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The role of melanism in oncillas on the temporal segregation of nocturnal activity

The role of melanism in oncillas on the temporal segregation of nocturnal activity The occurrence of coat colour polymorphisms in populations may promote the ecological success of species by permitting a wider spectrum of use of different subsets of available resources. We conducted an analysis of temporal segregation by comparing night brightness with nocturnal activity of spotted and melanistic oncillas (Leopardus tigrinus). Melanistic oncillas were more active during bright nights and spotted oncillas and other species were more active during dark nights. Each colour morph occupied a temporal niche outside the confidence interval of the other colour morph, demonstrating the ecological significance of polymorphic colour patterns in this felid species. Keywords: Leopardus tigrinus, activity, moonlight, camera trap. O papel do melanismo na segregação temporal da atividade noturna do gato-do-mato-pequeno Resumo A ocorrência de polimorfismo no padrão de pelagem de populações pode promover o sucesso ecológico das espécies por permitir um amplo espectro de uso de diferentes parcelas de recursos disponíveis. Nós testamos a existência de diferença na segregação temporal do gato-do-mato-pequeno (Leopardus tigrinus), comparando a luminosidade durante períodos de atividade noturna de indivíduos pintados e melânicos. Indivíduos melânicos de gato-do-mato-pequeno foram mais ativos durante noites claras e indivíduos pintados de gato-do-mato-pequeno e outras espécies foram mais ativas durante noites escuras. Cada forma de coloração ocupou um nicho temporal fora do intervalo de confiança do outro, demonstrando a significância ecológica dos padrões de polimorfismo de colorações nesta espécie de felino. Palavras-chave: Leopardus tigrinus, atividade, luminosidade da lua, armadilha fotográfica. 1. Introduction Melanism is a ubiquitous phenomenon in the animal some populations and hence it is apparently adaptative in kingdom that has been used to investigate evolutionary nature (Eizirik et al., 2003). Similar to other animal groups shifts (Majerus and Mundy, 2003). Most mammal species (invertebrates, reptiles and birds), this polymorphism has possess variations in coat colour within populations, but originated several times independently in felids. However, only a few species exhibit discontinuous variations (Majerus the functional advantages of melanism remain an open and Mundy, 2003). Eleven out of 38 felid species carry field for empirical and theoretical development (Caro, mutations that increase the amount of melanin, resulting in 2005). While the genetic and molecular basis of melanism black or nearly black coats of melanistic individuals that in mammals have been clarified (Nachman et al., 2003; coexist with wild-type individuals with spotted or uniform Eizirik et al., 2003), we have little evidence linking the tan coat colours. Top cats such as the jaguar Panthera onca genetic information to maintenance of the melanistic (Linnaeus), or small cats such as the domestic cat Felis phenotype in the wild. The melanistic phenotype must bring silvestris catus, exhibit melanism in high frequencies in adaptative advantages. Therefore, ecological information 142 142 Braz. J. Biol., 2014,  vol. 74, no. 3 (suppl.), p. S142-S145 Melanism and moonlight about polymorphic individuals in the wild will be determinant hour apart as independent. The percentage of nocturnal in the understanding of the genetic maintenance of these brightness was obtained by Moonrise 3.5 software for polymorphisms. each nocturnal independent record, discarding crepuscular The increase in the frequency of the black moth after records. Nocturnal records obtained before moonrise and industrialisation (Grant et al., 1996; Majerus, 1998), and its after moonset were considered zero percent brightness. decline after air quality improvement (Saccheri et al., 2008) Nebulosity data were obtained in a meteorological station are emblematic examples of natural selection decreasing 40 km far from our study area, which