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60-427: See text Etheostomatinae is a species-rich subfamily of freshwater ray-finned fish , the members of which are commonly known as the darters . The subfamily is part of the family Percidae which also includes the perches , ruffes and pikeperches . The family is endemic to North America. It consists of three to five different genera and well over 200 species. A common name for these fish in southern Indiana

120-597: A botanical subfamily. Detarioideae is a subdivision of the family Fabaceae (legumes), containing 84 genera. Stevardiinae is an example of a zoological subfamily. Stevardiinae is a large subdivision of the family Characidae , a diverse clade of freshwater fish . This biology article is a stub . You can help Misplaced Pages by expanding it . Carotenoid Carotenoids ( / k ə ˈ r ɒ t ɪ n ɔɪ d / ) are yellow, orange, and red organic pigments that are produced by plants and algae , as well as several bacteria, archaea, and fungi . Carotenoids give

180-455: A genus of darters (Percidae: Etheostoma). Copeia , 103 (1), 99–124. Near, T. J. (2002). Phylogenetic relationships of Percina (Percidae: Etheostomatinae). Copeia , 2002 (1), 1–14. Paine, M. D. (1990). Life history tactics of darters (Percidae: Etheostomatiini) and their relationship with body size, reproductive behaviour, latitude and rarity. Journal of Fish Biology , 37 (3), 473–488.m Sherman, A. N. (2022). Investigating Differences in

240-438: A mound-tube papillae allowing females to completely bury themselves as they lay eggs under the substrate. Egg clustering species tend to express complexes of pleated rosette papillae with ventral genital pores allowing the females to attach their eggs to the underhangs of rocks and other objects. Selective traits may vary across species and genera, female darters have been observed to lay 500 to 1000 eggs sequentially, alluding to

300-427: A number of different steps within the carotenoid biosynthetic pathway. Glyceraldehyde 3-phosphate and pyruvate , intermediates of photosynthesis , are converted to deoxy-D-xylulose 5-phosphate (DXP) catalyzed by DXP synthase (DXS). DXP reductoisomerase catalyzes the reduction by NADPH and subsequent rearrangement. The resulting MEP is converted to 4-(cytidine 5’-diphospho)-2-C-methyl-D-erythritol (CDP-ME) in

360-408: A promiscuous mating system with limited female mate preference and no parental care. Observations have been made that the majority of species across the diverse genus Etheostoma and that they attach eggs to their surroundings for protection. These egg attaching species mate when a female selects a location above the substrate such as a plant or rock where she will deposit her eggs, the male then mounts

420-461: A single enzyme, the bacterial phytoene desaturase (CRTI) for the catalysis. Plants and cyanobacteria however utilize four enzymes for this process. The first of these enzymes is a plant-type phytoene desaturase which introduces two additional double bonds into 15-cis-phytoene by dehydrogenation and isomerizes two of its existing double bonds from trans to cis producing 9,15,9’-tri-cis-ζ-carotene. The central double bond of this tri-cis-ζ-carotene

480-453: A species. Coloration has been viewed to be heightened through ingestion of carotenoids , which could explain how coloration is phenotypically correlated to fitness across chromatic species of Etheostomatinae . Another prominent feature held by females across darter lineage is the presence of gential papillae , fleshy protuberances between the anus and anal fin and are associated with egg laying abilities. Phylogenetic analyses expressed that

540-456: A subgenus of Etheostoma . Etheostomatinae, as a species rich and diverse sub-family within the family Percidae of freshwater ray-finned fishes, is composed of multiple genera including Ammocrypta, Crystalluria, Etheostoma, Nothonotus, and Percina, all of which have been observed as monophyletic groupings through molecular phylogenies. Speciation events can occur to a culmination of factors as both sympatric and allopatric speciation

600-407: Is a higher energy state and is essential in photoprotection. Light produces damaging species during photosynthesis, with the most damaging being reactive oxygen species (ROS). As these high energy ROS are produced in the chlorophyll the energy is transferred to the carotenoid’s polyene tail and undergoes a series of reactions in which electrons are moved between the carotenoid bonds in order to find

