Harvesting Training Images for Fine-Grained Object Categories Using Visual Descriptions
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The Genetics and Evolution of Iridescent Structural Colour in Heliconius Butterflies
The genetics and evolution of iridescent structural colour in Heliconius butterflies Melanie N. Brien A thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy The University of Sheffield Faculty of Science Department of Animal & Plant Sciences Submission Date August 2019 1 2 Abstract The study of colouration has been essential in developing key concepts in evolutionary biology. The Heliconius butterflies are well-studied for their diverse aposematic and mimetic colour patterns, and these pigment colour patterns are largely controlled by a small number of homologous genes. Some Heliconius species also produce bright, highly reflective structural colours, but unlike pigment colour, little is known about the genetic basis of structural colouration in any species. In this thesis, I aim to explore the genetic basis of iridescent structural colour in two mimetic species, and investigate its adaptive function. Using experimental crosses between iridescent and non-iridescent subspecies of Heliconius erato and Heliconius melpomene, I show that iridescent colour is a quantitative trait by measuring colour variation in offspring. I then use a Quantitative Trait Locus (QTL) mapping approach to identify loci controlling the trait in the co-mimics, finding that the genetic basis is not the same in the two species. In H. erato, the colour is strongly sex-linked, while in H. melpomene, we find a large effect locus on chromosome 3, plus a number of putative small effect loci in each species. Therefore, iridescence in Heliconius is not an example of repeated gene reuse. I then show that both iridescent colour and pigment colour are sexually dimorphic in H. -
The Speciation History of Heliconius: Inferences from Multilocus DNA Sequence Data
The speciation history of Heliconius: inferences from multilocus DNA sequence data by Margarita Sofia Beltrán A thesis submitted for the degree of Doctor of Philosophy of the University of London September 2004 Department of Biology University College London 1 Abstract Heliconius butterflies, which contain many intermediate stages between local varieties, geographic races, and sympatric species, provide an excellent biological model to study evolution at the species boundary. Heliconius butterflies are warningly coloured and mimetic, and it has been shown that these traits can act as a form of reproductive isolation. I present a species-level phylogeny for this group based on 3834bp of mtDNA (COI, COII, 16S) and nuclear loci (Ef1α, dpp, ap, wg). Using these data I test the geographic mode of speciation in Heliconius and whether mimicry could drive speciation. I found little evidence for allopatric speciation. There are frequent shifts in colour pattern within and between sister species which have a positive and significant correlation with species diversity; this suggests that speciation is facilitated by the evolution of novel mimetic patterns. My data is also consistent with the idea that two major innovations in Heliconius, adult pollen feeding and pupal-mating, each evolved only once. By comparing gene genealogies from mtDNA and introns from nuclear Tpi and Mpi genes, I investigate recent speciation in two sister species pairs, H. erato/H. himera and H. melpomene/H. cydno. There is highly significant discordance between genealogies of the three loci, which suggests recent speciation with ongoing gene flow. Finally, I explore the phylogenetic relationships between races of H. melpomene using an AFLP band tightly linked to the Yb colour pattern locus (which determines the yellow bar in the hindwing). -
Genetic Differentiation Without Mimicry Shift in a Pair of Hybridizing Heliconius Species (Lepidoptera: Nymphalidae)
