Plumage polymorphism in a newly colonized black sparrowhawk population: classification, temporal stability and inheritance patterns

Size: px
Start display at page:

Download "Plumage polymorphism in a newly colonized black sparrowhawk population: classification, temporal stability and inheritance patterns"

Transcription

1 bs_bs_bannerjournal of Zoology Plumage polymorphism in a newly colonized black sparrowhawk population: classification, temporal stability and inheritance patterns A. Amar, A. Koeslag & O. Curtis* Journal of Zoology. Print ISSN Percy FitzPatrick Institute of African Ornithology, DST/NRF Centre of Excellence, University of Cape Town, Rondebosch, South Africa Keywords polymorphism; raptors; inheritance; Mendelian; morphs; pedigree data. Correspondence Arjun Amar, Zoology, Percy FitzPatrick Institute for African Ornithology, University of Cape Town, Private Bag X3, Rondebosch, Cape Town 7701, South Africa. Tel: arjun.amar@uct.ac.za *Current address: Overberg Lowlands Conservation Trust, 3 de Kock St., Napier Editor: Andrew Kitchener Received 7 June 2012; revised 9 August 2012; accepted 14 August 2012 doi: /j x Abstract Persistent plumage polymorphism occurs in around 3.5% of bird species, although its occurrence is not distributed equally across bird families or genera. Raptors show a disproportionately high frequency of polymorphism, and among raptors it is particularly frequent among the Accipiter hawks. However, no systematic study of polymorphism in this genus exists. Using a long-term study of the black sparrowhawk (Accipiter melanoleucus), a widespread polymorphic African Accipiter, we first demonstrate that the species shows discrete polymorphism (cf. continuous polymorphism), occurring as either dark or light morph adults, and that morph type and plumage pattern are invariant with age. We then demonstrate that adult morph type follows a typical Mendelian inheritance pattern, suggesting a one-locus, two-allele system within which the allele coding for the light morph is dominant. This inheritance pattern provides further support for classifying polymorphism in this species as discrete. In most of the species range the dark morph is the rarer morph; however, in our study population where the species is a recent colonist, over 75% of birds were dark and this remained fairly constant over the 10 years of our study. This reversal in morph ratio may represent an adaptive response to different environmental conditions or could be a founder effect with colonizing individuals having been mostly dark morph birds simply by chance. The extreme differences in environment conditions (seasonality of rainfall) that occur across the species range in South Africa provide support for an adaptive explanation, but further work is needed to test this hypothesis. Introduction Plumage polymorphism, in which different plumage morphs occur within the same age and sex of a breeding population, occurs in around 3.5% of bird species (Roulin, 2004). Evolutionary ecologists have long been fascinated by this phenomenon because the occurrence of two morphs in the same population runs counter to the notion that selective pressure should favour the optimal form for an environment, and any lesser quality individuals should be quickly eliminated (Huxley, 1955). Various explanations have been postulated for the occurrence and maintenance of polymorphism in birds (Galeotti et al., 2003; Roulin, 2004), and some of the most established hypotheses include: (1) apostatic selection (Fowlie & Krüger, 2003); (2) disruptive selection (Mather, 1955); (3) allopatric evolution (Cooke, Rockwell & Lank, 1995); (4) sexual selection (O Donald, 1983). Underpinning all these theories is the notion that an individual s phenotype is heritable, intransient, and not influenced by environmental variation, and that the colour variant is under selective pressure. However, in many studies these factors often remain untested. Polymorphism is particularly common in raptorial species (Fowlie & Krüger, 2003; Galeotti et al., 2003). Plumage colour in polymorphic raptors can vary continuously or may show two or more discrete morphs, for example, the polymorphic Swainson s hawk (Buteo swainsoni) and common buzzard (Buteo buteo) show continuous polymorphism, although they are often classified as dark, light or intermediate for analyses (Krüger & Lindström, 2001; Briggs, Collopy & Woodbridge, 2011). By contrast, discrete polymorphism exists in ferruginous hawk (Buteo regalis Schmutz & Schmutz, 1981) and Eleonora s falcon (Falco eleonorae Gangoso et al., 2011) with either dark or light morph birds. The type of phenotypic plumage polymorphism is likely to be influenced by the mode of genetic inheritance. Many studies have shown that polymorphic phenotypes are genetically determined in birds and follow a Mendelian mode of segregation (Roulin, 2004). To date, the use of pedigree data to study the genetic pattern of inheritance of plumage morphs Journal of Zoology (2012) 2012 The Authors. Journal of Zoology 2012 The Zoological Society of London 1

