltose hydrolysis, was relatively ineffective in the former species at pH 5.1, The data suggest that several separate enzymes or active sites were responsible for maltose and sucrose hydrolysis in aphids and H. melpomone.},
  Keywords = {alkaloid castanospermine glycosidase hemiptera inhibitor lepidoptera polyhydroxyalkaloid swainsonine insect inhibition disaccharidases},
  Year = {1995}
}
• [1995,article] bibtex
  M. R. Weiss, "Associative colour learning in a nymphalid butterfly," Ecological Entomology, vol. 20, pp. 298-301, 1995.
  @article{ 6216,
  author = {Weiss, M. R.},
  Title = {Associative colour learning in a nymphalid butterfly},
  Journal = {Ecological Entomology},
  Volume = {20},
  Pages = {298-301},
  Abstract = {{IAgraulis vanillae}. Butterflies learned that the yellow flowers of {ILantana camara} contained nectar only after experience with red non-nectar containing, and yellow nectar-containing flowers.},
  Keywords = {BEHAVIOURAL ECOLOGY, SOCIOBIOLOGY; HELICONIUS},
  Year = {1995}
}
• [1995,article] bibtex
  L. Passos and M. Sazima, "Reproductive-Biology of the Distylous Manettia-Luteo-Rubra (Rubiaceae)," Botanica Acta, vol. 108, iss. 4, pp. 309-313, 1995.
  @article{ Author = {Passos, L. and Sazima, M.},
  Title = {Reproductive-Biology of the Distylous Manettia-Luteo-Rubra (Rubiaceae)},
  Journal = {Botanica Acta},
  Volume = {108},
  Number = {4},
  Pages = {309-313},
  Note = {Rw749 Times Cited:11 Cited References Count:41},
  Abstract = {The reproductive biology of Manettia luteo-rubra was studied in the coastal montane Atlantic rain forest of southeastern Brazil. This Rubiaceae is a perennial vine that flowers all the year round, but has a flowering peak during the dry season. It presents reciprocal herkogamy, thus the plants are morphologically distylous. The morphs occur in a 1:1 ratio, and pollen diameter and corolla length vary between short and long-styled flowers. Manettia luteo-rubra displays typical heterostylous self-incompatibility and sets almost no fruits from self- or intramorph pollinations. The flowers are tubular, red with yellow lobes, and odourless. Anthesis is asynchronous, and the Bowers last about four days. Concentration of sugars in nectar is similar in both morphs, ca. 24%, this concentration being typical for hummingbird flowers. Three species of hummingbirds are the major pollinators of the flowers of M. luteo-rubra at the study site: the hermits Phaethornis eurynome and P. squalidus, and the trochiline Thalurania glaucopis. Three species of Heliconius butterflies act as minor pollinators. Both morphs of M. luteo-rubra exhibit natu ral fruit-set of about 80%, this reproductive output being maintained throughout the year by the pollinators' constancy to the flowers.},
  Keywords = {heterostyly breeding system floral biology hummingbird pollination butterfly pollination flowering phenology individual flowers butterflies pollination community longevity forest plants},
  Year = {1995}
}
• [1995,article] bibtex
  L. V. Lutz and A. M. Araújo, "Variation and selection on the wing colour pattern of IHeliconius erato phyllis (Lepidoptera: Nymphalidae) in natural populations from Southern Brazil," Biological Journal of the Linnean Society, vol. rejected, pp. 1-27+3figs, 1995.
  @article{ 6400,
  author = {Lutz, L. V. and Araújo, A. M.},
  Title = {Variation and selection on the wing colour pattern of {IHeliconius erato phyllis} (Lepidoptera: Nymphalidae) in natural populations from Southern Brazil},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {rejected},
  Pages = {1-27+3figs},
  Abstract = {Very bad morphological paper},
  Keywords = {HELICONIUS},
  Year = {1995}
}
• [1994,article] bibtex
  A. V. Z. Brower, "Rapid Morphological Radiation and Convergence among Races of the Butterfly Heliconius-Erato Inferred from Patterns of Mitochondrial-DNA Evolution," Proceedings of the National Academy of Sciences of the United States of America, vol. 91, iss. 14, pp. 6491-6495, 1994.
