Colour and colour change in

Kate Umbers Dr Kate Umbers Draco maculatus Macquarie University & Australian National University (soon to be University of Wollongong) Why are animals colourful?

Kate Umbers Draco maculatus Functions of colouration in animals crypsis masquerade mimcry sensory exploitation aposematism sexual thermoregulation protection from damaging radiation burrowing efficiency crypsis

Nikolai Tatarnic Happarana pallida signalling - masquerade

tumblr.com Uroplatus phantasticus signalling - mimcry

amphibiancare.com Dendrobates imitator signalling - sensory exploitation

Kate Umbers Hymenopus ornatus signalling - aposematism

asknature.org Notaden bennetti signalling - sexual

Kate Umbers

Physignathus lesueurii thermoregulation

allposters.co.uk Coccinella novemnotata protection from damaging radiation

healthpost.com.au Euphausia superba burrowing efficiency

Kate Umbers Polyzosteria viridissima costs of being colourful

Kate Umbers colour production - pigments

Robert Strickland Carpodacus mexicanus colour production - structures

Cor Zonneveld Callophrys rubi conspicuousness

nationalgeographic.com Morphinae colour change reversible and irreversible

Kate Umbers Octopus tetricus Best of both worlds?

Kate Umbers Octopus tetricus Best of both worlds? Why not change colour?

Kate Umbers Octopus tetricus colour change surprisingly common

Mollusca, Chordata, Arthropoda, +?

Kate Umbers Octopus tetricus that change colour Araneus quadratus Ocypode ceratophthalma Corethra plumicornis Argia oculata Cyrtophora cicatrosa Uca capricornus Euphausia superba Argia sedula Gea heptagon Uca maracoani Nezara viridula Argia tibialis Floronia bucculenta Uca minax Plautia stali Argia vivida Argyria venusta Uca mordax Anilocra physodes aspersum Micrommata virescens Uca pugilator Idotea emarginata Enallagma cardenium evanida Uca pugnax Idotea japonica Enallagma cyathigerum vatia Uca pugnax Idotea montereyensis Enallagma durum Misumenoides formosipes Uca rapax Idotea neglecta Enallagma geminatum labefactus Uca speciosa Itodea metallica Ischnura heterosticta Uca thayeri Ligia baudinian Xanthagrion erythroneurum Thomisus spectabilis Leander serratus Ligia exotica Diphlebia lestoides Tmesisternus isabellae Macrobrachium acanthurus Ligia occidentalis Austrolestes annulosus Aspidomorpha tecta Palaemon paucidens Ligia oceanica Austrolestes leda Charidotella egregia Palaemonetes paladosus Idotea balthica Sympetrum pedemontanum Charidotella sexpuncata Palaemonetes pugio Sphaeroma quadridenatum Sympetrum sanguineum granti Palaemonetes vulgaris Sphaeroma serratum Sympetrum striolatum Dynastes hercules Palaemontes vulgaris Biston betularia Sympetrum vulgatum Dynastes hyllus Callinectis sapidus Battus philenor Kosciuscola tristis Carcinus maenas Aeshna brevistyla Locusta migratoria Cryptoglossa verrucosa Sesarma dehaani Aeshna caerulea Locustana pardalina Astacus leptodactylus Sesarma haematocheir Aeshna mixta Nomadacris septemfasciata Cambarellus shufeldtii Sesarma intermedia Anax imperator Schistocerca gregaria Oroconectes clypeatus Sesarma reticulatum Anax junius Schistocerca lineata Oroconectes immunis Upogebia affinis Argia apicalis Anabrus simplex Crago vulgaris Chasmagnathus granulatus Argia bipunctulata Carausius morosus Hippolyte cranchii Hemigrapsus oregonensis Argia extranea Squilla empusa Hippolyte varians Chaoborus crystallinus Argia moestra Cambarius clarkii Umbers et al (in review) Biological Reviews Arthropods that change colour

Types of colour change

ontogenetic morphological mechanical physiological each can occur via a diverse range of mechanisms ontological colour change mostly irreversible

Pat Backwell

Uca capricornis morphological colour change mostly reversible, slow

flickr.com Thomisus spectabilus mechanistic colour change reversible, fast

Kate Umbers Acripeza reticulata physiological colour change reversible fast

Kate Umbers Kosciuscola tristis fast, reversible colour change

physiological mechanistic mechanistic fast, reversible colour change

physiological mechanistic mechanistic Temperature dependent colour change in the chameleon grasshopper