2 Polymorphism in black sparrowhawks A. Amar, A. Koeslag and O. Curtis in raptors has been reported in the common buzzard (Krüger & Lindström, 2001), ferruginous hawk (Schmutz & Schmutz, 1981), Swainson s hawk (Briggs, Woodbridge & Collopy, 2010a), Eleonora s falcon (Gangoso et al., 2011) and gyrfalcon (Falco rusticolus Chang, Lejeune & Cheng, 2010). The first four studies suggest a simple one-locus, two-allele autosomal inheritance pattern. For the ferruginous hawk and Eleonora s falcon that show discrete polymorphism, dark alleles are dominant and dark morph birds are thus either homozygous (with the alleles designated DD, capital letters for dominant alleles) or heterozygous (Dl), and light birds are homozygous for the recessive light allele (ll). For common buzzard and Swainson s hawk, dark (d) and light (l) alleles show incomplete dominance and heterozygous (dl) individuals therefore display intermediate plumage between the two homozygous morphs [dark (dd) or light (ll)], and hence give rise to continuous polymorphism along the plumage spectrum. For gyrfalcons, which show a full spectrum from pure white to pure black and many variants in-between, a more complex inheritance pattern is suggested, with colour being controlled by two genes, one controlling pigment production and the other restricting pigment distribution in feathers, with alleles in one gene having dominance and alleles in the other gene being co-dominant (Chang et al., 2010). Although several studies support a genetic basis to plumage variation, relatively few have demonstrated the stability of an individual s morph as it ages (Lowther, 1961; Lank et al., 1995; Brommer, Ahola & Karstinen, 2005). Criticism has been made of some studies which assume that plumage morph patterns remain constant over the course of an individual s life (Roulin, 2004). However, in the only study to examine this question in raptors, Briggs, Woodbridge & Collopy (2010b) found that plumage morph and patterning was invariant over time in 18 Swainson s hawks that were photographed at least 2 years apart. Among raptors, polymorphism occurs frequently in the genus Accipiter, with 11 of the 46 species displaying different colour morphs (Ferguson-Lees & Christie, 2001). However, no empirical research into polymorphism has focused on any species from this genus. Polymorphic species in this genus often show a similar polymorphic adult plumage, with a standard type, for example, common morph (light breast and underwing coverts) and a rarer dark adult morph, which tends to be black on the breast and underwing coverts (Ferguson- Lees & Christie, 2001). The black sparrowhawk (Accipiter melanoleucus) is a widely distributed Accipiter species occurring throughout much of sub-saharan Africa (Ferguson-Lees & Christie, 2001) and is usually described as showing these two morphs, although with varying degrees of white under the chin. The adult dark morph of this species is usually classed as rare (Steyn, 1982; Kemp & Kemp, 1998; Ferguson-Lees & Christie, 2001; Hockey, Dean & Ryan, 2005), although there are no published details on morph frequencies across the species range. The juveniles also occur as two morphs (pale or rufous) although these do not apparently reflect their subsequent adult morphs (Ferguson-Lees & Christie, 2001). The species has recently expanded into South Africa s Western Cape (Sebele, 2012) where it has successfully colonized the Cape Peninsula. The first breeding attempt on the Peninsula was recorded in 1993 (Oettlé, 1994; Curtis, Hockey & Koeslag, 2007) and the current population is estimated to be at least 40 breeding pairs. In this paper, using a long-term study of the black sparrowhawk on the Cape Peninsula, we undertake the first detailed study of polymorphism in an Accipiter species. Using photographs to score plumage characteristics we: (1) describe the type of polymorphism present and establish whether polymorphism in this species is best quantified as discrete or continuous, and (2) determine whether an individual s plumage pattern is invariant over time. Then, using pedigree data from wild, colour-ringed birds with known parental morphs, we explore plumage inheritance patterns to test for a genetic basis to the trait, and whether this follows conventional Mendelian inheritance patterns. Lastly, we examine the morph ratio of this newly colonized population and explore whether (1) it differs between the sexes of breeding adults, and (2) it has changed over time. Methods We monitored the black sparrowhawk population on the Cape Peninsula between 2001 and The study area features a matrix of habitats including urban gardens, alien pine (Pinus spp.) and Eucalyptus (Eucalyptus spp.) plantations, and small pockets of indigenous Afromontane forest and Fynbos. Altitudes where the birds breed range from sea level to about 300 m, and the climate is temperate, with locally variable winter rainfall (Cowling, MacDonald & Simmons, 1996). Mean annual rainfall is c.1250 mm, with average minimum and maximum monthly temperatures of 12 and 21 C, respectively (South African Weather Service). Monitoring was conducted during the breeding season (March November; Sebele, 2012) each year. Nests were located by surveying suitable stands of trees during the breeding season, searching for calling sparrowhawks, prey remains, whitewash and nest structures. Territories were visited regularly (approximately monthly) throughout the season until breeding was detected and then breeding attempts were monitored until conclusion. Where possible, we identified the morphs (dark or light) and sex of both parents attending a nest, which was possible in around 90% of breeding attempts. The species is easy to sex, with males weighing around 40% less than the females (unpublished data, see also Ferguson- Lees & Christie, 2001). In this study, data were only used for pairs where we knew the morph of each pair member. We fitted unique colour-ring combinations to as many breeding adult birds and 2 3-week-old nestlings as possible. Adults were trapped on territories using a bal-chatri baited with live white pigeons (Columba livia Berger & Mueller, 1959). Some of the birds ringed as chicks entered the breeding population during the course of the study, and enabled us to determine their adult morphs, which they began to acquire after their first year. In total, 102 adults (50 males, 52 females) and 131 chicks (76 males, 55 females of which 33 were later resighted in adult plumage) were fitted with colour rings during the course of the study. 2 Journal of Zoology (2012) 2012 The Authors. Journal of Zoology 2012 The Zoological Society of London