is an index varying or increasing predation rates of the different colour forms. from 0 to 10. Desert rodent species that present intraspecific coat colour Differences between brightness associated with activity variation show a strong correlation between their coat colour of spotted and melanistic morphs, and other wild felid and the substrate colour, and this camouflage is an adaptive species and small mammals [rodents and Philander frenatus phenotype against predation by rapine birds (reviewed (Olfers, 1818)] captured during the sampling effort, were by Majerus and Mundy, 2003). Hoekstra and Nachman compared by analysis of variance. In addition, we used the (2003) have demonstrated the link between genotype and logistic regression to test the relationship between each phenotype in these rodents, where black rodents in dark record of melanistic/spotted animal (binary variable) with moon brightness and nebulosity. substrate habitat present high frequency of the mutated alleles responsible for melanism. In contrast to moths and desert rodents, the adaptive benefits of melanism in felids 3. Results remain obscure. We recorded 170 independent observations of oncillas, Forsman et al. (2008) hypothesised that the evolutionary 139 of spotted and 31 of melanistic forms (Figure 1). persistence of polymorphisms in populations is because they Additionally, we recorded 15 independent nocturnal promote the ecological success of species by permitting brightless observations of margay, 35 of ocelot, 19 of the use of a wider spectrum of available resources. This cougar and 132 of small mammals. hypothesis argues that behavioural observations of resource There was no trend towards either nocturnal or diurnal use by individuals that belong to different colour morphs activity of oncillas (Figure 2), and there was no difference will demonstrate that each morph explores different in the distribution of arrhythmic activity patterns between niche dimensions. Thus, we conducted a test for temporal spotted and melanistic forms (Watson’s U test = 0.056; segregation by melanistic and spotted oncilla Leopardus n1 = 31; n2 = 139; P > 0.5) (Figure 2). tigrinus (Schreber), one of the smallest spotted wild cats of America, which has a silhouette and size resembling a house cat (Nowell and Jackson, 1996). We compared the temporal segregation of nocturnal activity by lunar brightness for each colour morph using the largest known sample of melanistic individuals of L. tigrinus. We expected that moon brightness has some role in the activity pattern of melanistic L. trigrinus. So we expect that one of the two morphs of this felid species could be favoured by the brightness of moonlight, in this case, the melanistic form. It is known that moonlight influences the activity of some mammal species like marsupials (Julien-Laferrière, 1997) and bats (Esbérard, 2007). In this sense, we expected something similar to L. tigrinus, related to frequency of coat colour and moonlight. 2. Material and Methods Between January 2005 and July 2009 the activity of oncillas was recorded by remote camera traps (n = 30; Tigrinus®) in a sampling effort of 8,500 trap-days in four areas inside two Tropical Rain Forest Reserves in southern Brazil (Caraguatá Ecological Reserve: 27°27’S, 48°57’W – 4,200 ha; Serra do Tabuleiro State Park: 28°26’S, 48°50’W – 85.000 ha). A detailed description of the study areas and sampling method is available in Goulart et al. (2009). We compared the distribution of the circadian activity pattern of spotted and melanistic morphs using Watson’s U Test for circular data, using the Figure 1. Spotted (top) and melanistic (bottom) oncillas Oriana 3.0 software. We considered only records taken one recorded by camera-traps. Braz. J. Biol., 2014,  vol. 74, no. 3 (suppl.), p. S142-S145 143 143 Graipel, ME. et al. However, melanistic oncillas were more active during p = 0.007 for moon brightness, and X = 0.17; df = 1; bright nights (mean nocturnal brightness = 75.3%; n = 11; p = 0.687 for nebulosity). SE = 10.1; CI = 55.5 – 95.0) than spotted oncillas (mean nocturnal brightness = 38.5%; n = 67; SE = 5.0; CI = 28.8 4. Discussion – 48.2), and the activity of melanistic oncillas was also Oncillas presented arrhythmic activity pattern similar significantly higher during bright nights than other wild to that obtained by Tortato and Oliveira (2005) and felid species, L.wiedii (Schinz), L. pardalis (Linnaeus,), Oliveira-Santos et al. (2012) for the same region and and Puma concolor (Linnaeus), and small mammals species. However, the latter authors recorded variations (F = 2.601; P = 0.025) (Figure 3). in activity distribution of oncillas associated with the The multiple logistic regression between melanistic presence of other felid species. and spotted frequencies with regard to moon brightness Individuals from a single population may differ in several and nebulosity confirmed the same relationship above, aspects in the use of resources, for instance, between sexes being significant (X = 7.71; df = 2; p = 0.021); however, (Schoener, 1967), age-group (Cordero and Nicolas, 1987) only brightness contributed to explain melanistic and 2 and among trophic polymorphic individuals (Swanson et al., spotted frequencies in simple analysis (X = 7.35; df = 1; 2003). The temporal segregation between different colour coat phenotypes observed in our study corroborates Forsman´s prediction of the ecological significance of polymorphic colour patterns (Forsman et al., 2008). In the partitioning of resources between and within species, food and habitat sources are considered more important than the daily distribution of activity (Schoener, 1974). On the other hand, the activity pattern through interference competition increases in importance (Carothers and Jaksic, 1984), because territorial Neotropical carnivores could present potentially lethal damage between each other (Palomares and Caro, 1999; Di Bitetti et al., 2009). Melanistic individuals could be more effectively cryptic on bright nights than spotted individuals. According to our data, the space and the food resources available during bright nights are under-explored by other nocturnal predators present in the study area (e.g. L. wiedii, L. pardalis and Puma concolor). Most rodents, the main prey consumed by oncillas and other felid competitors (Oliveira et al., 2010), could present moon phobia, and thus decrease in Figure 2. Circadian activity of oncillas in Atlantic Rain availability to the predators during these moon phases Forest of southern Brazil. White bars represent registers (Lockard and Owings, 1974; Kaufman and Kaufman, of spotted (n = 139), and black bars, registers of melanistic 1982; Price et al., 1984). These ideas lead us to two non- oncillas (n = 31). excluding hypotheses to explain the observed pattern: (i) the melanistic individuals are more cryptic for their prey, ambushing prey more easily than other competitors on bright nights, compensating for the decrease in prey activity; (ii) poor quality foraging on bright nights did not compensate the high exposure of spotted oncillas to predators, but dark coat colour could offset this risk. Irrespective of the process underlying the observed pattern of reduction in intraspecific competition, attributed to differential use of time, our results suggest that melanistic individuals could occupy an alternative and wider ecological niche in relation to spotted individuals. While urban moths and desert rodents use melanism for cryptic protection in their habitats (Majerus, 1998; Majerus and Mundy, 2003), the oncilla, in its multicolor tropical habitat, should benefit Figure 3. Activity of melanistic oncilla (n = 11), spotted from the variation in moonlight. These results help us oncilla (n = 67), margay (n = 15), ocelot (n = 35), cougar to shed some light onto the way natural selection acts to (n = 19) and small mammals (n = 132) regarding nocturnal maintain polymorphisms (Skúlason and Smith, 1995), brightless (mean ± 95% IC) in Atlantic Rain Forest of southern Brazil. in this case, melanistic forms in populations of oncillas. 