bs_bs_banner Biological Journal of the Linnean Society, 2013, 109, 830–847. With 5 figures Genetic differentiation without mimicry shift in a pair of hybridizing Heliconius species (Lepidoptera: Nymphalidae) CLAIRE MÉROT1, JESÚS MAVÁREZ2,3, ALLOWEN EVIN4,5, KANCHON K. DASMAHAPATRA6,7, JAMES MALLET7,8, GERARDO LAMAS9 and MATHIEU JORON1* 1UMR CNRS 7205, Muséum National d’Histoire Naturelle, 45 rue Buffon, 75005 Paris, France 2LECA, BP 53, Université Joseph Fourier, 2233 Rue de la Piscine, 38041 Grenoble Cedex, France 3Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Panama 4Department of Archaeology, University of Aberdeen, St. Mary’s Building, Elphinstone Road, Aberdeen AB24 3UF, UK 5UMR CNRS 7209, Muséum National d’Histoire Naturelle, 55 rue Buffon, 75005 Paris, France 6Department of Biology, University of York, Wentworth Way, Heslington, York YO10 5DD, UK 7Department of Genetics, Evolution and Environment, University College London, Gower Street, London WC1E 6BT, UK 8Department of Organismic and Evolutionary Biology, Harvard University, 16 Divinity Avenue, Cambridge, MA 02138, USA 9Museo de Historia Natural, Universidad Nacional Mayor San Marcos, Av. Arenales, 1256, Apartado 14-0434, Lima-14, Peru Received 22 January 2013; revised 25 February 2013; accepted for publication 25 February 2013 Butterflies in the genus Heliconius have undergone rapid adaptive radiation for warning patterns and mimicry, and are excellent models to study the mechanisms underlying diversification. In Heliconius, mimicry rings typically involve distantly related species, whereas closely related species often join different mimicry rings. Genetic and behavioural studies have shown how reproductive isolation in many pairs of Heliconius taxa is largely mediated by natural and sexual selection on wing colour patterns. -
Ecological Divergence and Speciation in Heliconius Cydno and H
Ecological divergence and speciation in Heliconius cydno and H. melpomene by Russell Edward Naisbit A thesis submitted for the degree of Doctor of Philosophy of the University of London September 2001 Department of Biology University College London To my family, for their support and encouragement throughout this crazy endeavour 2 “It is hardly an exaggeration to say, that whilst reading and reflecting on the various facts given in this Memoir, we feel to be as near witnesses, as we can ever hope to be, of the creation of a new species on this earth.” Charles Darwin, Natural History Review: Quarterly Journal of Biological Science, 1863. From a review of “Contributions to an Insect Fauna of the Amazon Valley,” in which Henry Walter Bates gave an adaptive explanation for mimicry in Amazonian butterflies and argued that variation in mimicry might cause speciation 3 Abstract We are in the midst of a renaissance in speciation research. There is a return to Darwin’s belief in the role of natural selection in driving speciation, after a lengthy focus on geographic isolation and hybrid sterility. Here I describe the ecological, behavioural, and genetic bases of speciation in Heliconius cydno and Heliconius melpomene (Lepidoptera: Nymphalidae). The two species are sympatric in tropical rainforest across most of Central America and the foothills of the Andes. Ecological differentiation allows coexistence of these sister species despite rare hybridisation. Divergence in microhabitat and larval host plant use has reduced both the potential for gene flow and for competition. In Panama H. cydno uses most Passiflora species in closed canopy forest, whilst H. -
Insect Seminal Fluid Proteins: Identification and Function
EN56CH02-Wolfner ARI 14 October 2010 9:49 Insect Seminal Fluid Proteins: Identification and Function Frank W. Avila, Laura K. Sirot, Brooke A. LaFlamme, C. Dustin Rubinstein, and Mariana F. Wolfner Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York 14853; email: [email protected], [email protected], [email protected], [email protected], [email protected] Annu. Rev. Entomol. 2011. 56:21–40 Key Words First published online as a Review in Advance on egg production, mating receptivity, sperm storage, mating plug, sperm September 24, 2010 by University of California - San Diego on 08/13/13. For personal use only. competition, feeding, reproduction, Acp Annu. Rev. Entomol. 