3 A. Amar, A. Koeslag and O. Curtis Polymorphism in black sparrowhawks Resightings of individual birds occurred mainly at breeding territories, but also through occasional observation away from breeding territories, via reporting and photographs taken by the authors and by members of the public. Photographs were taken of birds in adult plumage after being caught for ringing, or by using a 300-mm telephoto lens when birds were perched near the nest. Only photographs that showed the chin, throat, breast and flanks (hereafter front) were used for scoring coloration. Two observers (A. A. s data were used in the final analysis for ease of analysis) scored the percentage of white plumage on the front of birds in each photo. This approach to scoring plumage visually has been used in other studies, both with the birds in the hand (Brommer et al., 2005) and using photographs (Briggs et al., 2010b). Neither observer had knowledge of the other s scores, nor of the identity of the photographed bird. To explore morph frequencies between sexes and years, we used the data from all breeding pairs whose morphs were identified in each year (n = 245 pair years, between 2001 and 2010). However, this led to some level of pseudo-replication, since some birds featured in multiple years. For this reason, we ran a separate analysis, analysing the morph ratio of all pairs in the first year of breeding only (i.e. excluding known or suspected pairs from subsequent years; n = 130 pairs). Pairs and individuals were relatively faithful to nesting territories (unpublished data), and previously established or new pairs were classified based either on the pair s colour rings (n = 43), or in situations where only one bird was colour ringed, the combination of its colour rings and its partner s morph and lack of colour rings (n = 48), or in cases where neither bird was colour ringed (n = 39) we used only morph and sex combinations that were previously present on that territory to determine whether they were new or previously established pairs. We tested whether there were any differences between the sexes in the percentage of white on the front using a general linear model, with arcsine square root transformed percentage data as the response variable and the sex (M or F) as the explanatory variable. Analyses of the two morph types were conducted separately. Changes in frequency of morphs over time were analysed using a generalized linear model, with a binomial distribution and a logit link function. Whether a bird was a dark morph or a white morph (1/0) was specified as the response variable and year (as a continuous variable) was then specified as the explanatory variable. These analyses were carried out for males and females separately. All analyses were carried out in SAS version 9.1 (SAS Inc., 2004). Means are presented 1 standard error. Results Polymorphism classification and individual variance over time We had 135 photographs of individual adult birds which showed their front adequately to score the percentage of white plumage. These photos came from 42 different territories in the study area. Scores from the two observers did not differ significantly (paired t-test t 1,134 =-1.18, P = 0.23) and Frequency % white plumage on front Figure 1 The frequency distribution of birds displaying different amounts of white plumage on their throat, chin, breast, and flanks, and whether they were classified as either dark (solid bars) or light (open bars) in the field. White plumage percentages were visually estimated from 135 photos. The data show there is a clear bimodal pattern and justify the division of birds into either dark or light morphs. there was close agreement between different observers in the scores allocated to individual birds (R 2 = 0.96, P < ). A histogram of the scores clearly showed a discrete bimodal distribution in plumage coloration (Fig. 1). Birds described as dark morphs or light morphs had on average 8% (range 0 33%) and 84% (68 92%) white plumage on their front, respectively. For a subset of these data, where the sex was known (n = 125), we explored, for each morph type, whether the percentage of white plumage on the front differed between the sexes. For light morphs, although sample size was small (n = 28), there appeared to be no difference in the percentage of white plumage between the sexes (males: %, females: %; c 2 1 = 0.54; P = 0.46). For dark morphs (n = 97), although not quite statistically significant, there was a tendency for males to have less white than females (males: %, females: %; c 2 1 = 2.90; P = 0.08). For six colour-ringed individuals (three males and three females; three dark morphs and three light morphs), we had photos taken at least 4 years apart (range 4 11 years). From the scores given to these birds, it was apparent that the morphs did not change over the years, with the small difference between years not showing any directional change over time and was most likely attributable to observer error (Table 1). Inheritance patterns of morphs We observed 33 birds (17 males, 16 females) in adult plumage that were colour ringed as chicks at nests where we also knew the morphs of both attending parents. These birds were produced in 18 different territories and from the parental colouring combinations they came from 18 different pairings. Thirteen dark x dark pairings produced 13 dark morph offspring (8 males, 5 females) and no light morphs, whereas from 19 dark x light pairings, 13 dark morph offspring (6 Journal of Zoology (2012) 2012 The Authors. Journal of Zoology 2012 The Zoological Society of London 3