144 144 Braz. J. Biol., 2014,  vol. 74, no. 3 (suppl.), p. S142-S145 Melanism and moonlight of Mammalogy, vol. 55, no. 1, p. 189-193. http://dx.doi. Acknowledgements org/10.2307/1379266. PMid:4819590 To Russel W. Coffin and FATMA for allowing field research MAJERUS, MEN., 1998. Melanism: evolution in action. Oxford: at the Reserves. To Conservation International-Brazil, Oxford University Press. Reserva Ecológica do Caraguatá and FUNPESQUISA/ MAJERUS, MEN. and MUNDy , NI., 2003. Mammalian melanism: UFSC for logistical support. LGROS is supported by natural selection in black and white. Trends in Genetics, vol. 19, CNPq. NCC is a CNPq research fellow in Brazil. no. 11, p. 585-588. http://dx.doi.org/10.1016/j.tig.2003.09.003. PMid:14585605 References NACHMAN, MW., HOEKSTRA, HE. and D’AGOSTINO, SL., 2003. The genetic basis of adaptive melanism in pocket mice. CORDERO, GA. and NICOLAS, RA., 1987. Feeding Habits Proceedings of the National Academy of Sciences of the United of the Opossum (Didelphis marsupialis) in Northern Venezuela. States of America, vol. 100, no. 9, p. 5268-5273. http://dx.doi. Fieldiana. Zoology, vol. 39, p. 125-132. org/10.1073/pnas.0431157100. PMid:12704245 CARO, T., 2005. The adaptative significance of coloration in NOWELL, K. and JACKSON, P., 1996. Wild cats: status survey mammals. Bioscience, vol. 55, no. 2, p. 125-136. http://dx.doi. and conservation action plan. Gland: IUCN. org/10.1641/0006-3568(2005)055[0125:TASOCI]2.0.CO;2. OLIVEIRA-SANTOS, LGR., GRAIPEL, ME., TORTATO, MA., CAROTHERS, JH. and JAKSIC, FM., 1984. Time as a niche ZUCCO, CA., CáCERES, NC. and GOULART, FVB., 2012. difference: the role of interference competition. Oikos, vol. 42, Density and activity flexibility of oncilla Leopardus tigrinus no. 3, p. 403-406. http://dx.doi.org/10.2307/3544413. (CARNIVORA, FELIDAE) in the Atlantic Forest of Southern Brazil. Zoologia. In press DI BITETTI, MS., DI BLANCO, YE., PEREIRA, JA., PAVIOLO, A. and PEREZ, IJ., 2009. Time partitioning favors the coexistence OLIVEIRA, TG., TORTATO, MA., SILVEIRA, L., KASPER, of sympatric crab-eating foxes (Cerdocyon thous) and pampas CB., MAZIM, FD., LUCHERINI, M., JáCOMO, AT., SOARES, foxes (Lycalopex gymnocercus). Journal of Mammalogy, vol. 90, JBG., MARQUES, RV. and SUNQUIST, M., 2010. Ocelot ecology no. 2, p. 479-490. http://dx.doi.org/10.1644/08-MAMM-A-113.1. and its effect on the small-felid guild in the lowland Neotropics. In MACDONALD, D. and LOVERIDGE, A. (Eds.). The Biology EIZIRIK, E., YUHKI, N., JOHNSON, WE., MENOTTI-RAyMOND, and Conservation of Wild Felid. Oxford: Oxford University M., HANNAH, SS. and O’BRIEN, SJ., 2003. Molecular genetics Press. p.559-580. and evolution of melanism in the cat family. Current Biology, vol. 13, no. 5, p. 448-453. http://dx.doi.org/10.1016/S0960- PALOMARES, F. and CARO, TM., 1999. Interspecific killing 9822(03)00128-3. PMid:12620197 among mammalian carnivores. American Naturalist, vol. 153, no. 5, p. 492-508. http://dx.doi.org/10.1086/303189. ESBéRARD, CEL., 2007. Influência do ciclo lunar na captura de morcegos Phyllostomidae. Iheringia Série Zoologia, vol. 97, no. PRICE, MV., WASER, NM. and BASS, TA., 1984. Effects 1, p. 81-85. http://dx.doi.org/10.1590/S0073-47212007000100012. of moonlight on microhabitat use by desert rodents. Journal of Mammalogy, vol. 65, no. 2, p. 353-356. http://dx.doi. FORSMAN, A., AHNESJö, J., CAESAR, S. and KARLSSON, org/10.2307/1381183. M., 2008. A model of ecological and evolutionary consequences of color polymorphism. Ecology, vol. 89, no. 1, p. 34-40. http:// SACCHERI, IJ., ROUSSET, F., WATTS, PC., BRAKEFIELD, dx.doi.org/10.1890/07-0572.1. PMid:18376544 PM. and COOK, LM., 2008. Selection and gene flow on a diminishing cline of melanic peppered moths. 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Effect moonlight on activity and microhabitat use by Ord’s kangaroo rat (Dipodomys minckleyi. Ecology, vol. 84, no. 6, p. 1441-1446. http://dx.doi. ordii). Journal of Mammalogy, vol. 63, no. 2, p. 309-312. http:// org/10.1890/02-0353. dx.doi.org/10.2307/1380644. TORTATO, M. and OLIVEIRA, TG., 2005. Ecology of the Oncilla LOCKARD, RB. and OWINGS, DH., 1974. Seasonal variation (Leopardus tigrinus) at Serra do Tabuleiro State Park, Southern in moonlight avoidance by bannertail kangaroo rats. Journal Brazil. Cat News, vol. 42, p. 28-30. Braz. J. Biol., 2014,  vol. 74, no. 3 (suppl.), p. S142-S145 145 145 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Brazilian Journal of Biology Unpaywall