2011.56:21-40. Downloaded from www.annualreviews.org The Annual Review of Entomology is online at ento.annualreviews.org Abstract This article’s doi: Seminal fluid proteins (SFPs) produced in reproductive tract tissues 10.1146/annurev-ento-120709-144823 of male insects and transferred to females during mating induce nu- Copyright c 2011 by Annual Reviews. merous physiological and behavioral postmating changes in females. All rights reserved These changes include decreasing receptivity to remating; affecting 0066-4170/11/0107-0021$20.00 sperm storage parameters; increasing egg production; and modulating sperm competition, feeding behaviors, and mating plug formation. In addition, SFPs also have antimicrobial functions and induce expression of antimicrobial peptides in at least some insects. Here, we review re- cent identification of insect SFPs and discuss the multiple roles these proteins play in the postmating processes of female insects. 21 EN56CH02-Wolfner ARI 14 October 2010 9:49 INTRODUCTION SFPs provide intriguing targets for the control of disease vectors and agricultural pests. -
Genomic Hotspots of Adaptation in Butterfly Wing
Author's personal copy Available online at www.sciencedirect.com Genomic hotspots of adaptation in butterfly wing pattern evolution Riccardo Papa, Arnaud Martin and Robert D Reed What is the genetic architecture of morphological evolution? Is underlies adaptation in multiple lineages. There are there uniform potential for novelty across a genome or, on the now numerous cases known of specific genes underlying contrary, can a small number of large-effect genes explain the parallel and convergent evolution [3]. For example, regu- phenotypic variation observed within and between species? latory elements of yellow underlie multiple cases of con- Here we highlight recent work on butterfly wing pattern vergent evolution of both abdomen and wing genetics showing that a small set of loci can be repeatedly pigmentation in Drosophila species [4,5], independent involved in the evolution of complex traits. These loci behave as events of albinism in cavefish were linked to mutations genomic hotspots for diversification because they underlie in Oca2, a determinant of pigmentation in human popu- adaptive variation within and between species with both lations [6,7], and derived pigmentation in several stickle- convergent and highly divergent wing patterns. These findings back populations is linked to a regulatory allele of Kitlg,a suggest that certain loci may be more likely than others to gene associated with skin color in humans [8]. Repeated facilitate rapid evolutionary change. fixation of an allele of the Eda gene is responsible for the reduction of armor plates in sticklebacks [9–11], while its Address receptor Edar matches a quantitative trait locus (QTL) Department of Ecology and Evolutionary Biology, University of for hair thickness in humans [12]. -
Genetic Evidence for a Sibling Species of Heliconius Charithonia (Lepidoptera; Nymphalidae)
Biological Journal of the Linnean Society (1998), 64: 57±67. With 1 ®gure Article ID: bj970211 Genetic evidence for a sibling species of Heliconius charithonia (Lepidoptera; Nymphalidae) CHRIS D. JIGGINS* The Galton Laboratory, Department of Biology, University College London, 4 Stephenson Way, London NW1 2HE NEIL DAVIES University of Hawaii at Manoa, Center for Conservation Research and Training, Honolulu, HI 96822, U.S.A. Received 29 May 1997; accepted for publication 22 December 1997 Heliconius charithonia is a widespread species which, unlike many Heliconius, is non-mimetic and shows little racial diVerentiation. 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 diVerentiation was ten times 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 ®ve diagnostic loci and was most likely an F1 hybrid between H. charithonia and H. peruvianus. These data imply that H. charithonia and H. peruvianus are distinct species which hybridize occasionally. This species pair show many similarities with H. -
Dissecting Comimetic Radiations in Heliconius Reveals Divergent Histories of Convergent Butterflies