4 Polymorphism in black sparrowhawks A. Amar, A. Koeslag and O. Curtis Table 1 Percentage of white plumage estimated from photographs of breeding black sparrowhawks, with photos spanning 4 or more years. Percentage of white on the front (chin, throat, breast and flanks) Sex Morph Mean ( SD) Male L 78.2 ( 2.3) Male L 84.4 ( 1.4) Male D 5.3 ( 1.2) Female L 88.8 ( 3.0) Female D 2.6 ( 0.6) Female D 4.6 ( 0.6) SD, standard deviation. Table 2 Inheritance patterns according to the morph of the male and female parents and the morph (dark/light) of the offspring for all offspring and for male and female offspring separately. Male parental morph Dark Light All offspring Female parental morph Dark 13/0 5/0 Light 8/6 1/0 Male offspring Female parental morph Dark 8/0 3/0 Light 3/3 NA Female offspring Female parental morph Dark 5/0 2/0 Light 5/3 1/0 Birds were ringed as chicks with unique colour-ring combination. Inheritance patterns appear to follow a simple one-locus, two-allele system whereby the light allele is dominant. Light birds are therefore heterozygous with the genotype LL, or Ld, and dark birds are homozygous and have the genotype dd. Percentage of dark/light morphs Year Figure 2 Percentage ( 1 standard error) of dark morph male (solid line) and female (dashed line) black sparrowhawks in the population on the Cape Peninsula between 2001 and 2010, from the 244 pairs for which the morphs of both birds were known. There were significantly more dark morph males than females (P < 0.001), but the frequency of dark morphs did not differ over time for either sex (P > 0.60). males, 7 female) and 6 light morphs (3 males, 3 females) were produced. We had only one record of a light x light pairing and this produced one dark offspring (Table 2). Inheritance was apparently autosomal rather than sex linked because, on several occasions, birds produced offspring of the same sex that differed from their own morph (Table 2). However, it was interesting to note that light morph males produced no light morph offspring (from six cases), whereas light females produced light offspring at c. the 1:1 ratio predicted, although admittedly sample size for this finding is small. From these data, inheritance of morph type appears to follow a simple Mendelian one-locus, two-allele system, whereby the light allele is dominant. Light birds are therefore homozygous or heterozygous (LL or Ld) and dark birds are invariably homozygous (dd). Seventy-six per cent of birds in our population were dark morphs (see later). Assuming these dark morphs are homozygous (dd based on the inheritance patterns observed in this investigation), we can calculate the expected frequencies of L and d alleles in our study population under Hardy Weinberg equilibrium conditions. We calculated an allelic frequency of 0.87 for the d allele and consequently 0.13 for the L allele. We therefore expect that a minimum of 22% of birds are heterozygous light morphs (Ld) and only around 2% of birds are homozygous light morphs (LL). Stated differently, of the white morphs the vast majority (92%) are heterozygous (Ld) while only 8% are homozygous (LL). Morph frequencies over time and between sexes Over the entire study period, 76% of birds were dark morph, with the frequency being significantly higher for males than females (males: 83 2%, females 68 3%; c 2 1 = 14.85; P = ; Fig. 2). Between 2001 and 2010, the frequency of the two morphs did not change, either for the population as a whole (c 2 1 = 0.16, P = 0.68), or for either sex (males: c 2 1 = 0.12, P = 0.72; females: c 2 1 = 0.06, P = 0.80; Fig. 2). Using data only from the first year pairs were found breeding (to avoid issues of pseudoreplication), the same patterns emerged. There was no change 4 Journal of Zoology (2012) 2012 The Authors. Journal of Zoology 2012 The Zoological Society of London