  @article{ Author = {Brower, A. V. Z.},
  Title = {Rapid Morphological Radiation and Convergence among Races of the Butterfly Heliconius-Erato Inferred from Patterns of Mitochondrial-DNA Evolution},
  Journal = {Proceedings of the National Academy of Sciences of the United States of America},
  Volume = {91},
  Number = {14},
  Pages = {6491-6495},
  Note = {Nv420 Times Cited:434 Cited References Count:40},
  Abstract = {The neotropical Heliconius butterflies are famous examples of Mullerian mimicry, due to the diverse array of shared, brightly colored wing patterns that advertise the butterflies' unpalatability. The parallel geographical variation in these patterns within several widespread species has been invoked to support the controversial Pleistocene refugium hypothesis of tropical diversification. However, in no Heliconius species have either evolutionary rates or relationships among geographical races been explicitly examined. I present a phylogenetic hypothesis based on mitochondrial DNA sequences for 14 divergent races of Heliconius erato, which reveals that similar wing patterns have evolved rapidly and convergently within the species. There is a basal split between groups of races from east and west of the Andes, reflecting a vicariant separation at the base of the Pleistocene. Within each of these clades, sequence divergence is very low, and some haplotypes are shared between allopatric races with radically different wing patterns. The topology implies a simultaneous radiation of races in these two areas within the last 200,000 years. Ages for the clades are estimated by comparing sequence divergence to a plot of mitochondrial divergence in several arthropod taxa with independently dated divergence times. This plot is linear and suggests that mitochondrial DNA in arthropods evolves in a clocklike manner, at least initially, when sequence divergence is low.},
  Keywords = {molecular clock pleistocene refugia biogeography mullerian mimicry natural-selection divergence amplification population polymerase mimicry genome rates},
  Year = {1994}
}
• [1994,article] bibtex
  A. V. Z. Brower, "The case of the missing H: Heliconius charithonia (L., 1767), not ‘Heliconius charitonia (L., 1767)’," Journal of the Lepidopterists’ Society, vol. 48, pp. 166-168, 1994.
  @article{ Author = {Brower, A. V. Z.},
  Title = {The case of the missing H: Heliconius charithonia (L., 1767), not 'Heliconius charitonia (L., 1767)'},
  Journal = {Journal of the Lepidopterists' Society},
  Volume = {48},
  Pages = {166-168},
  Year = {1994}
}
• [1994,book] bibtex
  H. Holzinger and R. Holzinger, IHeliconius and related Genera. Lepidoptera: Nymphalidae. The Genera IEueides, INeruda and IHeliconius., Vennette, France: Sciences Nat., 1994.
  @book{ 5224,
  author = {Holzinger, H. and Holzinger, R.},
  Title = {{IHeliconius} and related Genera. Lepidoptera: Nymphalidae. The Genera {IEueides},
  {INeruda} and {IHeliconius}.},
  Publisher = {Sciences Nat.},
  Address = {Vennette, France},
  Abstract = {himera x erato favorinus, m, Rodriguez de Mendoza; himera x erato cyrbia, m, Guayquichuma. Also cydno x melpomene, ethilla x besckei and numata x melpomene hybrids.},
  Keywords = {HELICONIUS; HYBRID ZONES, CLINES (FACT)},
  Year = {1994}
}
• [1994,article] bibtex
  A. Alonso-Mej¡a and M. Marquez, "Dragonfly predation on butterflies in a tropical dry forest," Biotropica, vol. 26, pp. 341-344, 1994.