Dissecting comimetic radiations in Heliconius reveals divergent histories of convergent butterflies Swee-Peck Queka, Brian A. Countermanb, Priscila Albuquerque de Mourac, Marcio Z. Cardosoc, Charles R. Marshalld, W. Owen McMillanb, and Marcus R. Kronforsta,1 aFaculty of Arts and Sciences Center for Systems Biology, Harvard University, Cambridge, MA 02138; bDepartment of Genetics, North Carolina State University, Raleigh, NC 27695; cDepartment of Botany, Ecology and Zoology, Universidade Federal do Rio Grande do Norte, Natal RN 59072-970, Brazil; and dDepartments of Organismic and Evolutionary Biology and Earth and Planetary Sciences, Harvard University, Cambridge, MA 02138 Edited by May R. Berenbaum, University of Illinois at Urbana–Champaign, Urbana, IL, and approved March 16, 2010 (received for review October 6, 2009) Mimicry among Heliconius butterflies provides a classic example of tandem, from location to location (4, 9, 10). The racial variation coevolution but unresolved relationships among mimetic subspe- in H. erato and H. melpomene involves both major phenotypic cies have prevented examination of codiversification between shifts, such as the difference between rayed and postman pat- species. We present amplified fragment length polymorphism terns, as well as relatively minor variations, such as the subtle and mtDNA datasets for the major comimetic races of Heliconius differences among rayed populations or among postman pop- erato and H. melpomene. The AFLP data reveal unprecedented ulations. Two primary hypotheses have been proposed -
Mallet 1986C
RNATIONAL ❑ GEOGRAPHIC A SCIENTIFIC JOURNAL SPRING 1986 GREGARIOUS BUTTERFLIES ❑ YORUBA RITUAL I=1 DINOSAUR EVOLUTION ❑ TROPICAL ISLAND WATER BALANCE 0. , James Mallet Gregarious Roosting and Home Range in Heliconius Butterflies Individual Heliconius butterfliesfly within a restricted area by day and roost gregariously at night. In premosting gatherings, individuals `fan" already roosted conspecifics. Fanning does not have an aggressive function, as has been previously suggested; it is a courtshiplike behav- ior, although it never leads to mating. The behavior occurs with high frequencyjust before roosting and torpidity, rather Mart during the day when mating takes place. Individuals also fan detul ir rdividuals of Their own and other species, and even crude cardboard models. Fan n ing is probably a means of testingfor species-specific stimuli that enable indi- viduals to aggregate. Roosts of H. erato do not consist of discrete farn P groups; both ten- eral and older butterflies disperse among roosting groups. The home ranges of individual erato are independent of those of their roostmates, and frequently overlap with home ranges of individuals from other roosts. The loose population structure found in erato contrasts with that required for kin selection models of the evolution of warning color and unpalatability. There is little predation of Heliconius near the roost, but foraging birds often disturb erato during the early morning, before the roosts disperse. The selective value ofgregarious roosting is discussed. Heliconius butterflies inhabit tropical forests and woods from the south- ern United States to Argentina. Their long wings, which allow a lazy, hovering flight, are usually black with gaudy patches of red, orange, yellow, white, or iridescent blue. -
Seminal Fluid Protein Genes of the Brown Planthopper, Nilaparvata Lugens Bing Yu, Dan-Ting Li, Jia-Bao Lu, Wen-Xin Zhang and Chuan-Xi Zhang*
Yu et al. BMC Genomics (2016) 17:654 DOI 10.1186/s12864-016-3013-7 RESEARCH ARTICLE Open Access Seminal fluid protein genes of the brown planthopper, Nilaparvata lugens Bing Yu, Dan-Ting Li, Jia-Bao Lu, Wen-Xin Zhang and Chuan-Xi Zhang* Abstract Background: Seminal fluid proteins (SFPs) are produced mainly in the accessory gland of male insects and transferred to females during mating, in which they induce numerous physiological and post-mating behavioral changes. The brown plant hopper (BPH), Nilaparvata lugens, is an economically important hemipterous pest of rice. The behavior and physiology of the female of this species is significantly altered by mating. SFPs in hemipteran species are still unclear. Results: We applied high-throughput mass spectrometry proteomic analyses to characterize the SFP composition in N. lugens. We identified 94 putative secreted SFPs, and the expression levels of these proteins was determined from the male accessory gland digital gene expression database. The 94 predicted SFPs showed high expression in the male accessory gland. Comparing N. lugens and other insect SFPs, the apparent expansion of N. lugens seminal fluid trypsins and carboxylesterases was observed. The number of N. lugens seminal fluid trypsins (20) was at least twice that in other insects. We detected 6 seminal fluid carboxylesterases in N. lugens seminal fluid, while seminal fluid carboxylesterases were rarely detected in other insects. Otherwise, new insect SFPs, including mesencephalic astrocyte–derived neurotrophic factor, selenoprotein, EGF (epidermal growth factor) domain–containing proteins and a neuropeptide ion transport-like peptide were identified. Conclusion: This work represents the first characterization of putative SFPs in a hemipeteran species. -