5 A. Amar, A. Koeslag and O. Curtis Polymorphism in black sparrowhawks over time detected for either sex (males: c 2 1 = 0.02, P = 0.87; females: c 2 1 = 0.70, P = 0.40) and the frequency of dark morphs was significantly higher for males than females (males: 85 3%, females 73 4%; c 2 1 = 5.41; P = 0.02). Discussion Our study showed that (1) black sparrowhawks in our study area display discrete polymorphism in plumage coloration; (2) morph type and plumage patterns of individual birds change little, if at all, with age; (3) the observed inheritance pattern suggests that morph type is under genetic control consistent with a single Mendelian locus at which the light allele is dominant; (4) the Cape Peninsula population shows a reversal of the morph frequencies generally encountered in the rest of this species range, with over 75% of birds being dark morphs, and with significantly more dark morph males than females. From our plumage scores, it was clear that although there was considerable variation in the percentage of white plumage, black sparrowhawks show two distinct morphs, revealed as two clear modes on the histogram of percentage white plumage (Fig. 1). White morphs usually had around 85 90% and never less than 70% white plumage on their front whereas dark morphs usually had between 0 10% and never more than 35% white frontal plumage. There was clear separation of these modes, with no birds of intermediate plumage. Although not statistically significant, the percentage of white on the front of dark morph birds tended to be greater for females than for males, potentially indicating that darker males may enjoy a selective advantage in this population. Similar sex differences have been observed for polymorphic barn owls (Tyto alba), with females on average more reddish brown with more prominent black spotting than males (Roulin, 1999). It is also interesting that there were significantly more light morph females in the population than males (see later). However, whether the frequency of white morphs in the population and the percentage of white on the front of dark morph birds are in any way linked through similar selective pressures is unknown. Observation of known colour-ringed birds over time (4 11 years) provided strong evidence that plumage coloration in this species is fixed and does not vary with age (within this study s time frame). The species average life expectancy is around 10 years (based on average annual adult survival rates of 89% unpublished data). Data from males and females of both plumage morphs suggested that the proportion of white plumage on their front varied little with age. Indeed, the variation between years is probably attributable to observer error. Closer examination of the photographs of known birds suggests that plumage variability in this species might be successfully used to identify individuals over time. For example, obvious features of the plumage, such as the location of dark or light streaks, also appear to remain constant over time. This work therefore adds to that of Briggs et al. (2010b) in suggesting that plumage patterns in polymorphic raptors persist over an individual s lifetime, and provide further support for other studies that identify individuals in highly polymorphic populations by plumage alone (e.g. Krüger & Lindström, 2001; Curtis et al., 2005). Furthermore, because it is unlikely that these individuals would have experienced the same environmental conditions in the different years of their lives, it does suggest that environmental variables, such as weather, or body condition is relatively unimportant in determining the plumage patterns in this species, as Briggs et al. (2010b) also concluded. This result, together with our other findings on inheritance patterns (see later), further strengthens the argument that these plumage patterns may well be under genetic control. Observations of birds in adult plumage which were colour ringed as chicks with known parental morphs revealed that morph inheritance showed a classic Mendelian pattern. Dark x dark crosses produced only dark offspring, dark x light crosses produced both dark and light morph offspring, and in one instance a light x light cross produced a dark offspring. Hartley (1976) also reports an instance of light x light black sparrowhawk pairing producing a dark morph bird. From these findings, it therefore appears that polymorphism in this species is under genetic control with a one-locus, two-allele inheritance pattern, and that the light allele is dominant. Thus, light birds can be either homozygous (LL) or heterozygous (dl), whereas dark morph birds are always homozygous (dd). In this study, we have assumed no extra-pair paternity, which although unlikely to be true, occurs at a relatively low rate among raptors (Mougeot, 2004). Our finding of no impossible offspring from any pairings, given our predictions, also suggests that extra-pair paternity was not particularly high or not high enough to disrupt these results. Several Accipiter spp. occurring across the globe show similar polymorphic patterns to the black sparrowhawk (i.e. with dark or light morphs Ferguson-Lees & Christie, 2001): it would be interesting to know whether the same inheritance patterns also operate in these species. We could not use our data on the morphs of offspring from known parents to test accurately whether the phenotypes of these offspring were as predicted from our assumed genetic inheritance patterns (i.e. one locus, two alleles, with the light allele being dominant), because the offspring s morph was only assigned once a bird was recruited into the adult population rather than as a chick. Thus, any morph-linked differences in dispersal or juvenile survival would produce biased results. However, if we do assume equal juvenile survival rates between morphs, and given our estimated distribution of genotypes [i.e. 92% of light birds being heterozygous (dl)], we would predict an approximate dark : light morph ratio of 1:1.17 from our mixed-pair crosses. However, from the 19 offspring recruited from mixed pairs, we found a ratio of 2.2:1 in favour of dark morphs. Thus, more than twice as many dark birds as expected were subsequently found breeding from these crosses. This in turn suggests that either survival of white morphs prior to recruitment is lower than that of dark morphs, or that light morphs disperse further away from our study areas, or that we have incorrectly identified the inheritance mechanism. Although inheritance in our study was apparently not sex linked, no light offspring were ever recruited from light fathers, and this result largely explains the lower than predicted occurrence of light birds recruited from mixed pairs; however, the sample Journal of Zoology (2012) 2012 The Authors. Journal of Zoology 2012 The Zoological Society of London 5