The role of melanism in oncillas on the temporal segregation of nocturnal activity

Brazilian Journal of BiologyAug 1, 2014

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Abstract

The occurrence of coat colour polymorphisms in populations may promote the ecological success of species by permitting a wider spectrum of use of different subsets of available resources. We conducted an analysis of temporal segregation by comparing night brightness with nocturnal activity of spotted and melanistic oncillas (Leopardus tigrinus). Melanistic oncillas were more active during bright nights and spotted oncillas and other species were more active during dark nights. Each colour morph occupied a temporal niche outside the confidence interval of the other colour morph, demonstrating the ecological significance of polymorphic colour patterns in this felid species. Keywords: Leopardus tigrinus, activity, moonlight, camera trap. O papel do melanismo na segregação temporal da atividade noturna do gato-do-mato-pequeno Resumo A ocorrência de polimorfismo no padrão de pelagem de populações pode promover o sucesso ecológico das espécies por permitir um amplo espectro de uso de diferentes parcelas de recursos disponíveis. Nós testamos a existência de diferença na segregação temporal do gato-do-mato-pequeno (Leopardus tigrinus), comparando a luminosidade durante períodos de atividade noturna de indivíduos pintados e melânicos. Indivíduos melânicos de gato-do-mato-pequeno foram mais ativos durante noites claras e indivíduos pintados de gato-do-mato-pequeno e outras espécies foram mais ativas durante noites escuras. Cada forma de coloração ocupou um nicho temporal fora do intervalo de confiança do outro, demonstrando a significância ecológica dos padrões de polimorfismo de colorações nesta espécie de felino. Palavras-chave: Leopardus tigrinus, atividade, luminosidade da lua, armadilha fotográfica. 1. Introduction Melanism is a ubiquitous phenomenon in the animal some populations and hence it is apparently adaptative in kingdom that has been used to investigate evolutionary nature (Eizirik et al., 2003). Similar to other animal groups shifts (Majerus and Mundy, 2003). Most mammal species (invertebrates, reptiles and birds), this polymorphism has possess variations in coat colour within populations, but originated several times independently in felids. However, only a few species exhibit discontinuous variations (Majerus the functional advantages of melanism remain an open and Mundy, 2003). Eleven out of 38 felid species carry field for empirical and theoretical development (Caro, mutations that increase the amount of melanin, resulting in 2005). While the genetic and molecular basis of melanism black or nearly black coats of melanistic individuals that in mammals have been clarified (Nachman et al., 2003; coexist with wild-type individuals with spotted or uniform Eizirik et al., 2003), we have little evidence linking the tan coat colours. Top cats such as the jaguar Panthera onca genetic information to maintenance of the melanistic (Linnaeus), or small cats such as the domestic cat Felis phenotype in the wild. The melanistic phenotype must bring silvestris catus, exhibit melanism in high frequencies in adaptative advantages. Therefore, ecological information 142 142 Braz. J. Biol., 2014,  vol. 74, no. 3 (suppl.), p. S142-S145 Melanism and moonlight about polymorphic individuals in the wild will be determinant hour apart as independent. The percentage of nocturnal in the understanding of the genetic maintenance of these brightness was obtained by Moonrise 3.5 software for polymorphisms. each nocturnal independent record, discarding crepuscular The increase in the frequency of the black moth after records. Nocturnal records obtained before moonrise and industrialisation (Grant et al., 1996; Majerus, 1998), and its after moonset were considered zero percent brightness. decline after air quality improvement (Saccheri et al., 2008) Nebulosity data were obtained