Zoology - Theses

Permanent URI for this collection

Search Results

Now showing 1 - 8 of 8
  • Item
    Thumbnail Image
    The function of female and male ornaments in the lovely fairy-wren
    Leitao, Ana V. ( 2019)
    Ornaments like plumage colours or complex song are generally regarded as male traits that are shaped by sexual selection. By contrast, the factors that shape female elaborate traits have often been overlooked, though they are expressed in females across many taxa. Understanding how trade-offs and selective pressures shape female ornamentation is crucial for advancing our understanding of trait evolution. In this thesis, I investigate the form and function of female and male plumage colour and song in the Lovely fairy-wren (Malurus amabilis), a tropical species in which females and males are both highly colourful and vocal. This was investigated over three consecutive years and field seasons in Far North Queensland, Australia. My thesis research employed field observations, behavioural experiments, and genetic analysis, to test the adaptive function(s) and mechanisms for the evolution of female and male ornamental traits. I explicitly contrast females and males so that we can address, in the light of the abundant work done on males, how females may or may not differ from males. To provide context for the ornamental traits that are exhibited by this species, I first provide a comprehensive overview of the ecology and breeding biology of the Lovely fairy-wren, since a detailed description on the species natural history prior to this work was lacking. To understand the function of plumage colouration, I studied whether plumage colour in females and males is a signal and experimentally tested if it functions in a competitive context. Additionally, I assessed whether plumage colour is sexually selected, by examining its signalling content, costs (survival), and its relationship with reproductive and paternity success. Lastly, I investigated the song function, by describing female and male song structure and examining sex-specific variation in song rate across different contexts. I also used experimental data to examine female and male responses to simulated territorial intrusion. Overall this thesis provides insight into the form and function of both female and male plumage colours and song. First, it shows that visual and acoustic ornaments are important signalling components in different contexts, suggesting that female ornaments are not just a correlated genetic by-product of traits in males, and that selection favours female (and male) expression of traits. Second, the information conveyed by plumage colouration seems to be context-dependent in relation to the sex of the bearer: in males, it may follow the classical pattern of sexual selection, functioning in mate choice and male-male competition, while in females, plumage colours do not seem to be influenced by male choice, but function in same-sex competitive contexts. Third, it highlights that song has convergent functions in both sexes, as females and males have similar song structure and used song year-round in identical contexts for within-pair communication and joint territorial defence. The fact that females and males sing and have bright colours year-round in parallel with their territorial and breeding behaviour, suggests that individuals use their traits to maintain (sexual and non-sexual) resources. This work highlights the importance of studying and considering the fundamental differences in females and males, a necessary step for a realistic understanding of ornament expression, and contributes to the ongoing discussion on the evolution of elaborate female signal traits.
  • Item
    Thumbnail Image
    Evolution and biogeography of Australian tropical freshwater fishes
    Shelley, James ( 2016)
    Australia’s freshwater fish fauna is the most depauperate of any continent (256 formally recognised species), although endemism is exceptionally high (74%), largely due to its arid climate and history of isolation from other land masses. The Australian Monsoonal Tropics (AMT) biome in the tropical north is an exception. The AMT encompasses 33% of the Australia landmass, but contains 65% of the Australian fish fauna and, in a global context, the biome and many of its catchments contain moderate to high species richness relative to their size. However, the biodiversity, evolution, and biogeography of the AMT’s fish fauna remain poorly studied relative to the rest of the continent. In this thesis I utilise samples from the most comprehensive region-wide collection of freshwater fish molecular and distributional data in the AMT to help answer three fundamental questions regarding the regions freshwater fish fauna: (1) what is the true biodiversity of the AMT; (2) what are the key evolutionary processes driving and maintaining freshwater fish diversity across the region, in particular the highly endemic fauna of the Kimberley bioregion; and (3) what