6 Polymorphism in black sparrowhawks A. Amar, A. Koeslag and O. Curtis size (n = 6) for this finding was small. To investigate this further will require the use of genetic markers to test the genotypes of the chicks produced. Many studies have now shown the importance of the melanocortin-1 receptor gene in determining similar plumage polymorphisms in other bird species (Mundy, 2005) including for some (Gangoso et al., 2011) but not all raptor species (Hull et al., 2010). In future studies, we hope that the use of molecular markers will allow us to determine the morph genotype of the young chicks and to examine whether juvenile and sub-adult survival differs between morphs. Our Hardy Weinberg estimates suggested that 92% of the light morphs in our population were heterozygous. However, while this statistic is useful, we acknowledge that the accuracy of Hardy Weinberg equation relies on a number of assumptions, such as no selection, no non-random genetic drift and no gene flow, which are unlikely to be strictly true in our study population. For the current study, we have focused on polymorphism of adult black sparrowhawks. However, juvenile black sparrowhawks are also polymorphic, displaying a pale morph and a rufous morph (Ferguson-Lees & Christie, 2001). Initially, juvenile morphs were thought to be sex linked; however, pale and rufous juveniles of both sexes have been found in the same nest (Ferguson-Lees & Christie, 2001). Juvenile morph is also apparently not directly linked to adult phenotype, because of two rufous juveniles that were followed into adulthood, one became a light morph and the other a dark morph adult (Ferguson-Lees & Christie, 2001). Rather, we hypothesize that juvenile morphs could be linked to the adult morph genotype in the following way: birds that carry the recessive dark allele would be rufous juveniles and those birds that are homozygous light morphs (LL) would be pale morphs. Unfortunately, we do not yet have the data to test this hypothesis (e.g. from enough chicks with known morphs). The higher frequency of dark morph males as compared with females was an interesting result. One possible explanation for this could be that dark morph males survive better than dark morph females, and thus more dark morph males are recruited into the population. Again, an alternative explanation could be that females disperse further than males and that immigrant females come from population with a lower frequency of dark morphs. However, of the six light birds recruited into the population, three were male and three were female. Alternatively, this difference could have come about through mate choice, for example, if males actively select light females or if females actively select dark males, such similar mate choice has been reported for other species (Knapton & Falls, 1983; O Donald, 1983), and therefore remains a potential explanation for this finding. Dark morph adults were the more common (> 75%) morph in our population, which contrasts with most other sources that suggest that dark morphs are the rarer morph for this species (Steyn, 1982; Kemp & Kemp, 1998; Ferguson-Lees & Christie, 2001; Hockey et al., 2005). We are unaware of any published data on the morph frequency from other populations. However, unpublished data from a population studied in eastern Mpumalanga (formerly known as the Transvaal), South Africa (for details, see Tarboton & Allan, 1984), supported the idea that dark morphs are usually the rare morph for this species, with only around 22% (n = 36) of birds being dark morphs (W. Tarboton, unpublished data). What might explain the almost complete reversal in the frequency of the different morphs in these two study populations? There are two possible explanations for this pattern. Firstly, that it is simply a founder effect, with the first colonizing birds arriving on the Cape Peninsula being dark morphs, purely by chance. Alternatively, that this is an adaptive response to different environmental conditions (e.g. the different rainfall seasonality of the two regions). Both populations breed over the winter months (Sebele, 2012): this period coincides with the dry season in the north and east of South Africa, but with the wet season in the south west of the country. Thus, individuals breeding in our study population are exposed to far higher rainfall levels during the breeding period. Recent reviews on the causes and functions of polymorphism (Galeotti et al., 2003; Roulin, 2004) have both suggested a strong link between light conditions and variations in plumage, with crypsis (background matching) likely to play a key role. Thus, it may be that dark morph birds in our study population are at a selective advantage because they benefit from improved hunting efficiency in the poorer light conditions that would be associated with higher rainfall. As with most Accipiters, males provide most of the food during the breeding season, feeding the female during the incubation and early nestling stage, and this could explain the different morph frequencies between sexes if there is greater selection pressure for males to be dark than females. Similar relationships between habitat background and colour-morph ratio have been identified for other bird species (e.g. Rohwer, 1990). Alternatively, pressure from parasites may be greater in these wet conditions which may favour dark morphs, as different raptor morphs can show different immune responses (Gangoso et al., 2011) and subsequent parasite loads (Chakarov, Boerner & Kruger, 2008). It is also possible that dark morph birds have a thermal advantage in these colder wet conditions or that darker birds are able to withstand the feather degrading bacteria which may be more abundant or virulent in such conditions (Burtt & Ichida, 2004). To test further whether the unique morph ratios found in this population are likely to be an adaptive trait, future research will focus on whether dark morphs have any selective advantage by comparing reproductive output and survival with light morph birds: the findings presented here lay the foundations for this future research. Acknowledgements We are very grateful to Sharon Yodaiken and Gerry Meihuizen for all their help in the field. We are also grateful to Jacqueline Bishop and Gareth Tate who helped improve the manuscript immensely. We thank Warwick Tarboton for providing unpublished information on the morphs of his study population. Thanks are also due to Lovelater Sebele, Fitzum Baldi and Rowan Martin for useful discussions and advice. We thank M. Boerner and an anonymous reviewer for their comments which greatly improved the manuscript. We are 6 Journal of Zoology (2012) 2012 The Authors. Journal of Zoology 2012 The Zoological Society of London