660-468: Is a virulence factor with an antioxidant action that helps the microbe evade death by reactive oxygen species used by the host immune system. The basic building blocks of carotenoids are isopentenyl diphosphate (IPP) and dimethylallyl diphosphate (DMAPP). These two isoprene isomers are used to create various compounds depending on the biological pathway used to synthesize the isomers. Plants are known to use two different pathways for IPP production:

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720-441: Is cyclized; cyclization gives rise to carotenoid diversity, which can be distinguished based on the end groups. There can be either a beta ring or an epsilon ring, each generated by a different enzyme ( lycopene beta-cyclase [beta-LCY] or lycopene epsilon-cyclase [epsilon-LCY]). α-Carotene is produced when the all-trans lycopene first undergoes reaction with epsilon-LCY then a second reaction with beta-LCY; whereas β-carotene

780-512: Is derived from herbicides applied to soil and binds to DXP synthase. This inhibits DXP synthase, preventing synthesis of DXP and halting the MEP pathway. The use of this toxin leads to lower levels of carotenoids in plants grown in the contaminated soil. Fosmidomycin , an antibiotic , is a competitive inhibitor of DXP reductoisomerase due to its similar structure to the enzyme. Application of said antibiotic prevents reduction of DXP, again halting

840-437: Is directly related to their structure, especially the length of the conjugation. Xanthophylls are often yellow, giving their class name. Carotenoids also participate in different types of cell signaling. They are able to signal the production of abscisic acid , which regulates plant growth, seed dormancy , embryo maturation and germination , cell division and elongation, floral growth, and stress responses. The length of

900-460: Is eliminated from the CDP-ME2P molecule. MECDP is then converted to (e)-4-hydroxy-3-methylbut-2-en-1-yl diphosphate (HMBDP) via HMBDP synthase in the presence of flavodoxin and NADPH. HMBDP is reduced to IPP in the presence of ferredoxin and NADPH by the enzyme HMBDP reductase . The last two steps involving HMBPD synthase and reductase can only occur in completely anaerobic environments. IPP

960-554: Is imperative that female birds carefully select high quality mates. Current literature supports the theory that vibrant carotenoid coloration is correlated with male quality—either though direct effects on immune function and oxidative stress, or through a connection between carotenoid metabolizing pathways and pathways for cellular respiration. It is generally considered that sexually selected traits, such as carotenoid-based coloration, evolve because they are honest signals of phenotypic and genetic quality. For instance, among males of

1020-400: Is isomerized by the zeta-carotene isomerase Z-ISO and the resulting 9,9'-di-cis-ζ-carotene is dehydrogenated again via a ζ-carotene desaturase (ZDS) . This again introduces two double bonds, resulting in 7,9,7’,9’-tetra-cis-lycopene. CRTISO , a carotenoid isomerase, is needed to convert the cis -lycopene into an all-trans lycopene in the presence of reduced FAD . This all-trans lycopene

1080-403: Is no correlation between consumption of foods high in carotenoids and vitamin A and the risk of Parkinson's disease . Humans and other animals are mostly incapable of synthesizing carotenoids, and must obtain them through their diet. Carotenoids are a common and often ornamental feature in animals. For example, the pink color of salmon , and the red coloring of cooked lobsters and scales of

1140-440: Is not common within the darter subfamily, it has developed across Etheostomatinae's evolutionary history multiple times, once in the clade Goneaperca , and a more novel evolutionarily derived clade Nothonotus . Male parental care across these clades have expressed evolutionary reversals back to the lack of care for their offspring multiple times. There are at least 250 species of darters with known reproductive methodologies across

1200-552: Is observed across darter evolutionary history. Factors related to spatiotemporal habitat stability and life history strategies can play into speciation and genetic differentiation of darters, such as dispersal ability and gene flow in headwater reproductive habitats impacting lineage divergence . It was seen that habitat preference and isolation is a major factor contributing to speciation in darters. Speciation events are common across species of this subfamily, one example includes Barcheek darters, an ancient monophyletic grouping residing in

1260-730: Is pollywog. Species within the Etheostomatinae are all small fish, mostly less than 11 centimetres (4.3 in) in length, and their bodies are slightly compressed or fusiform in shape. They have two pterygiophores between the first and second dorsal fins which do not have spines and a reduced swimbladder which may be completely lacking. The common name "darter" owes to the behavior of the fish, which dart around their benthic habitat. They are sexually dimorphic ; most species have males with bright colors and patterning, particularly when breeding. These colors and patterns are used to attract females and allow for recognition of species, as

