These throids feed selectively on the pipe wine family, Aristolochiaceae, and sequester toxic aristolochic acids (AAs) (Fig.11.3) in the body tissues (Euw et al.1968; Nishida and Fukami1989; Wu et al.2000). Further studies are needed to substantiate possible unpalatability of E. 11.5 The red-bodied butterflies and moths. a) Pachliopta aristolochiae female, (b) Atrophaneura alcinous female, (c) Epicopeia hainesii female, (d) Histia flabellicornis female secreting defensive foam containing linamarin (orange arrow).
Discussion
Nishida R, Schulz S, Kim CS et al (1996) Lalaki a sekso a pheromone ti higante a kulibangbang a danaine, Idea leuconoe. Ti modelo para iti dinamika ti populasion ti mimetiko a kulibangbang a Papilio polytes idiay Is-isla Sakishima, Hapon (II).
Introduction
The model was created on the basis of field data on the islands as well as experimental data in the laboratory. In particular, they discussed the temporal change of relative abundance (RA) since 1975 on Miyako-jima Island and the variation of RA on the Sakishima Islands.
Field Records of Papilio polytes Observed in Sakishima Islands
Observation of Temporal Change in the Population of the Mimetic Female of P. polytes in Miyako-jima
The Sakishima Islands are located at the southeastern tip of Japan and are part of the Ryukyu Islands, which include both the Miyako Islands and the Yaeyama Islands. The Miyako Islands include Miyako-jima Is., Tarama-jima Is., etc., and the Yaeyama Islands include Ishigaki-jima Is., Hateruma-jima Is., Iriomote-jima Is., Taketomi-jima Is. ., Kohama-jima Is., etc.
Variation in the Relative Abundance (RA) in Sakishima Islands
Extended Mathematical Model for Population Dynamics of P. polytes
Fundamental Facts on the Mimicry of P. polytes
From the point of view of producing offspring, the non-mimetic f.cyrushes have an advantage over the mimicking f.polytes. In any case, the result shows that the lifespan of f.cyrus is slightly longer than that of f.polytes.
Mathematical Model of Three ODEs for Population Dynamics of P. polytes with Intraspecific Competition
For example, the number 16.50 in the left column of condition “A” is the average lifespan (days) of 10 (f. cyrus) individuals. According to our statistical analysis of the data, there is no statistically significant difference in the physiological lifespan between two forms f.
Mathematical Analysis of the System Equations and Computer Simulations
Mathematical Analysis
12.1) and (12.2) represents the effect of mimicry, i.e. the effect of negative density which means that the increase in the density of f. In Section 12.3.1.1, we noted the result in beak marks from predators indicating that non-mimetic females were selectively attacked, while males, mimetic females and model butterflies were attacked less. Inequality (12.6) provides analytical evidence for the field data on the positive dependence of RA on AI in the Sakishima Islands in Fig. 12.5b (Uesugi 1992) noted in Section 12.2.2.
Parameter values used in the numerical simulation are all the same as in Fig. 12.5a.
Summary and Discussions
The positive dependence of RA on AI originates from the result that changes in both relative abundance (RA) and superiority index (AI) with respect to K1 carrying capacity are positive. The results in Section 12.4 showed that it would be possible to reproduce the experimental data on the population dynamics of the mimetic butterfly P. Sekimura T, Fujihashi Y, Takeuchi Y (2014) Model for the population dynamics of the mimetic butterfly Papilio polytes in the Sakishima Islands, Japan.
This developmental flexibility of color pattern can result in extremely different seasonal phenotypes in a single species.
Introduction
Elements of butterfly color patterns are developmentally semi-autonomous, allowing detailed developmental and evolutionary changes in the overall color pattern. We show that forewing shape and eyespot size both vary seasonally and that the methods by which phenotypic elements change in dry season forms are different in different clades and therefore have independent and diverse evolutionary origins can have. The elements of butterfly color patterns are developmentally semi-autonomous, allowing detailed developmental and evolutionary changes in the overall color pattern (Nijhout 1991).
We show that forewing shape and eyespot size both vary seasonally in Junoniini and that the mechanisms by which phenotypic elements change in the dry.