  @article{ 5925,
  author = {Alonso-Mej¡a, A. and Marquez, M.},
  Title = {Dragonfly predation on butterflies in a tropical dry forest},
  Journal = {Biotropica},
  Volume = {26},
  Pages = {341-344},
  Abstract = {Quite a few butterflies eaten, including especially Pieridae; also a few unpalatable species like Heliconiini. Total no. 205; 14 Agraulis vanillae, 1 Dryadula phaetusa, 2 Dryas iulia, and 1 Heliconius charitonia},
  Keywords = {HELICONIUS; MIMICRY, APOSEMATISM, PREDATOR DEFENCE},
  Year = {1994}
}
• [1994,article] bibtex
  E. I. Deinert, J. T. Longino, and L. E. Gilbert, "Mate competition in butterflies," Nature (London), vol. 370, pp. 23-24, 1994.
  @article{ 5280,
  author = {Deinert, E. I. and Longino, J. T. and Gilbert, L. E.},
  Title = {Mate competition in butterflies},
  Journal = {Nature (London)},
  Volume = {370},
  Pages = {23-24},
  Abstract = {pupal mating and mate competition - effects of size},
  Keywords = {HELICONIUS},
  Year = {1994}
}
• [1994,article] bibtex
  N. E. Rank and J. T. Smiley, "Host-Plant Effects on Parasyrphus-Melanderi (Diptera, Syrphidae) Feeding on a Willow Leaf Beetle Chrysomela-Aeneicollis (Coleoptera, Chrysomelidae)," Ecological Entomology, vol. 19, iss. 1, pp. 31-38, 1994.
  @article{ Author = {Rank, N. E. and Smiley, J. T.},
  Title = {Host-Plant Effects on Parasyrphus-Melanderi (Diptera, Syrphidae) Feeding on a Willow Leaf Beetle Chrysomela-Aeneicollis (Coleoptera, Chrysomelidae)},
  Journal = {Ecological Entomology},
  Volume = {19},
  Number = {1},
  Pages = {31-38},
  Note = {Mv437 Times Cited:17 Cited References Count:42},
  Abstract = {1. Generalist predators are repelled by chrysomelid (Chrysomela spp., Phratora vitellinae L.) larval defensive secretions that are obtained from salicin in their host plants. But little is known about the effect of these secretions on specialist predators. 2. In this study, we describe the feeding behaviour of a fly, Parasyrphus melanderi Curran (Diptera: Syrphidae), which feeds on Chrysomela aeneicollis Schaeffer (Coleoptera: Chrysomelidae). Parasyrphus melanderi lays its eggs on C.aeneicollis egg clutches, and its larvae consume C.aeneicollis eggs and larvae. 3. Chrysomela aeneicollis hatching rates were significantly lower (20%) on clutches with fly eggs than on clutches without them (40%). Half of the clutches with one fly egg had survival rates below 5%, and when two fly eggs were present (four clutches), the entire clutch was consumed. 4. In nature, P.melanderi eggs were 3 times more abundant on a salicylate-rich willow species S. orestera Schneider, than on the medium-salicylate S. geyeriana Anderss. (1.8 v 0.6 eggs per clutch). On 18% of the S.orestera clones, all the beetle clutches contained fly eggs. In laboratory-choice tests, P.melanderi larvae fed equally rapidly on C.aeneicollis larvae that were chemically defended (feeding on S.orestera) as on larvae that produced no secretion (feeding on the salicylate-poor S.lutea Nutt.). This predator does not appear to be deterred by C.aeneicollis's defensive secretion. We discuss the implications of specialist predators on determining host suitability to herbivorous insects.},
  Keywords = {parasyrphus-melanderi syrphidae predation 3-trophic level interactions chrysomela-aeneicollis salix spp 3 trophic levels enemy-free space defensive secretion group-size larvae survivorship heliconius passiflora selection mortality},
  Year = {1994}
}
• [1994,article] bibtex
  L. M. Silva and A. M. Araújo, "The genetic structure of Heliconius erato populations (Lepidoptera; Nymphalidae)," Revista Brasileira de Genetica, vol. 17, pp. 19-24, 1994.