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1320-417: Is predominantly found within species of large geographic ranges, larger populations, and egg-burying reproductive mechanisms. Darters are composed of fusiform shaped bodies that are typically less than 11 centimeters in length, and characteristically host a reduced or lack an air bladder, allowing them to reside in benthic regions of lakes, streams, and rivers. A prominent feature across many species of darters

1380-513: Is produced by two reactions with beta-LCY. α- and β-Carotene are the most common carotenoids in the plant photosystems but they can still be further converted into xanthophylls by using beta-hydrolase and epsilon-hydrolase, leading to a variety of xanthophylls. It is believed that both DXS and DXR are rate-determining enzymes, allowing them to regulate carotenoid levels. This was discovered in an experiment where DXS and DXR were genetically overexpressed, leading to increased carotenoid expression in

1440-628: Is their diverse coloration. Color preferences have been found in common darter predators such as the largemouth bass, imposing selective pressures on certain species of darters limiting nuptial coloration. Chromatic species of darters are more prominent in fast flowing habitats with coarse substrate present within the water column and low flow accumulation, more light penetration, and fewer predators present. Species with blue fins are present in habitats supporting coarse substrate along with low flow accumulation, while blue body and red fins are commonly present in rifle ecosystems, and species with red coloration on

1500-414: Is then able to isomerize to DMAPP via IPP isomerase. Two GGPP molecules condense via phytoene synthase (PSY), forming the 15-cis isomer of phytoene . PSY belongs to the squalene/phytoene synthase family and is homologous to squalene synthase that takes part in steroid biosynthesis. The subsequent conversion of phytoene into all-trans- lycopene depends on the organism. Bacteria and fungi employ

1560-506: The Etheostomatinae sub-family with roughly 40 species experiencing paternal care and the rest showing no parental care at all. Males, while predominantly giving no parental care, do invest in courtship along with egg guarding among certain species. Intra-sexual and intersexual interactions affect speciation via sexual selection , predominantly via behavioral isolation through male aggression more so than female biases, likely due to

1620-745: The Mississippi River basin and the drainages of the Great Lakes , Hudson Bay , the seaboards of the Atlantic and the Gulf of Mexico and the Pacific coast of Mexico. Fishbase recognises five genera as follows: However, Fishbase places Crystallaria within the subfamily Percinae while the 5th Edition of Fishes of the World regard it as a subgenus of Ammocrypta and Nothonotus as

1680-488: The Gene Expression Profiles of Rainbow Darter (Percidae: Etheostoma caeruleum) Populations Across Space and Time (Doctoral dissertation, Southeastern Louisiana University). Williams, T. H., Gumm, J. M., & Mendelson, T. C. (2013). Sexual selection acting on a speciation trait in darters (Percidae: Etheostoma). Behavioral Ecology , 24 (6), 1407–1414. Subfamily Detarioideae is an example of

1740-549: The ability to utilize these pigments for external coloration has evolved independently many times throughout avian evolutionary history. Carotenoid coloration exhibits high levels of sexual dimorphism , with adult male birds generally displaying more vibrant coloration than females of the same species. These differences arise due to the selection of yellow and red coloration in males by female preference . In many species of birds, females invest greater time and resources into raising offspring than their male partners. Therefore, it

1800-669: The bird species Parus major , the more colorfully ornamented males produce sperm that is better protected against oxidative stress due to increased presence of carotenoid antioxidants . However, there is also evidence that attractive male coloration may be a faulty signal of male quality. Among stickleback fish, males that are more attractive to females due to carotenoid colorants appear to under-allocate carotenoids to their germline cells. Since carotinoids are beneficial antioxidants, their under-allocation to germline cells can lead to increased oxidative DNA damage to these cells. Therefore, female sticklebacks may risk fertility and

1860-403: The body are present in small streams. The size of darters can be linked to increased fitness as larger darters tend to grow faster, produce more offspring, mature earlier at a larger size, and have longer lifespans along with reproductive lifespans. Diversity of life species traits across darters have been linked to influences such as reproductive behavior, geographic range, and population size of