7 A. Amar, A. Koeslag and O. Curtis Polymorphism in black sparrowhawks also grateful to all the land owners who have granted us access to carry out fieldwork on their land, and we are particularly grateful to South African National Parks for access to the Table Mountain National Park. References Berger, D.D. & Mueller, H.C. (1959). The bal-chatri: a trap for the birds of prey. Bird-Banding 30, Briggs, C.W., Collopy, M.W. & Woodbridge, B. (2011). Plumage polymorphism and fitness in Swainson s hawks. J. Evol. Biol. 24, Briggs, C.W., Woodbridge, B. & Collopy, M.W. (2010a). Inheritance patterns of plumage morph in Swainson s hawks. J. Raptor Res. 44, Briggs, C.W., Woodbridge, B. & Collopy, M.W. (2010b). Temporal morph invariance of Swainson s hawks. J. Raptor Res. 44, Brommer, J.E., Ahola, K. & Karstinen, T. (2005). The colour of fitness: plumage coloration and lifetime reproductive success in the tawny owl. Proc. Roy. Soc. Lond. B 272, Burtt, E.H. & Ichida, J.M. (2004). Gloger s rule, featherdegrading bacteria, and colour variation among song sparrows. Condor 106, Chakarov, N., Boerner, M. & Kruger, O. (2008). Fitness in common buzzards at the cross-point of opposite melanin-parasite interactions. Functional Ecology 22, Chang, V.S., Lejeune, J. & Cheng, K.M. (2010). The pattern of inheritance of melanin-based plumage color variants in the gyrfalcon (Falco rusticolus). J. Raptor Res. 44, Cooke, F., Rockwell, R.F. & Lank, D.B. (1995). The snow geese of La Pérouse Bay, natural selection in the wild. Oxford: Oxford University Press. Cowling, R.M., MacDonald, I.A.W. & Simmons, M.T. (1996). The Cape Peninsula, South Africa: physiographical, biological and historical background to an extraordinary hot-spot of biodiversity. Biodivers. Conserv. 5, Curtis, O., Malan, G., Jenkins, A. & Myburgh, N. (2005). Multiple brooding in birds of prey: South African black sparrowhawks Accipiter melanoleucus extend the boundaries. Ibis 147, Curtis, O.E., Hockey, P.A.R. & Koeslag, A. (2007). Competition with Egyptian geese Alopochen aegyptiaca overrides environmental factors in determining productivity of black sparrowhawks Accipiter melanoleucus. Ibis 149, Ferguson-Lees, J. & Christie, D. (2001). Raptors of the world. London: Christopher Helm. Fowlie, M.K. & Krüger, O. (2003). The evolution of plumage polymorphism in birds of prey and owls: the apostatic selection hypothesis revisited. J. Evol. Biol. 16, Galeotti, P., Rubolini, D., Dunn, P.O. & Fasola, M. (2003). Colour polymorphism in birds: causes and functions. J. Evol. Biol. 16, Gangoso, L., Grande, J.M., Ducrest, A.L., Figuerola, J., Bortolotti, G.R., Andrés, J.A. & Roulin, A. (2011). MC1R-dependent, melanin-based colour polymorphism is associated with cell-mediated response in the Eleonora s falcon. J. Evol. Biol. 24, Hartley, R. (1976). Some notes on the plumages of the black sparrowhawk. Bokmakierie 28, Hockey, P.A.R., Dean, W.R.J. & Ryan, P.G. (2005). Roberts birds of southern Africa. 7th edn. Cape Town: Trustees of the John Voelcker Bird Book Fund. Hull, J.M., Mindell, D.P., Talbot, S.L., Kay, E.H., Hoekstra, H.E. & Ernest, H.B. (2010). Population structure and plumage polymorphism: the intraspecific evolutionary relationships of a polymorphic raptor, Buteo jamaicensis harlani. BMC Evol. Biol. 10, 224. Huxley, J.S. (1955). Morphism in birds. Acta 6th International Ornithological Congress, Basel, 1954, pp Kemp, A. & Kemp, M. (1998). Birds of prey of Africa and its islands. Cape Town: New Holland. Knapton, R.W. & Falls, J.B. (1983). Differences in parental contribution among pair types in the polymorphic whitethroated sparrow. Can. J. Zool. 61, Krüger, O. & Lindström, J. (2001). Lifetime reproductive success in common buzzard, Buteo buteo: from individual variation to population demography. Oikos 93, Lank, D.B., Smith, C.M., Hanotte, O., Burke, T. & Cooke, F. (1995). Genetic polymorphism for alternative mating behaviour in lekking male Ruff Philomachus pugnax. Nature 378, Lowther, J.K. (1961). Polymorphism in the white-throated sparrow, Zonotrichia albicollis (Gmelin). Can. J. Zool. 39, Mather, K. (1955). Polymorphism as an outcome of disruptive selection. Evolution 9, Mougeot, F.S. (2004). Breeding density, cuckoldry risk and copulation behaviour during the fertile period in raptors: a comparative analysis. Anim. Behav 67, Mundy, N.I. (2005). A window on the genetics of evolution: MC1R and plumage colouration in birds. Proc. Roy. Soc. Lond. B 272, O Donald, P. (1983). The arctic skua: a study of the ecology and evolution of a seabird. Cambridge: Cambridge University Press. Oettlé, E. (1994). Black sparrowhawk breeds on the Cape Peninsula. Promerops 212, 7 7. Rohwer, S. (1990). Foraging differences between white and dark morphs of the Pacific Reef Heron Egretta sacra. Ibis 132, Roulin, A. (1999). Nonrandom pairing by male barn owls Tyto alba with respect to a female plumage trait. Behav. Ecol. 10, Journal of Zoology (2012) 2012 The Authors. Journal of Zoology 2012 The Zoological Society of London 7