  @article{ 5545,
  author = {Silva, L. M. and Araújo, A. M.},
  Title = {The genetic structure of Heliconius erato populations (Lepidoptera; Nymphalidae)},
  Journal = {Revista Brasileira de Genetica},
  Volume = {17},
  Pages = {19-24},
  Abstract = {5 localities in Rio Grande do Sul. 4 loci: Pgm, Lap, Mdh1, Mdh2 (the latter monomorphic). Very great similarities Fst=0.025, P<0.001). Fis also significant, =0.1859., consisting of Lap 0.28, Mdh1 0.59, Pgm 0.06. (I STILL THINK IT IS SCORING ERRORS!!). Only the Mdh1 results were significant, giving overall significance. "Confirms the island nature of the population previously found on the basis of ecological investigations".},
  Keywords = {HELICONIUS; POPULATION STRUCTURE (GENES)},
  Year = {1994}
}
• [1994,article] bibtex
  J. Vanderplank, "A growing passion," The Garden, vol. 119, pp. 28-33, 1994.
  @article{ 5120,
  author = {Vanderplank, J.},
  Title = {A growing passion},
  Journal = {The Garden},
  Volume = {119},
  Pages = {28-33},
  Note = {reprint in file; ; ;,},
  Abstract = {About Passiflora and hybridization},
  Keywords = {HELICONIUS; PASSIFLORA; SYSTEMATICS, TAXONOMY},
  Year = {1994}
}
• [1994,article] bibtex
  L. Maurico-Da-Silva and A. M. Dearaujo, "The Genetic-Structure of Heliconius-Erato Populations (Lepidoptera, Nymphalidae)," Revista Brasileira De Genetica, vol. 17, iss. 1, pp. 19-24, 1994.
  @article{ Author = {Maurico-Da-Silva, L. and Dearaujo, A. M.},
  Title = {The Genetic-Structure of Heliconius-Erato Populations (Lepidoptera, Nymphalidae)},
  Journal = {Revista Brasileira De Genetica},
  Volume = {17},
  Number = {1},
  Pages = {19-24},
  Note = {Nm227 Times Cited:3 Cited References Count:0},
  Abstract = {Samples of butterflies (Heliconius erato, Nymphalidae) were drawn from five different localities in Rio Grande do Sul (Brazil) to study allozyme polymorphisms and genetic structure. Four loci were investigated, Pgm, Lap, Mdh1 and Mdh2 (the latter being monomorphic). The five localities, which were less than 30 km apart, showed a high genotypic similarity, ranging from 0.86 to 0.95. However, they also showed a significant isolation component when considered as part of a larger population (F(ST) = 0.0247; P < 0.001). When they were taken as forming only two subdivisions of a larger population, F(ST) was even greater, 0.0798 (P < 0.001). The component due to inbreeding (F(IS))) was also significant in both cases, being respectively 0. 1859 (P < 0.05) and 0.2153 (P < 0.001). These results confirm the island nature of H. erato populations previously suggested on the basis of ecological information. The high levels of inbreeding could be associated with certain behavioural traits shown by these insects.},
  Year = {1994}
}
• [1994,article] bibtex
  J. Smiley and L. E. Gilbert, "Habitat segregation in coexisting mimicry complexes," Unpublished manuscript, 1994.
  @article{ 3980,
  author = {Smiley, J. and Gilbert, L. E.},
  Title = {Habitat segregation in coexisting mimicry complexes},
  Journal = {Unpublished manuscript},
  Keywords = {HELICONIUS; MIMICRY, APOSEMATISM, PREDATOR DEFENCE},
  Year = {1994}
}
• [1994,article] bibtex
  G. Holzkamp and A. Nahrstedt, "Biosynthesis of Cyanogenic Glucosides in the Lepidoptera – Incorporation of [U-C-14]-2-Methylpropanealdoxime, 2s-[U-C-14]-Methylbutanealdoxime and D,L-[U-C-14]-N-Hydroxyisoleucine into Linamarin and Lotaustralin by the Larvae of Zygaena-Trifolii," Insect Biochemistry and Molecular Biology, vol. 24, iss. 2, pp. 161-165, 1994.