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1920-590: The characteristic color to pumpkins , carrots , parsnips , corn , tomatoes , canaries , flamingos , salmon , lobster , shrimp , and daffodils . Over 1,100 identified carotenoids can be further categorized into two classes – xanthophylls (which contain oxygen) and carotenes (which are purely hydrocarbons and contain no oxygen). All are derivatives of tetraterpenes , meaning that they are produced from 8 isoprene units and contain 40 carbon atoms. In general, carotenoids absorb wavelengths ranging from 400 to 550 nanometers (violet to green light). This causes

1980-433: The color divergence is due to intra-sexual mechanisms in certain species lacking parental care, and that male coloration and aggressiveness may have coevolved. It has been viewed across species exhibiting parental care that achromatic coloration can be a trade off between paternal care and energy allocation for reproduction. Nuptial color preference has been observed across females of the darter species E. barrenses supporting

2040-429: The colors and patterns are particular to each species. Many species use typical sites for spawning and they care for their eggs and fry. They have evolved a variety of methods of depositing eggs and these include burying them, which may be the basal habit as it is found in all genera, as well as attaching eggs to a substrate and egg clustering. Etheostomatinae darters are endemic to North America where they are found in

2100-427: The common ancestor of Etheostomatinae buried eggs with the help of tube-like appendage with a genital pore, a pore that releases the females eggs, held at the dorsal-posterior end known as tube papillae, leading researchers to believe this is the ancestral state of darters. Tube papillae are present in majority egg laying species across Etheostomatinae , with the exception of the genus Nothonotus , which instead have

2160-629: The compounds to be deeply colored yellow, orange, or red. Carotenoids are the dominant pigment in autumn leaf coloration of about 15-30% of tree species, but many plant colors, especially reds and purples, are due to polyphenols . Carotenoids serve two key roles in plants and algae: they absorb light energy for use in photosynthesis , and they provide photoprotection via non-photochemical quenching . Carotenoids that contain unsubstituted beta-ionone rings (including β-carotene , α-carotene , β-cryptoxanthin , and γ-carotene ) have vitamin A activity (meaning that they can be converted to retinol ). In

2220-558: The cytosolic mevalonic acid pathway (MVA) and the plastidic methylerythritol 4-phosphate (MEP). In animals, the production of cholesterol starts by creating IPP and DMAPP using the MVA. For carotenoid production plants use MEP to generate IPP and DMAPP. The MEP pathway results in a 5:1 mixture of IPP:DMAPP. IPP and DMAPP undergo several reactions, resulting in the major carotenoid precursor, geranylgeranyl diphosphate (GGPP). GGPP can be converted into carotenes or xanthophylls by undergoing

2280-468: The development of monoclonal antibodies to trans- lycopene it was possible to localise this carotenoid in different animal and human cells. Beta-carotene , found in pumpkins , sweet potato , carrots and winter squash , is responsible for their orange-yellow colors. Dried carrots have the highest amount of carotene of any food per 100-gram serving, measured in retinol activity equivalents (provitamin A equivalents). Vietnamese gac fruit contains

2340-432: The disappearance of chlorophyll. Carotenoids are responsible for the brilliant yellows and oranges that tint deciduous foliage (such as dying autumn leaves ) of certain hardwood species as hickories , ash , maple , yellow poplar , aspen , birch , black cherry , sycamore , cottonwood , sassafras , and alder . Carotenoids are the dominant pigment in autumn leaf coloration of about 15-30% of tree species. However,

2400-441: The eastern highlands which under time-calibrated molecular phylogenetic analysis of the mitochondrial genome has presented five novel cryptic species previously thought to be associated with Etheostoma basilare . Hybridization has been accounted for multiple times across Etheostomatinae. It is more common between species with similar egg-laying behavioral mechanisms and within the same phylogenetic clade, with sperm viability being

2460-446: The evolution of male parental care in darters (Teleostei: Percidae: Etheostomatinae). Journal of Evolutionary Biology , 25 (5), 836–846. Martin, M. D., & Mendelson, T. C. (2016). Male behaviour predicts trait divergence and the evolution of reproductive isolation in darters (Percidae: Etheostoma). Animal Behaviour , 112 , 179–186. Martin, Z. P., & Page, L. M. (2015). Comparative morphology and evolution of genital papillae in