are the key patterns in diversity and distributions across the landscape and how can they be arranged into a cohesive biogeographic framework? First, I conducted a multigene molecular assessment of species boundaries in the AMTs most speciose freshwater family (Terapontidae) in order to assess the phylogenetic relatedness of terapontids in northwestern Australia (including the Kimberley) to the level of population, and to identify any unique genetic lineages that likely represent undescribed ‘candidate species’. I demonstrated the presence of 13 new candidate species within the Kimberley, more than doubling previous estimates of terapontid diversity in the region. Second, I conducted an assessment of morphological (morphometric and meristic) data from seven of the genetically defined candidate taxa, and the four previously described species within the genus Syncomistes to see if the seven candidates can be discriminated morphologically and to determine which characters best delimit taxa. I found an impressive array of meristic and morphometric character differences between species within Syncomistes and determined that the head, particularly feeding structures such as the jaw and dentition, were the most important morphological features in discriminating between taxa. Third, I looked for congruence between phylogenetic patterns in Kimberley terapontids and both past (low sea-level)/present (high sea-level) geological barriers and pathways as identified by GIS analysis, and tested the general hypothesis that geographic isolation of terapontid lineages during Pliocene and Pleistocene high sea-levels triggered the onset of reproductive isolation between taxa thus driving rapid speciation in the region. I found that most Kimberley terapontid species arose during the Plio-Pleistocene glacial cycles and are at different stages of allopatric divergence and speciation caused by the same vicariant processes. The results support the hypothesis that changing sea levels during late Pliocene and Pleistocene glacial cycles are a key driver of speciation and distributional patterns in the Kimberley. Fourth, I combined phylogenetic, biogeographical and diversification analyses to examine the nature of the Kimberley as a mesic refugium. Specifically, I investigate the tempo and timing of endemic diversification to see if the Kimberley has been a ‘museum’ or a ‘cradle’ of diversification. My combined molecular clock estimates and likelihood-based historical biogeographic reconstructions suggest that terapontids recently transitioned into the Kimberley from the east during the late-Miocene. Outstandingly, ~80% of Kimberley terapontids diversified within the region in the last 3 Ma. Further diversification analyses identified a single significant shift in diversification rates ~1.4 Ma that corresponds with a significant change in global climate midway through the Pleistocene. Given these finding my findings suggest that the Kimberley has been acting as a cradle of Neoendemism. Fifth, I generate a bioregionalisation of the freshwater fish in the AMT using the Simpson’s beta dissimilarity metric, and then assess the relationships of the biogeographic regions to their current environment using generalised dissimilarity modeling (GDM). I also estimate true species richness across catchments using the Chao 2 index in order to identify major sampling gaps. I propose three major freshwater fish bioregions and 14 subregions that differ substantially from the current bioregionalisation scheme. I found that species turnover was most strongly influenced by environmental variables that reflect changes in terrain (catchment relief and confinement) and productivity (NPP and forest cover). Current river orientation and historic connectivity between rivers during low sea-level events also appear to be influential. Three focal points of species richness and two of endemism were identified in the AMT, considerably expanding upon the spatial understanding of these patterns. Finally, a number of key sampling gaps are identified that need to be filled in order to fully refine the proposed regionalisation. Overall the results of this thesis add considerably to biodiversity estimates and the taxonomic knowledge of freshwater fish communities in the AMT. It also helps determine the major drivers of speciation in the Kimberley, the mode of diversification, and provides insight into the regions function as an evolutionarily important mesic refugium. Finally, it provides a modern freshwater bioregionalisation of the AMT and helps to determine the environmental variables driving community change across the landscape. These findings have important ramifications for the conservation of Australia’s tropical freshwater fishes. The Kimberley in particular is highlighted as not only an important evolutionary refugium, but also as a catalyst for narrow range endemic speciation. As a result the regions contains some of the most threatened freshwater fish communities in Australia.