  @article{ Author = {Holzkamp, G. and Nahrstedt, A.},
  Title = {Biosynthesis of Cyanogenic Glucosides in the Lepidoptera - Incorporation of [U-C-14]-2-Methylpropanealdoxime, 2s-[U-C-14]-Methylbutanealdoxime and D,L-[U-C-14]-N-Hydroxyisoleucine into Linamarin and Lotaustralin by the Larvae of Zygaena-Trifolii},
  Journal = {Insect Biochemistry and Molecular Biology},
  Volume = {24},
  Number = {2},
  Pages = {161-165},
  Note = {Mr834 Times Cited:9 Cited References Count:21},
  Abstract = {[U-C-14]-2-methylpropanealdoxime was incorporated into the cyanogenic glucoside linamarin (1-cyano-1-methylethyl-beta-D-glucopyraraoside) and D,L-[U-C-14]-N-hydroxyisoleucine and 2S-[U-C-14]-methylbutanealdoxime were incorporated into the cyanogenic glucoside lotaustralin (R-1-cyano-1-methylpropyl-beta-D-glucopyranoside) after injection into the haemocoele of larvae of Zygaena trifolii. The incorporation rates of the aldoximes were 17-22% and of the N-hydroxyamino acid 4-5%. The results indicate that both aldoximes and N-hydroxyamino acids are intermediates in the biosynthesis of linamarin and lotaustralin by the larvae by Zygaena trifolii.},
  Keywords = {zygaena trifolii zygaenidae lepidoptera cyanogenic glucosides linamarin lotaustralin biosynthesis aldoxime intermediates n-hydroxyamino acid intermediates 2-methylpropanealdoxime 2s-methylbutanealdoxime d,l-n-hydroxyisoleucine heliconius species lepidoptera sorghum-bicolor invitro biosynthesis dhurrin butterflies glycosides moench plants moths},
  Year = {1994}
}
• [1994,article] bibtex
  B. Karlsson, "Feeding-Habits and Change of Body-Composition with Age in 3 Nymphalid Butterfly Species," Oikos, vol. 69, iss. 2, pp. 224-230, 1994.
  @article{ Author = {Karlsson, B.},
  Title = {Feeding-Habits and Change of Body-Composition with Age in 3 Nymphalid Butterfly Species},
  Journal = {Oikos},
  Volume = {69},
  Number = {2},
  Pages = {224-230},
  Note = {Na703 Times Cited:36 Cited References Count:39},
  Abstract = {Three species of nymphalid butterflies with different types of adult feeding behaviour were examined to see how the allocation of larval-derived resources to abdomen vs thorax change with age. Adults of Pararge aegeria and Speyeria mormonia feed on poor-quality diets. They are expected to be more dependent on larval-derived reserves stored in the abdomen for reproduction than are species with high-quality diets. In accordance with these expectations abdomen mass decreases with age in both males and females. Furthermore, thorax mass of females also decreases with age. Since the bulk of the thorax consists of flight muscles, this suggests that old females use resources derived from flight muscles for egg production. However, since thorax ratio (thorax mass/total mass) increases with age, flight properties such as speed and maneuverability are likely to be unchanged or even improved with advancing age. Adults of Heliconius hecale, which feed on a more rich-quality diet, show a reversed pattern compared with the other two species. In both sexes, abdomen mass and thorax mass increase with age.},
  Keywords = {pararge-aegeria neotropical butterflies resource-allocation reproductive effort oviposition rate egg weight population insects size palatability},
  Year = {1994}
}
• [1994,article] bibtex
  H. F. Nijhout, "Developmental Perspectives on Evolution of Butterfly Mimicry," Bioscience, vol. 44, iss. 3, pp. 148-157, 1994.