Hypotheses tested:

aposematism thermoregulation female choice male competition Temperature dependent colour change in the chameleon grasshopper

Hypotheses tested:

aposematism thermoregulation female choice male competition Temperature dependent colour change in the chameleon grasshopper

Hypotheses tested:

aposematism thermoregulation female choice male competition* Temperature dependent colour change in the chameleon grasshopper

Hypotheses tested:

aposematism thermoregulation female choice male competition* males fight in Autumn

video: Nikolai J Tatarnic

Umbers, et al (2012) PLoS ONE, Umbers et al (2012) Behavioural Ecology and Sociobiology

temperature-controlled colour change colour

Umbers (2011) Journal of Physiology Key & Day (1954) Australian Journal of Zoology Filshie et al (1975) Journal of Insect Physiology threshold: temperature-controlled • black below 10°C • turquoise above 25°C

speed: • 30 min black to turquoise • 300 min turquoise to black

variation: change colour • maximum brightness varies among males • dramatic change in males • weak change in females

Umbers (2011) Journal of Insect Physiology Key & Day (1954) Australian Journal of Zoology Filshie et al (1975) Journal of Insect Physiology Colour change in thermoregulation

100

% reflectance 0

Umbers, Herberstein & Madin (2012) Journal of Insect Physiology Colour change in thermoregulation

100

% reflectance 0

difference: 0.55°C (0.1ms-1), 0.05°C (10ms-1)

Umbers, Herberstein & Madin (2012) Journal of Insect Physiology Colour change in thermoregulation

100

% reflectance 0

difference: 0.55°C (0.1ms-1), 0.05°C (10ms-1) how do we know if this is adaptive?

Umbers, Herberstein & Madin (2012) Journal of Insect Physiology Do use colour and colour change to thermoregulate?

We don’t really know. Dark colour Light colour Polymorphism TempDiff difference meridiana black black painted with an aqueous solution of lamp- -0.5 N Eristalis tenax black brown black artistsblack oil paint -0.3 N Sarcophaga carnaria black dull blue black black artists oil paint -0.3 N Phormia caerulia black metallic blue black painted with an aqueous solution of lamp- -0.2 N Lucilia caesar black metallic green painted with an aqueousblack solution of lamp- 0.0 N Schistocerca gregaria red green naturalblack 0.1 N Eleodes armata black white white paint 0.1 N Austrolestes annulosus black blue natural colour change 0.2 N Phymata americana black yellow both painted 0.3 Y Tetrix subulata black white natural colour morphs 0.4 Y Calliphora erythrocephala black dull blue-black black artists oil paint 0.4 N Carausius morosus black green painted with an aqueous solution of lamp- 0.7 N Periplaneta americana black brown painted with an aqueousblack solution of lamp- 0.8 N Vespa vulgaris black yellow and black painted with an aqueousblack solution of lamp- 1.1 N Periplaneta americana black brown black artistsblack oil paint 1.3 N Coccinella septempunctata large spots small spots natural (size of spots) 1.5 Y Melanoplus differentialis black yellow natural 1.6 N Adalia bipunctata two black spots orange natural (number of spots) 1.9 Y Pterid species moreon orange melanin less melanin natural variation in number of dark 2.0 Y Drepanosiphum platanoides melanic non-melanic colouredseasonal scales 2.1 Y Adalia bipunctata 2 spots 6 spots natural variation in number of spots of 2.1 Y Colias eurytheme black orange differentseasonal morphs 2.5 Y Schistocerca gregaria black dull yellow green painted with an aqueous solution of lamp- 3.3 N Brachystola magna black tan bothblack painted 3.7 Y Colias nastes melanic less melanin natural variation in number of dark 4.0 Y Carausius morosus black green painted with ancoloured aqueous scales solution of lamp- 4.4 N Calliptamus coelesyriensis chocolate (rarer) buff naturalblack fixed 5.0 Y Coccinella septempunctata black orange nail polish (black & clear) 6.0 Y Onymacris brincki black white sister species 6.0 Y Brachystola magna tan white both painted 8.3 Y Brachystola magna black white both painted 12.0 Y Parnassius phoebus more melanin less melanin natural variation in number of dark significant N coloured scales ly Temperature difference between insect colour morphs

8" p=0.003 7"

6"

5"

4"

3"