  • Item
    Thumbnail Image
    The evolution of colour patterns in chrysomelines
    Tan, Eunice Jingmei ( 2016)
    The risk of predation is pervasive, and numerous anti-predator strategies have evolved in response to this selection pressure. The same animal can be cryptic in one habitat but conspicuous in another, depending on its background. Chrysomeline leaf beetles have a variety of colour patterns that, to varying degrees, fit or contrast with their environments. Chrysomelines also possess glands that produce defensive chemicals, and these toxic defence mechanisms, together with their diurnal lifestyle, encourage the view that the colour patterns of chrysomelines are aposematic, a defensive mechanism in which a combination of conspicuousness and colour pattern is thought to signal the unprofitability of the prey to visual predators. However, attributing an aposematic role to colour patterns commonly associated with toxicity may not necessarily reflect the evolutionary significance of the colour patterns. Through comparative and experimental studies, I suggest that the diversity of colours and patterns of chrysomeline beetles do not serve a single purpose of aposematic colouration, despite the presence of chemical defences in these taxa. My studies suggest that the ancestor of Australian chrysomelines was likely an inconspicuous, non-iridescent, disruptively coloured, medium-sized beetle. In general, chrysomelines progressively evolved conspicuous and disruptive signals, while iridescence was gained and lost frequently. Chrysomelines that are not disruptively coloured appear to be either cryptic or conspicuously coloured, both of which are visual forms of anti-predator strategies. The variety of anti-predator strategies in chrysomelines may explain their successful radiation into a variety of habitats. Although closely related chrysomeline species utilise similar ecological habitats and hosts, differences in colour and pattern between beetle and host plant mostly do not have a strong phylogenetic signal. Species inhabiting darker environments tend to be larger than beetles inhabiting lighter environments. Host specialisation of leaf beetles may have led to similar levels of difference, across cryptic species, between the colour pattern of the beetle and its host plant and light environment. However, this association is not evident among conspicuous species. This could be a product of the diverse colour pattern differences that species adopt to be conspicuous: as long as they are detectable from the background, conspicuous species have a much larger range of colour patterns that they can adopt, compared with cryptic species. Chrysomeline larvae also show striking patterns and are chemically protected. Experimentally, I show that effective aposematic warning signals in larvae involve a combination of colour and pattern. Müllerian mimicry could have evolved in response to intense predation in the adaptation of similar colour patterns in some species of chrysomeline larvae. Avian predators are usually thought to be driving these systems, but the natural enemies of chrysomeline beetles include other animals, including parasitoids, and future studies would profit from considering the role of these additional natural enemies.
  • Item
    Thumbnail Image
    Geographic variation and speciation in the colour polymorphic tawny dragon lizard
    McLean, Claire Alice ( 2014)
    Polymorphism, the co-occurrence of multiple discrete variants (or morphs) within a single population, represents extreme intraspecific diversity and can provide critical insights into evolutionary processes. Many polymorphic species vary in the number, type, and frequency of morphs present in different populations, which is likely to be shaped by the interaction between spatially and temporally variable selection, genetic drift, and gene flow. This geographic variation can potentailly act as a precursor to speciation as assortative mating and/or genetic incompatabilities can arise between populations that differ in morph composition. Despite this, few studies have focused on geographic variation in polymorphism, its causes and evolutionary consequences. In this thesis, I studied geographic variation in the tawny dragon, Ctenophorus decresii, a small agamid lizard in which males are polymorphic for throat coloration. Using extensive field surveys and multi-locus phylogenetic data, I first investigated how biogeographic and demographic histories have shaped the current distribution of colour variation in C. decresii. I identified three geographically oriented lineages, each characterised by unique male throat coloration. Within South Australia there was a northern and a southern lineage, and the level of divergence between them indicated early quaternary separation. The northern lineage was polymorphic with four discrete throat colour morphs (orange, yellow, orange and yellow combined, and grey), and populations differed in the relative frequencies of morphs, while the southern lineage was monomorphic with blue-throated males. Populations from New South Wales were deeply genetically divergent, and subsequent morphological analyses confirmed that this lineage was a separate species, thus I described the Barrier Range dragon, Ctenophorus mirrityana. There was also notable morphological divergence between the South Australian lineages, and limited genetic introgression at a zone of secondary contact, suggesting potential barriers to gene flow. Throat coloration is likely to be a social signal in C. decresii, as males display their throats during courtship and territorial encounters. Using visual models, I assessed whether divergent coloration between the southern and northern lineage reflected local adaptation for increased detectability to conspecific lizards and/or increased crypsis to predators in different environments. I found that lineage specific throat colours (blue and orange) were more conspicuous to conspecifics against native lichen colours, therefore it appears that background colours have influenced the evolution of social signals in these lineages. Finally, I investigated the interaction between environmentally driven selection and gene flow in shaping geographic variation in morph frequencies in the northern lineage of C. decresii. I found that gene flow between populations was generally low and populations with similar morph frequencies were not necessarily genetically similar. Conversely, there were strong association between morph frequencies and measures of aridity and vegetation cover. This suggests an important role for selection in shaping population morph frequencies in C. decresii, and further research focusing on the northern lineage may help to answer questions about polymorphism maintenance and the link between geographic variation in polymorphism and speciation.