Rapid Speciation, Hybridization and Adaptive Radiation in the Heliconius Melpomene Group James Mallet
//FS2/CUP/3-PAGINATION/SPDY/2-PROOFS/3B2/9780521883184C10.3D 177 [177–194] 19.9.2008 3:02PM CHAPTER TEN Rapid speciation, hybridization and adaptive radiation in the Heliconius melpomene group james mallet In 1998 it seemed clear that a pair of ‘sister species’ of tropical butterflies, Heliconius melpomene and Heliconius cydno persisted in sympatry in spite of occasional although regular hybridization. They speciated and today can coexist as a result of ecological divergence. An important mechanism in their speciation was the switch in colour pattern between different Mu¨ llerian mimicry rings, together with microhabitat and host-plant shifts, and assortative mating pro- duced as a side effect of the colour pattern differences. An international con- sortium of Heliconius geneticists has recently been investigating members of the cydno superspecies, which are in a sense the ‘sisters’ of one of the original ‘sister species’, cydno. Several of these locally endemic forms are now recognized as separate species in the eastern slopes of the Andes. These forms are probably most closely related to cydno, but in several cases bear virtually identical colour patterns to the local race of melpomene, very likely resulting from gene transfer from that species; they therefore can and sometimes do join the local mimicry ring with melpomene and its more distantly related co-mimic Heliconius erato. I detail how recent genetic studies, together with ecological and behavioural observations, suggest that the shared colour patterns are indeed due to hybrid- ization and transfer of mimicry adaptations between Heliconius species. These findings may have general applicability: rapidly diversifying lineages of both plants and animals may frequently share and exchange adaptive genetic variation. -
Evolutionary and Ecological Processes Influencing Chemical Defense Variation in an Aposematic
1 Title: Evolutionary and ecological processes influencing chemical defense variation in an aposematic 2 and mimetic Heliconius butterfly 3 4 SUPPLEMENTARY FILE 4 5 SUPPLEMENTARY METHODS 6 7 Further information on the study species Heliconius erato 8 Heliconius erato (Lepidoptera: Nymphalidae) is a widespread neotropical butterfly typically occurring at the 9 edges of primary and secondary tropical rainforests. A striking diversity of color-pattern races with region- 10 specific wing patterns have evolved as mimics sharing aposematic coloration with other species, typically H. 11 melpomene, in order to deter common predators (Supple et al., 2013). Heliconius erato has a restricted home 12 range and a dispersal distance of around 1-2 km per generation (Turner, 1971). The larvae can feed on 13 several Passiflora species, depending on the local quantity and quality of available hosts (Kerpel & Moreira, 14 2005). In the Panamanian study area, the main natural hosts of H. erato are P. biflora, P. coriaceae and to a 15 lesser extent P. auriculata (Merrill et al., 2013), while the preferred host plants in Ecuador are not well 16 known (but possibly include P. punctate; (Hay-Roe & Nation, 2007). Cyanogenic toxicity of H. erato is 17 moderate compared to other Heliconius species and arise from both biosynthesized and sequestered toxins, 18 depending on the host on which it feeds (Arias et al., 2016; de Castro et al., 2019). 19 20 Study sites and field collection of butterflies and host plants 21 Heliconius erato demophoon butterflies and its typical host plant P. biflora were collected from forest and 22 forest-edge populations in Panama, and H.