Methods
13.1 (a) Phylogeny of Junoniini species (Lepidoptera: Nymphalidae: Nymphalinae) used in this study grouped by reference class. Measurements were normalized by size using (e) the perimeter of a triangle connecting the edges Rs and Cu2 to the root of the ventilation system [triangle RCR]. Tree topology from Kodandaramaiah and Wahlberg2007) Fig. Note the increasing sharpness of the angle formed by the fringes immediately flanking the M1 end as the peak increases in angularity from low to high.
To contrast pattern element data between seasonal forms, character state reconstructions of maturity were mapped onto an existing tree topology based on a molecular phylogeny for the tribe Junoniini (Kodandaramaiah and Wahlberg 2007) and mapped to make comparisons between seasonal forms.
Results
- Variation by Pattern Element
- Variation by Wing Cell
- Seasonal Eyespot Variation by Clade
- Seasonal Forewing Apex Shape Change by Clade
- Shape Type and Shape Change
- Discussion
Finally, the Rs, M1 and Cu1 eyespots are large in the wet season form and highly reduced in the dry season form (Fig.13.3). Furthermore, species whose forewing apex varied seasonally did so according to a pattern of increasing angularity in the dry season shape compared to the wet season shape (Fig.13.7 and 13.8 – bottom). Note the association between exhibiting seasonal shape change and having a higher forewing apex angle in the dry season shape (Tree Topology of Kodandaramaiah and Wahlberg 2007).
A very angular wing shape in the dry season form appears to have evolved very early in the tribe, but was independently lost in both Hypolimnas and upper Junonia (Fig. 13.4 right).
Introduction
The objectives of this study were to (1) document any changes that occur in the appearance of reproductive structures in male swallowtail butterflies (Battus philenor) as a result of mating, as well as the persistence of such changes after mating, in order to develop criteria for identifying males that have recently mated, and (2) examine the reproductive pathways of field-caughtB.
Materials and Methods .1 Source of Animals Used
Examination of Reproductive Tracts of Virgin and Mated Males
Estimation of Recent Mating Success of Field- Caught Male
Spectral Analyses of Iridescent Wing Areas
Results
- Virgin Male Reproductive Tract
- Reproductive Tract of Males Immediately After Mating
- Changes in the Male ’ s Reproductive Tract with Time Since Mating
- Mating Success of Field-Caught Males
For the eight possible groups, and to maximize contrasts, we labeled as having strong evidence of recent mating any group that met two or more of the criteria (groups E to H). We also confidently labeled as not recently mated males that did not meet any of the criteria (Group A). The characters included in the analysis were the intensity and shade of the iridescent area of male dorsal hindwings and age class.
In subsequent analysis, any group that met two or more criteria (groups E to H) and males that did not meet any of the criteria (group A) were used as recently mated males and recently mated males, respectively.
Discussion
Assessing the Mating History of Male Butterflies in the Field
We note that when using this method, there may be some characteristics that are not suitable for examining the relationship with mating history. For example, it is in principle impossible to investigate the relationship between recent mating success and spermatophore production capacity, which is closely linked to the reproductive success of both male and female butterflies.
Phenotypic Correlates of Mating Success in Male B. philenor in the Field
Harari AR, Handler AM, Landolt PJ (1999) Size-assortative mating, male choice and female choice in the curculionid beetle Diaprepes abbreviatus. Keyser AJ, Hill GE (1999) Condition-dependent variation in the blue-ultraviolet coloration of a structurally based plumage ornament. Rutowski RL, Newton M, Schaefer J (1983) Interspecific variation in the size of the nutrient investment made by male butterflies during copulation.
The images or other third party material in this chapter are included in the chapter's Creative Commons license, unless indicated otherwise in a credit line to the material.
Introduction
Among them, the color pattern of the body is particularly interesting, as it is important in visual recognition. The younger larva (first–fourth instar) mimics bird droppings with a black/white body color (marked mimetic pattern, Fig. 15.1a). A similar change in body color pattern is observed in other Papilio species (Prudic et al. 2007) and is considered a successful survival strategy for this genus (Tullberg et al. 2005).
Recent studies have reported that two critical insect hormones, ecdysone (Fig.15.2a, b) and juvenile hormone (JH), directly regulate pigmentation and color pattern change in larval P.