  • Item
    Thumbnail Image
    The regulation and development of the marsupial appendage
    Chew, Keng Yih ( 2013)
    The mammalian appendage is extremely diverse and complex none more diverse than the limb and phallus. These two seemingly unrelated appendages share many similarities particularly in their development and patterning. The parallels between the development of the two appendages are remarkable but there are some key differences between the two structures. The mouse model is predominantly used to investigate these pathways but alternative models which possess specialised traits and morphologies have helped to uncover new pathways involved in the development and evolution of these appendages. In particular, the tammar’s mode of reproduction and it specialised limbs have provided new avenues for investigation of the key mechanisms of development and evolution of the limb and phallus. The cellular, molecular and hormonal changes that that occur in the developing marsupial during limb and phallus formation are described in this thesis. Here, this study focussed on the macropodid marsupial limb, which demonstrates one of the most dramatic examples of heterochrony, which can be defined as the difference in developmental timing between the fore and hind-limb. Heterochrony is clearly evident across all marsupial species to varying degrees. In the tammar, I find that there was high conservation of gene function but clear differences in the timing and mechanism of heterochrony when compared to the only other marsupial species so far examined, the opossum. The heterochrony between the macropodid fore and hind-limb is achieved through a minor shift in the induction of the forelimb marked by TBX5 expression and a later acceleration of downstream growth and patterning elements as compared to the mouse. In contrast, the opossum heterochrony, in which the entire limb patterning program starting from induction of the limb bud was shifted forward in time, began much earlier compared to that of the tammar. In addition, there was variability in the physical presence of the key limb patterning signalling centre, the apical ectodermal ridge (AER). Interestingly the opossum has a reduced AER in the forelimb compared to the mouse. In contrast, the tammar had a more pronounced AER in the forelimb than the hindlimb when compared to the mouse and opossum, although FGF8 expression was present in the ectodermal ridge of both the fore- and hindlimbs. Thus this thesis highlights the variability between marsupials in their limb development and the different mechanisms developed to achieve heterochrony. This study also provides the first assessment of homeobox genes, HOXA13 and HOXD13 in a syndactylous marsupial. These two key digit patterning genes showed high conservation of gene structure. However, there was slightly divergent expression of HOXA13 from that observed in the chicken and mouse. The expression of these HOX limb markers reflected the heterochrony between the tammar fore and hindlimb. The results in this thesis reaffirm the key role in digit patterning that HOX genes play. In addition the results highlights the evolutionary flexibility of retaining in the most common digit that is lost throughout evolution, digit one. Thus the loss of this digit potentially reflects the least evolutionary constraint. In contrast to the limb, the phallus undergoes an androgen-dependant virilisation stage. In most species this occurs in utero in which examination is not easy due to accessibility and also there can be interference from placental hormones. However, in the tammar, the young in accessible in the pouch and readily manipulated free from such disruption. Thus, this study is the first functional examination into the influence of androgens on phallic patterning gene expression in a marsupial. The results in this thesis suggest that SHH has a possible role in the proliferation and development of the marsupial phallus and that SHH could also have a role to prime the phallus for a later acceleration in development after the androgen-dependent phase. However, this study was restricted to the first 50 days post-partum. This raises the question of how androgen interacts with key patterning genes during later stages of development when the majority of phallus growth occurs. Considering the diversity in form between the limb and the phallus, it is essential to investigate alternative models. The macropodid marsupial which, like all marsupials, delivers a highly altricial young, but also has evolved a hopping mode of locomotion, shows both changes in timing of the development and differentiation of the limbs and phallus, making it an ideal model to answer questions about developmental processes. My findings using the tammar wallaby exemplifies the balance between selection and developmental constraints during evolution. The limb and phallus are unique examples in Nature which elegantly display the evolution of form and function. Taken together, this study exemplifies evolutionary flexibility in the development of appendages.