  @article{ Author = {Nijhout, H. F.},
  Title = {Developmental Perspectives on Evolution of Butterfly Mimicry},
  Journal = {Bioscience},
  Volume = {44},
  Number = {3},
  Pages = {148-157},
  Note = {My533 Times Cited:16 Cited References Count:43},
  Keywords = {color patterns heliconius lepidoptera nymphalidae genetics constraints homologies},
  Year = {1994}
}
• [1994,article] bibtex
  W. M. Neukirchen, "Heliconius sara williami n. subsp. von Trinidad (West Indies) (Lepidoptera: Nymphalidae)," Entomologische Zeitschrift, vol. 104, pp. 141-160, 1994.
  @article{ 5509,
  author = {Neukirchen, W. M.},
  Title = {Heliconius sara williami n. subsp. von Trinidad (West Indies) (Lepidoptera: Nymphalidae)},
  Journal = {Entomologische Zeitschrift},
  Volume = {104},
  Pages = {141-160},
  Keywords = {HELICONIUS; SYSTEMATICS, TAXONOMY},
  Year = {1994}
}
• [1994,article] bibtex
  M. Baguette and G. N∑ve, "Adult movements between populations in the specialist butterfly Proclossiana eunomia (Lepidoptera, Nymphalidae)," Ecological Entomology, vol. 19, pp. 1-5, 1994.
  @article{ 5148,
  author = {Baguette, M. and N∑ve, G.},
  Title = {Adult movements between populations in the specialist butterfly Proclossiana eunomia (Lepidoptera, Nymphalidae)},
  Journal = {Ecological Entomology},
  Volume = {19},
  Pages = {1-5},
  Abstract = {Females were more likely to disperse between subpopulations than males. However, maximum distances were only of order 500m. Females also emerge approx 10 days after males, but overlap, of course.},
  Keywords = {HELICONIUS; POPULATION STRUCTURE, DISPERSAL (FIELD EXPT)},
  Year = {1994}
}
• [1994,article] bibtex
  K. L. Haag and A. M. Araújo, "Inbreeding, genetic load and morphometric variation in natural populations of IDryas iulia (Lepidoptera, Nymphalidae)," Revista Brasileira de Genetica, vol. 17, pp. 35-39, 1994.
  @article{ 5546,
  author = {Haag, K. L. and Araújo, A. M.},
  Title = {Inbreeding, genetic load and morphometric variation in natural populations of {IDryas iulia} (Lepidoptera, Nymphalidae)},
  Journal = {Revista Brasileira de Genetica},
  Volume = {17},
  Pages = {35-39},
  Abstract = {High inbreeding (F=0.07-0.12) estimated based on the mortality of eggs and inviability.},
  Keywords = {CONSERVATION GENETICS; HELICONIUS; POPULATION STRUCTURE, DISPERSAL (FIELD EXPT)},
  Year = {1994}
}
• [1994,article] bibtex
  A. V. Z. Brower, "Phylogeny of Heliconius butterflies inferred from mitochondrial DNA sequences (Lepidoptera: Nymphalidae)," Molecular Phylogenetics and Evolution, vol. 3, pp. 159-174, 1994.
  @article{ 5218,
  author = {Brower, A. V. Z.},
  Title = {Phylogeny of Heliconius butterflies inferred from mitochondrial DNA sequences (Lepidoptera: Nymphalidae)},
  Journal = {Molecular Phylogenetics and Evolution},
  Volume = {3},
  Pages = {159-174},
  Keywords = {HELICONIUS; MTDNA; PHYLOGENY},
  Year = {1994}
}
• [1994,article] bibtex
  D. A. West, "Unimodal Batesian Polymorphism in the Neotropical Swallowtail Butterfly Eurytides Lysithous (Hbn)," Biological Journal of the Linnean Society, vol. 52, iss. 3, pp. 197-224, 1994.