2"

difference"between"dark"and"light"(°C)" 1" n=15 n=16 0" important" not"important" author's"interpretaDon"of"temperature"difference" Limitations of past approaches

fixed / artificial colour polymorphisms

dead animals

artificial lighting

comparison between individuals / species evidence temperature differences are adaptive

selection experiments needed Fast, reversible colour change

physiological mechanistic mechanistic Startle colour in the alpine katydid

♀ ♂ Kate Umbers Kate Umbers Acripeza reticulata Startle colour in the alpine katydid

♀ ♂ Kate Umbers Kate Umbers Acripeza reticulata juveniles Startle colour in the alpine katydid

Questions:

What triggers their startle display? How unpalatable are Acripeza? Is unpalatability and display intensity linked? Who are their predators? How do display components work together? Startle colour in the alpine katydid

Questions:

What triggers their startle display? How unpalatable are Acripeza? Is unpalatability and display intensity linked? Who are their predators? How display components work together? Startle colour in the alpine katydid Startle colour in the alpine katydid

Treatments: Rated reaction: poke 1 no change pinch 2 blow on 3 over 4 poke next to 5 ‘full display’ 8 8 7 7 6 6 5 5 4 4 3 3 intensity of display 2 intensity of display 2 1 1 0 fly over poke next blow on pinch and poke on 0 Startle colourto in the alpinelift katydidabdomen fly over poke next blow on pinch treatmentand poke on to8 lift abdomen treatmentaverage before average after average7 before average after 6 5 4 3

intensity of display 2 1 0 fly over poke next blow on pinch and poke on to lift abdomen treatment average before average after Startle colour in the alpine katydid

Corvus mellori Fast, reversible colour change

physiological mechanistic mechanistic Startle colour in the myobatrachid frogs Startle colour in the myobatrachid frogs

Evolution of Myobatrachid frog colour Does colour convey toxicity? Is startle display effective against foes? How do display components work together?

Startle colour in the myobatrachid frogs

Evolution of Myobatrachid frog colour Does colour convey toxicity ? Is startle display effective against foes? How do display components work together? Evolution of Myobatrachid frog colour Evolution of Myobatrachid frog colour

Evolution of Myobatrachid frog colour

Fig 1. Plasticine model of Oophaga pumilio (A) and a brown coloured frog (B) placed on leaf litter

Saporito et al (2007) Experimental evidence for aposematism in the dendrobatid poison frog Oophaga pumilio Copeia 4: 1006 - 1011 Evolution of Myobatrachid frog colour

FROGBOT Evolution of Myobatrachid frog colour

to be continued...

(see you in July!) Acknowledgements

Arthropod colour change: University of Wollongong Scott Fabricant, Ainsley Seago, Australia & Pacific Science Foundation Felipe Gawryszewski and Marie Australian Biological Resource Study Herberstein Linnean Society of NSW Thredbo Sports Grasshopper colour change: The Orthopterists’ Society Nikolai Tatarnic, Marie Herberstein, Josh Madin and Greg Holwell

Katydid startle display: Johanna Mappes

Frog colour: Phillip Byrne, Scott Keogh, Dale Roberts and Allison Shaw Are the katydids toxic? diet heavy in Scenecio gunnii & S. pinnatifolius Is the grasshopper’s blue a warning signal?

50%#

n=64 n=64 40%#

30%#

20%#

10%#

0%# chi-square pearson: p = 0.04 Fisher’s exact: p = 0.07 Does brightness correlate with success?

1 Bradley Terry pairwise analysis 0.9 n interacons = 259 n grasshoppers = 74 0.8 Rank order correlaon 0.7 n = 74 r2 = 0.04, p = 0.77

0.6 effect F p 0.5 brightness 0.05 0.82

0.4 pronotum 0.27 0.61 L fore leg 2.18 0.15 Grasshopper brightness 0.3 R fore leg 0.01 0.98 0.2 L femur 2.03 0.16 0.1 R femur 3.11 0.08 0 mandible 1.01 0.32 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50 52 54 56 58 60 62 64 66 68 70 72 74 76 Grasshoppers in rank order of winningness Umbers et al (2013) Behavioural Ecology and Sociobiology Are males using their colour in assessment?

1 r = 0.53 0.9 p < 0.01 0.8 n = 32 0.7

0.6 0.5 colour matching in 0.4 field observations and laboratory 0.3 experiments Brightness of challenger 0.2

0.1

0 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 Brightness of primary

Umbers et al (2013) Behavioural Ecology and Sociobiology