  • Item
    Thumbnail Image
    Genomic imprinting in the post-natal marsupial
    Stringer, Jessica Miriam ( 2012)
    Genomic imprinting regulates several genes that control nutrition and growth of the developing mammalian fetus. In marsupials, the placental attachment is short lived and most growth and development of the young occurs post-natally. However, the role of imprinted genes in post-natal development, and in particular lactation, has received little attention in eutherians and none at all in marsupials. Therefore, this thesis focussed on three orthologous eutherian imprinted genes that are known to function in the mammary gland; insulin (INS), insulin-like growth factor 2 (IGF2) and growth-factor receptor binding protein 10 (GRB10). The expression, imprinting status and molecular regulation of these genes were examined in the mammary gland and in other adult and pouch young tissues of the marsupial, the tammar wallaby, Macropus eugenii. Direct sequencing confirmed that INS was tissue specifically imprinted and paternally epressed in the tammar liver and monoallelically expressed in the mammary gland. INS 5´RACE identified alternate transcription start sites (TSSs) located in the second to last exon of the tyrosine hydroxylase (TH) gene, which produced a novel TH-INS chimaera. Bisulphite sequencing showed that the INS TSS was highly methylated, but not differentially methylated, whereas the TH-INS TSS appeared to be a putative differentially methylated region (DMR) in both the mammary gland and liver. This is the first study to identify INS imprinting outside the yolk sac and suggests that genomic imprinting in the mother could regulate growth and development of the post-natal young. Tammar IGF2 5´RACE identified three promoters which were orthologous to eutherian promoters P1-P3. Expression of tammar IGF2 was similar to that seen in humans, but unlike humans, no DMRs were detected at any of the tammar IGF2 TSS. IGF2 had imprinted expression from all three promoters in the adult mammary gland while in the liver it switched from monoalleleic expression in the pouch young to biallelic in the adult. Interestingly, imprinting was restricted to the trilaminar layer of the yolk sac placenta. A putative placenta-specific DMR was identified in a region orthologous to the eutherian DMR2. The similarities between eutherian and marsupial IGF2 suggest that regulation and expression of this gene originated from three promoters, which have been selectively maintained for the last 160 million years. Tammar GRB10 was highly conserved with eutherian GRB10. However, unlike in eutherians, marsupials appear to have only one GRB10 promoter, othologous to the eutherian major-promoter. Tammar GRB10 was widely expressed in various tissues including the brain, but was not imprinted in any of the tissues examined. These results suggest that GRB10 imprinting evolved in the eutherian lineage after the eutherian-marsupial divergence, possibly subsequent to the acquisition of the brain-specific promoter. This thesis strongly supports the observations that the evolution of imprinting is ongoing and demonstrates it occurs in an organ that has largely been ignored in imprinting research. Clearly, imprinting in the mammary gland gland has evolved to provide a selective advantage for the provision of nutrients which are essential for the survival of the young mammal post-natally. Collectively, these results support the maternal-infant co-adaptation hypothesis and suggest that imprinting in the mammary gland has become an essential reproductive strategy during mammalian evolution.
  • Item
    Thumbnail Image
    The evolution and ecology of the Gadopsis marmoratus complex
    Sanger, Andrew Colin ( 1986)
    The Family Gadopsidae is the only family of fishes in South-eastern Australia which is both endemic to this region, and strictly confined to fresh water. Although several species of Gadopsis have been proposed, prior to the commencement of this study only one species, G. marmoratus Richardson, was formally recognised. This species was known to exhibit considerable geographic variation in morphology, colouration patterns, and size; and it had been suggested that the taxonomy of the genus was in need of review. This study is an attempt to: (i) determine the number of species in the, G. marmoratus complex; (ii) assess the relationships between G. marmoratus and any additional taxa in the complex; (iii) Formulate a biogeographic hypothesis to account for the geographic distributions and phylogenetic relationships of the taxa in the G. marmoratus complex; and, (iv) examine ecological characteristics of a population of Gadopsis from each side of the Great Dividing Range in Victoria. Analysis of meristic variation revealed that two sympatric taxa of Gadopsis were present in north-eastern Victoria. No evidence of hybridization between these taxa was found. The two taxa have different numbers of spines in the dorsal fin, and the specific status of the taxon with only two spines in the fin was recognised by the formal description of G. bispinosus Sanger. By adoption of an evolutionary species concept, and using univariate and multivariate statistical analyses of meristic variation, it was shown that G. marmoratus was comprised of two divergent taxa. It was proposed that these taxa be recognised as separate species, and for the purposes of this study are referred to as northern G. marmoratus and southern G. marmoratus. Multivariate statistical analyses of morphometric variation supported the proposal to recognise northern