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2520-482: The eye, lutein , meso -zeaxanthin , and zeaxanthin are present as macular pigments whose importance in visual function, as of 2016, remains under clinical research . Carotenoids are produced by all photosynthetic organisms and are primarily used as accessory pigments to chlorophyll in the light-harvesting part of photosynthesis. They are highly unsaturated with conjugated double bonds , which enables carotenoids to absorb light of various wavelengths . At

2580-421: The fatty tissues of animals, and exclusively carnivorous animals obtain the compounds from animal fat. In the human diet, absorption of carotenoids is improved when consumed with fat in a meal. Cooking carotenoid-containing vegetables in oil and shredding the vegetable both increase carotenoid bioavailability . The most common carotenoids include lycopene and the vitamin A precursor β-carotene. In plants,

2640-405: The female and as they vibrate the females attach eggs to her location of choosing while the male deposits sperm fertilizing the eggs. Other egg-laying behavioral characteristics include egg-guarding in which a female are courted by a male guarding a cavity, typically under flat stones, the couple then invert as the female lays eggs and the cavity ceiling as the male fertilizes them. When burying eggs,

2700-446: The highest known concentration of the carotenoid lycopene . Although green, kale , spinach , collard greens , and turnip greens contain substantial amounts of beta-carotene. The diet of flamingos is rich in carotenoids, imparting the orange-colored feathers of these birds. Reviews of preliminary research in 2015 indicated that foods high in carotenoids may reduce the risk of head and neck cancers and prostate cancer . There

2760-1424: The idea that female choice can drive the evolution of male coloration; while this may not apply to all species of darters, it does help support the general idea behind coloration and how it develops under the realm of sexual selection. Fluker, B. L., Kuhajda, B. R., & Harris, P. M. (2014). The influence of life-history strategy on genetic differentiation and lineage divergence in darters (Percidae: Etheostomatinae). Evolution , 68 (11), 3199–3216. Héjja-Brichard, Y., Renoult, J. P., & Mendelson, T. C. (2022). Preference for conspecific mates in sympatric and allopatric darters (genus Etheostoma): comparative evidence for geographical and sex effects. Hollingsworth Jr, P. R., & Near, T. J. (2009). Temporal patterns of diversification and microendemism in Eastern Highland endemic barcheek darters (Percidae: Etheostomatinae). Evolution , 63 (1), 228–243. Hopper, G. W. (2015). Ecological and morphological variation of darters among assemblages in Oklahoma streams (Doctoral dissertation, Kansas State University). Keck, B. P., & Near, T. J. (2009). Patterns of natural hybridization in darters (Percidae: Etheostomatinae). Copeia , 2009 (4), 758–773. Kelly, N. B., Near, T. J., & Alonzo, S. H. (2012). Diversification of egg‐deposition behaviours and

2820-419: The idea that males expending more energy into courtship reproduction than females. Male darters have been found to hold female preferences and mate choice and aggression correlated to their coloration differences expressing biases regarding intra and intersexual selection. Coloration has been viewed to be correlated more with male aggression as opposed to previously suggested female selective biases expressing that

2880-511: The limiting factor of hybridization. Sperm viability has also been witnessed to decrease as temperatures increase as an extraneous factor. Other factors impacting the viability of hybridization include life history of sympatry as opposed to allopatry, habitat preference, similar spawning periods, and duration of sperm viability. Species known to bury their eggs have been viewed to reproduce under sympatric conditions while egg-guarding species typically reproduce under allopatric conditions. Hybridization

2940-411: The male initiates with a courtship dance, in response the female wiggles until she is partially buried in the substrate. The male then mounts the female and similar to other species of darter, they vibrate as the females release eggs into her desired location while the male deposits sperm atop the eggs. Typically no parental care is associated with egg-burying species across the genus. While parental care

3000-571: The most balanced (lowest energy) state for the carotenoid. Carotenoids defend plants against singlet oxygen , by both energy transfer and by chemical reactions. They also protect plants by quenching triplet chlorophyll. By protecting lipids from free-radical damage, which generate charged lipid peroxides and other oxidised derivatives, carotenoids support crystalline architecture and hydrophobicity of lipoproteins and cellular lipid structures, hence oxygen solubility and its diffusion therein. Like some fatty acids , carotenoids are lipophilic due to