in a meteorological station are emblematic examples of natural selection decreasing 40 km far from our study area, which is an index varying or increasing predation rates of the different colour forms. from 0 to 10. Desert rodent species that present intraspecific coat colour Differences between brightness associated with activity variation show a strong correlation between their coat colour of spotted and melanistic morphs, and other wild felid and the substrate colour, and this camouflage is an adaptive species and small mammals [rodents and Philander frenatus phenotype against predation by rapine birds (reviewed (Olfers, 1818)] captured during the sampling effort, were by Majerus and Mundy, 2003). Hoekstra and Nachman compared by analysis of variance. In addition, we used the (2003) have demonstrated the link between genotype and logistic regression to test the relationship between each phenotype in these rodents, where black rodents in dark record of melanistic/spotted animal (binary variable) with moon brightness and nebulosity. substrate habitat present high frequency of the mutated alleles responsible for melanism. In contrast to moths and desert rodents, the adaptive benefits of melanism in felids 3. Results remain obscure. We recorded 170 independent observations of oncillas, Forsman et al. (2008) hypothesised that the evolutionary 139 of spotted and 31 of melanistic forms (Figure 1). persistence of polymorphisms in populations is because they Additionally, we recorded 15 independent nocturnal promote the ecological success of species by permitting brightless observations of margay, 35 of ocelot, 19 of the use of a wider spectrum of available resources. This cougar and 132 of small mammals. hypothesis argues that behavioural observations of resource There was no trend towards either nocturnal or diurnal use by individuals that belong to different colour morphs activity of oncillas (Figure 2), and there was no difference will demonstrate that each morph explores different in the distribution of arrhythmic activity patterns between niche dimensions. Thus, we conducted a test for temporal spotted and melanistic forms (Watson’s U test = 0.056; segregation by melanistic and spotted oncilla Leopardus n1 = 31; n2 = 139; P > 0.5) (Figure 2). tigrinus (Schreber), one of the smallest spotted wild cats of America, which has a silhouette and size resembling a house cat (Nowell and Jackson, 1996). We compared the temporal segregation of nocturnal activity by lunar brightness for each colour morph using the largest known sample of melanistic individuals of L. tigrinus. We expected that moon brightness has some role in the activity pattern of melanistic L. trigrinus. So we expect that one of the two morphs of this felid species could be favoured by the brightness of moonlight, in this case, the melanistic form. It is known that moonlight influences the activity of some mammal species like marsupials (Julien-Laferrière, 1997) and bats (Esbérard, 2007). In this sense, we expected something similar to L. tigrinus, related to frequency of coat colour and moonlight. 2. Material and Methods Between January 2005 and July 2009 the activity of oncillas was recorded by remote camera traps (n = 30; Tigrinus®) in a sampling effort of 8,500 trap-days in four areas inside two Tropical Rain Forest Reserves in southern Brazil (Caraguatá Ecological Reserve: 27°27’S, 48°57’W – 4,200 ha; Serra do Tabuleiro State Park: 28°26’S, 48°50’W – 85.000 ha). A detailed description of the study areas and sampling method is available in Goulart et al. (2009). We compared the distribution of the circadian activity pattern of spotted and melanistic morphs using Watson’s U Test for circular data, using the Figure 1. Spotted (top) and melanistic (bottom) oncillas Oriana 3.0 software. We considered only records taken one recorded by camera-traps. Braz. J. Biol., 2014,  vol. 74, no. 3 (suppl.), p. S142-S145 143 143 Graipel, ME. et al. However, melanistic oncillas were more active during p = 0.007 for moon brightness, and X = 0.17; df = 1; bright nights (mean nocturnal brightness = 75.3%; n = 11; p = 0.687 for nebulosity). SE = 10.1; CI = 55.5 – 95.0) than spotted oncillas (mean nocturnal brightness = 38.5%; n = 67; SE = 5.0; CI = 28.8 4. Discussion – 48.2), and the activity of melanistic oncillas was also Oncillas presented arrhythmic activity pattern similar significantly higher during bright nights than other wild to that obtained by Tortato and Oliveira (2005) and felid species, L.wiedii (Schinz), L. pardalis (Linnaeus,), Oliveira-Santos et al. (2012) for the same region and and Puma concolor (Linnaeus), and small mammals species. However, the latter authors recorded variations (F = 2.601; P = 0.025) (Figure 3). in activity distribution of oncillas associated with the The multiple logistic regression between melanistic presence of other felid species. and spotted frequencies with regard to moon brightness Individuals from a single population may differ in several and nebulosity confirmed the same relationship above, aspects in the use of resources, for instance, between sexes being significant (X = 7.71; df = 2; p = 0.021); however, (Schoener, 1967), age-group (Cordero and Nicolas, 1987) only brightness contributed to explain melanistic and 2 and among trophic polymorphic individuals (Swanson et al., spotted frequencies in simple analysis (X = 7.35; df = 1; 2003). The temporal segregation between different colour coat phenotypes observed in our study corroborates Forsman´s prediction of the ecological significance of polymorphic colour patterns (Forsman et al., 2008). In the partitioning of resources between and within species, food and habitat sources are considered more important than the daily distribution of activity (Schoener, 1974). On the other hand, the activity pattern through interference competition increases in importance (Carothers and Jaksic, 1984), because territorial Neotropical carnivores could present potentially lethal damage between each other (Palomares and Caro, 1999; Di Bitetti et al., 2009). Melanistic individuals could be more effectively cryptic on bright nights than spotted individuals. According to our data, the space and the food resources available during bright nights are under-explored by other nocturnal predators present in the study area (e.g. L. wiedii, L. pardalis and Puma concolor). Most rodents, the main prey consumed by oncillas and other felid competitors (Oliveira et al., 2010), could present moon phobia, and thus decrease in Figure 2. Circadian activity of oncillas in Atlantic Rain availability to the predators during these moon phases Forest of southern Brazil. White bars represent registers (Lockard and Owings, 1974; Kaufman and Kaufman, of spotted (n = 139), and black bars, registers of melanistic 1982; Price et al., 1984). These ideas lead us to two non- oncillas (n = 31). excluding hypotheses to explain the observed pattern: (i) the melanistic individuals are more cryptic for their prey, ambushing prey more easily than other competitors on bright nights, compensating for the decrease in prey activity; (ii) poor quality foraging on bright nights did not compensate the high exposure of spotted oncillas to predators, but dark coat colour could offset this risk. Irrespective of the process underlying the observed pattern of reduction in intraspecific competition, attributed to differential use of time, our results suggest that melanistic individuals could occupy an alternative and wider ecological niche in relation to spotted individuals. While urban moths and desert rodents use melanism for cryptic protection in their habitats (Majerus, 1998; Majerus and Mundy, 2003), the oncilla, in its multicolor tropical habitat, should benefit Figure 3. Activity of melanistic oncilla (n = 11), spotted from the variation in moonlight. These results help us oncilla (n = 67), margay (n = 15), ocelot (n = 35), cougar to shed some light onto the way natural selection acts to (n = 19) and small mammals (n = 132) regarding nocturnal maintain polymorphisms (Skúlason and Smith, 1995), brightless (mean ± 95% IC) in Atlantic Rain Forest of southern Brazil. in this case, melanistic forms in populations of oncillas. 144 144 Braz. J. 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