G. marmoratus and southern G. marmoratus as separate species. Electrophoretic analysis of protein variation also supported the recognition of northern G. marmoratus and southern G. marmoratus, and confirmed that hybridization was not occurring between G. bispinosus and northern G. marmoratus. The level of genetic divergence between northern G. marmoratus and southern G. marmoratus was less than that between both of these taxa and G. bispinosus. This result was interpreted to indicate a close phylogenetic relationship between northern G. marmoratus and southern G. marmoratus, and supported the relationships suggested by the phenetic analysis of meristic and morphometric variation. A biogeographic hypothesis to explain the geographic distributions and phylogenetic relationships of the three taxa was proposed. The ancestry of the Family Gadopsidae has been the subject of some dispute, and the alternative views on this subject were critically reviewed. Support was found for a basal percoid relationship for the family, and this result suggested that Gadopsis may have been of Gondwanan origin. The freshwater life-cycle of all the members of the family was interpreted as further support for a freshwater origin for Gadopsis. Several ecological characteristics of a population of G. bispinosus from the King Parrot Creek and G. marmoratus from the Yarra River system were compared. Gadopsis bispinosus in the King Parrot Creek is a small, lightly-built, short-lived species compared to G. marmoratus from the Yarra River system. The two species were found to have similar reproductive biologies. The discovery of additional species of Gadopsis has created the need for further examination of the evolution and ecology of the G. marmoratus complex, and suggestions for such further study are included at the end of this study.
  • Item
    Thumbnail Image
    Plumage pattern function and evolution: a phylogenetic and comparative approach
    GLUCKMAN, THANH-LAN ( 2011)
    Visual patterns, such as bars and spots, are common in the animal kingdom. In no other group are patterns so exquisite in their arrangement and coloration than in birds. Although bird plumage patterns appear to be visually diverse there are only four types of patterns, which can be broadly categorized into irregular and regular patterning. That these types of irregular and regular patterning are recursive is intriguing and speaks of an underlying shared mechanism on which selection can act. The prevailing assumption is that patterns predominantly function in camouflage, however evidence suggests that they also function in communication in a small number of birds. In particular it has been suggested that barred plumage patterns could be a signal of individual quality. In visual ecology, communication and camouflage seem to be in conflict with one another – visual signals are often conspicuous whereas camouflage has evolved to provide concealment. These ideas of pattern function need not be incongruous if patterns evolved a) for camouflage first and were subsequently co-opted by sexual selection for communication, and/or b) some patterns, specifically barred plumage, evolved for both camouflage and communication to overcome this functional compromise. To test these alternative ideas of pattern evolution I test whether a) patterns were co-opted for signaling in the model group waterfowl and gamebirds, and b) if the evolution of sexual dimorphism in barred plumage indicate camouflage and ⁄ or signaling functions across the class Aves. Additionally, I investigated whether development poses a constraint on pattern evolution in waterfowl and gamebirds. Tracing the most probable evolutionary pathway of plumage pattern evolution revealed that the ancestral state of plumage was uniform coloration. From uniform coloration, patterns initially evolved to be predominantly monomorphic, and subsequently evolved to be sexually dimorphic. In sexually dimorphic patterns, barred plumage frequently evolved in females and males, suggesting a role for both camouflage and communication. However, dimorphic spotted plumage only evolved in males suggesting it predominantly evolved for communication. Overall, it is likely Plumage pattern function and evolution: a phylogenetic and comparative approach ii that patterns originally evolved for camouflage and were subsequently co-opted for signaling. Focusing on the evolution of barred patterns by comparing their prevalence between the sexes I found a higher frequency of female- rather than male-biased sexual dimorphism, indicating that camouflage is its most common function. But I also found that, compared with other pigmentation patterns, barred plumage is more frequently biased towards males and its expression more frequently restricted to adulthood, suggesting that barred plumage often evolves or is maintained as a sexual communication signal. This illustrates how visual traits can accommodate the apparently incompatible functions of camouflage and communication. Lastly, I studied the recurrence of irregular and regular plumage patterns to explore why there are different kinds of patterns that are broadly recursive. By modeling pattern evolution I derived directionality and show that where species exhibit a single pattern, selection need not be constrained by development. However, instances of irregular and regular patterns in the same species are a result of selection on existing patterns. Together this demonstrates that the evolution of patterns is not difficult and that states of multiple pattern types are a result of selection.