Pigmentation of Larval Cuticle in P. xuthus
Futahashi and Fujiwara (2005) showed that the spatial expression of melanin synthesis genes (TH,DDC, andtan) corresponds perfectly to the presumed black pigment (Futahashi et al. 2010; Futahashi and Fujiwara2005) and that the expression of ebony is restricted to the red area of the eye (Futahashi and Fujiwara2005). Their findings indicate that cuticle color patterns are formed from spatially specific localization of melanin synthesis genes rather than the differential uptake of melanin precursors into individual epidermal cells. When Futahashi and Fujiwara (2005) examined the timing of expression of PxTH, PxDDC, Pxebony and Pxtan, they noted that the expression of these melanin synthesis genes precisely coincides with the onset of melanization.
Therefore, cuticular pigmentation is predictably strictly controlled by ecdysteroid, the molting hormone (Futahashi and Fujiwara2005,2007; Futahashi et al.2010).
Hormonal Regulation of Larval Pigmentation
Ecdysone-Induced Cuticular Pigmentation
In the ecdysone signaling pathway, 20E acts as a hormonal signal and regulates the expression of downstream transcription factors (Yamanaka et al.2013; Yao et al.1992). In an RNAi experiment, all four caused altered pigmentation when knocked down (Kalay et al.2016). 2012) used a dataset of EST data to identify E75AandE75B, which are transcription factors involved in ecdysone signaling, as candidates involved in specific marking patterning (Futahashi et al. 2012).
Because its function converts inactivated 3-dehydroecdysone to ecdysone, localized marker-specific ecdysone synthesis may be critical for complex cuticular pigmentation and pattern formation (Futahashi et al.2012).
Juvenile Hormone Directly Regulates Larval Color Pattern Switch
However, since only some regulatory genes have been identified, the detailed regulatory mechanisms remain to be discovered. Furthermore, the epidermis was observed to be sensitive to JHA only during the first 20 hours of the fourth stage. Therefore, this particular time window was called the "JH-sensitive period." In untreated species, the JH titer in the hemolymph was measured and found to continuously decrease during the early days of the fourth instar.
Taken together, this evidence suggests that the decline in JH titer within a limited developmental stage regulates the change in body color pattern in P.
Species-Specific Color Patterns in the Papilio Genus
A Combination of Yellow and Blue Makes the Larval Body Green
Due to our fragmentary knowledge of JH pathways, the molecular mechanisms underlying how JH alters color patterning and controls pigment synthesis are still under investigation (Jindra et al.2013). Subsequent research led to a model that postulates that pigments are tightly bound to specific proteins and that the pigment-conjugated protein complex accounts for visible coloration (Kawooya et al. 1985). Blue pigment-binding protein (or bilin-binding protein, BBP) has been isolated and identified in various lepidopterans (Riley et al. 1984; Huber et al. 1987; Saito and Shimoda1997; Kayser et al. 2009).
In addition, two putative carotenoid-binding proteins (PCBP1, PCBP2) and other members of BBP family were later identified, which were specifically expressed in the green epidermal regions during the final larval ecdysis (Futahashi et al.2012).
Species-Specific Color Pattern Among Papilio Species
Using next-generation sequencing (NGS) technology, whole genomes of several lepidopteran species have recently been released (Suetsugu et al.2013; Li et al. 2015; Nishikawa et al.2015; Kanost et al.2016). In 2015, whole genome sequences of those three species were released and made freely accessible (Li et al. 2015;. Expression of TH, DDC and yellow corresponded to the black regions in the eyespot, the V-shaped markings of P.
Regardless of the universal expression of BBP1 and YRG in the green areas among all three species, BBP1 was specifically expressed in the blue spots iP.
Trans-regulation of YRG in the Genus Papilio
Conclusion and Future Prospects
Futahashi R (2006) Molecular mechanisms of mimicry in larval body marking of the swallowtail butterfly, Papilio xuthus. Hiruma K, Riddiford LM (1990) Regulation of dopa decarboxylase gene expression in the larval epidermis of the tobacco hornworm by 20-hydroxyecdysone and juvenile hormone. Jindra M, Riddiford LM (1996) Expression of ecdysteroid-regulated transcripts in the silk gland of the wax moth, Galleria mellonella.
Riddiford LSLM (1975) Biology of the black larval mutant of the tobacco hornworm, Manduca sexta.
Introduction