  @article{ Author = {West, D. A.},
  Title = {Unimodal Batesian Polymorphism in the Neotropical Swallowtail Butterfly Eurytides Lysithous (Hbn)},
  Journal = {Biological Journal of the Linnean Society},
  Volume = {52},
  Number = {3},
  Pages = {197-224},
  Note = {Nt757 Times Cited:3 Cited References Count:74},
  Abstract = {The Brazilian swallowtail Eurytides lysithous is evidently a Batesian mimic of several Parides species. It is polymorphic for mimetic patterns in both sexes. Various populations contain from one to three major forms, and these seem to depend on two unlinked loci or supergenes. Samples from natural populations, and one reared brood, suggest that one locus controls two white-marked forms, with incomplete dominance producing a third heterozygous form. The heterozygotes are everywhere deficient from Hardy-Weinberg expectations. The second putative locus has an allele epistatic to the first locus which converts the white-marked forms to black, but epistasis is apparently incomplete in heterozygotes. The incomplete dominance and epistasis result in extraordinarily variable polymorphic populations and would allow a genetic analysis for comparison with those already done in the classic Batesian polymorphic swallowtails of the Old World.},
  Keywords = {batesian mimicry butterfly wing patterns dominance epistasis eurytides leptocircini color patterns heliconius lepidoptera hypolimnas-misippus papilio-polyxenes nymphalidae homologies genetics mimicry},
  Year = {1994}
}
• [1994,article] bibtex
  M. Burd, "Butterfly Wing Colour Patterns and Flying Heights in the Seasonally Wet Forest of Barro Colorado Island, Panama," Journal of Tropical Ecology, vol. 10, iss. 4, pp. 601-610, 1994.
  @article{ Author = {Burd, M.},
  Title = {Butterfly Wing Colour Patterns and Flying Heights in the Seasonally Wet Forest of Barro Colorado Island, Panama},
  Journal = {Journal of Tropical Ecology},
  Volume = {10},
  Number = {4},
  Pages = {601-610},
  Year = {1994}
}
• [1993,article] bibtex
  P. B. Koch, "Production of [C-14] Labeled 3-Hydroxy-L-Kynurenine in a Butterfly, Heliconius-Charitonia L (Heliconidae), and Precursor Studies in Butterfly Wing Ommatins," Pigment Cell Research, vol. 6, iss. 2, pp. 85-90, 1993.
  @article{ Author = {Koch, P. B.},
  Title = {Production of [C-14] Labeled 3-Hydroxy-L-Kynurenine in a Butterfly, Heliconius-Charitonia L (Heliconidae), and Precursor Studies in Butterfly Wing Ommatins},
  Journal = {Pigment Cell Research},
  Volume = {6},
  Number = {2},
  Pages = {85-90},
  Note = {Ld176 Times Cited:13 Cited References Count:17},
  Abstract = {A method was developed to produce radiolabeled 3-hydroxy-L-kynurenine by injection of [C-14]-L-tryptophan into pupae of the heliconid butterfly, Heliconius charitonia, which was converted into [C-14]-3-hydroxy-L-kynurenine and deposited as a wing pigment. Extractions of 3-hydroxykynurenine (3-OHK) with 60% methanol from wings yielded in 14.4 mug per mg dry weight. In extracts from yellow wing areas, 3-OHK represented 100% of detectable amino acids. Resulting specific radioactivity of [C-14]-3-OHK was between 0.05 and 0.07 mCi/mmol when 0.5 muCi [C-14]-tryptophan was injected into pupae 1 or 2 days before emergence of the butterfly. Incorporation of [C-14]-3-OHK into wing ommochromes was studied in nymphalid butterflies, Araschnia levana and Precis coenia. After injection into pupae [C-14]-3-OHK as well as [C-14]-tryptophan were specifically incorporated into red and red-brown wing scales as shown by autoradiography. The same incorporation occurred in isolated wings after incubation in Grace's medium containing [C-14]-3-OHK. In Araschnia levana, [C-14]-3-OHK offered to left wing pairs was incorporated into dihydroxanthommatin six times more effectively than [C-14]-tryptophan offered to right wing pairs from the same specimen. Therefore, 3-OHK seems to be the ultimate precursor of wing ommatins.},
  Keywords = {3-hydroxykynurenine tryptophan ommatins precursor wings lepidoptera heliconidae nymphalidae araschnia-levana l 3-hydroxykynurenine nymphalidae lepidoptera pattern},
  Year = {1993}
}
• [1993,article] bibtex
  Posla, "An unusual form of IHeliconius cydno from Costa Rica," Tropical Lepidoptera, vol. 4, p. 92, 1993.