3060-424: The multiple conjugated double bonds determines their color and photophysics. After absorbing a photon, the carotenoid transfers its excited electron to chlorophyll for use in photosynthesis. Upon absorption of light, carotenoids transfer excitation energy to and from chlorophyll . The singlet-singlet energy transfer is a lower energy state transfer and is used during photosynthesis. The triplet-triplet transfer

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3120-406: The presence of CTP using the enzyme MEP cytidylyltransferase. CDP-ME is then converted, in the presence of ATP , to 2-phospho-4-(cytidine 5’-diphospho)-2-C-methyl-D-erythritol (CDP-ME2P). The conversion to CDP-ME2P is catalyzed by CDP-ME kinase . Next, CDP-ME2P is converted to 2-C-methyl-D-erythritol 2,4-cyclodiphosphate (MECDP). This reaction occurs when MECDP synthase catalyzes the reaction and CMP

3180-399: The presence of long unsaturated aliphatic chains. As a consequence, carotenoids are typically present in plasma lipoproteins and cellular lipid structures. Carotenoids are located primarily outside the cell nucleus in different cytoplasm organelles, lipid droplets , cytosomes and granules. They have been visualised and quantified by raman spectroscopy in an algal cell. With

3240-524: The reds, the purples, and their blended combinations that decorate autumn foliage usually come from another group of pigments in the cells called anthocyanins . Unlike the carotenoids, these pigments are not present in the leaf throughout the growing season, but are actively produced towards the end of summer. Dietary carotenoids and their metabolic derivatives are responsible for bright yellow to red coloration in birds. Studies estimate that around 2956 modern bird species display carotenoid coloration and that

3300-417: The resulting seedlings. Also, J-protein (J20) and heat shock protein 70 (Hsp70) chaperones are thought to be involved in post-transcriptional regulation of DXS activity, such that mutants with defective J20 activity exhibit reduced DXS enzyme activity while accumulating inactive DXS protein. Regulation may also be caused by external toxins that affect enzymes and proteins required for synthesis. Ketoclomazone

3360-759: The same time, the terminal groups regulate the polarity and properties within lipid membranes . Most carotenoids are tetraterpenoids , regular C 40 {\displaystyle {\ce {C40}}} isoprenoids . Several modifications to these structures exist: including cyclization , varying degrees of saturation or unsaturation, and other functional groups . Carotenes typically contain only carbon and hydrogen, i.e., they are hydrocarbons . Prominent members include α-carotene , β-carotene , and lycopene , are known as carotenes . Carotenoids containing oxygen include lutein and zeaxanthin . They are known as xanthophylls . Their color, ranging from pale yellow through bright orange to deep red,

3420-409: The sweet floral smells present in black tea , aged tobacco , grape , and many fruits are due to the aromatic compounds resulting from carotenoid breakdown. Some carotenoids are produced by bacteria to protect themselves from oxidative immune attack. The aureus (golden) pigment that gives some strains of Staphylococcus aureus their name is a carotenoid called staphyloxanthin . This carotenoid

3480-453: The viability of their offspring by choosing redder, but more deteriorated partners with reduced sperm quality. Products of carotenoid degradation such as ionones , damascones and damascenones are also important fragrance chemicals that are used extensively in the perfumes and fragrance industry. Both β-damascenone and β-ionone although low in concentration in rose distillates are the key odor-contributing compounds in flowers. In fact,

3540-488: The xanthophyll lutein is the most abundant carotenoid and its role in preventing age-related eye disease is currently under investigation. Lutein and the other carotenoid pigments found in mature leaves are often not obvious because of the masking presence of chlorophyll . When chlorophyll is not present, as in autumn foliage, the yellows and oranges of the carotenoids are predominant. For the same reason, carotenoid colors often predominate in ripe fruit after being unmasked by

3600-403: The yellow morph of common wall lizards are due to carotenoids. It has been proposed that carotenoids are used in ornamental traits (for extreme examples see puffin birds) because, given their physiological and chemical properties, they can be used as visible indicators of individual health, and hence are used by animals when selecting potential mates. Carotenoids from the diet are stored in

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