  @article{ 5677,
  author = {Posla},
  Title = {An unusual form of {IHeliconius cydno} from Costa Rica},
  Journal = {Tropical Lepidoptera},
  Volume = {4},
  Pages = {92},
  Keywords = {HELICONIUS},
  Year = {1993}
}
• [1993,article] bibtex
  Tanner, "Courtship behavior of IDryadula phaetusa (Nymphalidae)," Tropical Lepidoptera, vol. 4, p. 106, 1993.
  @article{ 5676,
  author = {Tanner},
  Title = {Courtship behavior of {IDryadula phaetusa} (Nymphalidae)},
  Journal = {Tropical Lepidoptera},
  Volume = {4},
  Pages = {106},
  Keywords = {HELICONIUS; SEXUAL BEHAVIOUR},
  Year = {1993}
}
• [1993,article] bibtex
  D. A. Murawski, "A taste for poison," National Geographic Magazine, vol. 184, iss. 6, pp. 122-137, 1993.
  @article{ 5089,
  author = {Murawski, D. A.},
  Title = {A taste for poison},
  Journal = {National Geographic Magazine},
  Volume = {184},
  Number = {6},
  Pages = {122-137},
  Abstract = {About {IHeliconius}. Record for H. sapho from Ecuador laying on Passiflora macrophylla},
  Keywords = {HELICONIUS; INSECT/PLANT RELS (NON-CHEM); MIMICRY, APOSEMATISM, PREDATOR DEFENCE},
  Year = {1993}
}
• [1993,article] bibtex
  J. A. Salazar, "Notes on some populations of Heliconius heurippa in Colombia (Lepidoptera: Nymphalidae: Heliconiinae)," Tropical Lepidoptera, vol. 4, pp. 119-121, 1993.
  @article{ 5507,
  author = {Salazar, J. A.},
  Title = {Notes on some populations of Heliconius heurippa in Colombia (Lepidoptera: Nymphalidae: Heliconiinae)},
  Journal = {Tropical Lepidoptera},
  Volume = {4},
  Pages = {119-121},
  Abstract = {comparison with hybrids of cydno x melpomene. Figures of 3 melpomene x cydno hybrids from Victoria, Caldas, and Otanche, Boyaca. Figure of a new hybrid H.melpomene x H.heurippa (presumably) from the Villavicencio road, Chirajara},
  Keywords = {HELICONIUS; LEPIDOPTERA GENETICS},
  Year = {1993}
}
• [1993,article] bibtex
  C. Wiklund, A. Kaitala, V. Lindfors, and J. Abenius, "Polyandry and Its Effect on Female Reproduction in the Green-Veined White Butterfly (Pieris-Napi L)," Behavioral Ecology and Sociobiology, vol. 33, iss. 1, pp. 25-33, 1993.
  @article{ Author = {Wiklund, C. and Kaitala, A. and Lindfors, V. and Abenius, J.},
  Title = {Polyandry and Its Effect on Female Reproduction in the Green-Veined White Butterfly (Pieris-Napi L)},
  Journal = {Behavioral Ecology and Sociobiology},
  Volume = {33},
  Number = {1},
  Pages = {25-33},
  Note = {Lp947 Times Cited:106 Cited References Count:50},
  Abstract = {In many insects nutrients transferred by the male to the female at mating are later incorporated into both the eggs and soma of the mated females. Accordingly, it has been suggested that female insects can use these male-derived nutrients both for somatic maintenance and to increase both the number and quality of their offspring. Moreover, much discussion is presently devoted to whether the male nuptial gift represents paternal in