8 Polymorphism in black sparrowhawks A. Amar, A. Koeslag and O. Curtis Roulin, A. (2004). The evolution, maintenance and adaptive function of genetic colour polymorphism in birds. Biol. Rev. 79, SAS Institute Inc. (2004). SAS/STAT 9.1 User s Guide. North Carolina: SAS Institute Inc. Schmutz, S.M. & Schmutz, J.K. (1981). Inheritance of color phases of ferruginous hawks. Condor 83, Sebele, L. (2012). Factors influencing the timing of breeding in a range expanding raptor at two spatial scales. Unpublished MSc thesis, University of Cape Town. Steyn, P. (1982). Birds of prey of southern Africa: their identification and life histories. Cape Town: David Philip. Tarboton, W.R. & Allan, D.G. (1984). The status of birds of prey in the Transvaal. Pretoria: Transvaal Museum. 8 Journal of Zoology (2012) 2012 The Authors. Journal of Zoology 2012 The Zoological Society of London

A record of a first year dark plumage Augur Buzzard moulting into normal plumage.

A record of a first year dark plumage Augur Buzzard moulting into normal plumage. A record of a first year dark plumage Augur Buzzard moulting into normal plumage. Simon Thomsett The Peregrine Fund, 5668 West Flying Hawk Lane, Boise Idaho, 83709, USA Also: Dept. of Ornithology, National

More information

Genetics and Probability

Genetics and Probability Genetics and Probability Genetics and Probability The likelihood that a particular event will occur is called probability. The principles of probability can be used to predict the outcomes of genetic crosses.

More information

9-2 Probability and Punnett. Squares Probability and Punnett Squares. Slide 1 of 21. Copyright Pearson Prentice Hall

9-2 Probability and Punnett. Squares Probability and Punnett Squares. Slide 1 of 21. Copyright Pearson Prentice Hall 9-2 Probability and Punnett 11-2 Probability and Punnett Squares Squares 1 of 21 11-2 Probability and Punnett Squares Genetics and Probability How do geneticists use the principles of probability? 2 of

More information

Evolution and Gene Frequencies: A Game of Survival and Reproductive Success

Evolution and Gene Frequencies: A Game of Survival and Reproductive Success Evolution and Gene Frequencies: A Game of Survival and Reproductive Success Introduction: In this population of Bengal tigers, alleles exist as either dominant or recessive. Bengal tigers live high in

More information

TE 408: Three-day Lesson Plan

TE 408: Three-day Lesson Plan TE 408: Three-day Lesson Plan Partner: Anthony Machniak School: Okemos High School Date: 3/17/2014 Name: Theodore Baker Mentor Teacher: Danielle Tandoc Class and grade level: 9-10th grade Biology Part

More information

Question 3 (30 points)

Question 3 (30 points) Question 3 (30 points) You hope to use your hard-won 7.014 knowledge to make some extra cash over the summer, so you adopt two Chinchillas to start a Chinchilla breeding business. Your Chinchillas are

More information

Afring News. An electronic journal published by SAFRING, Animal Demography Unit at the University of Cape Town

Afring News. An electronic journal published by SAFRING, Animal Demography Unit at the University of Cape Town Afring News An electronic journal published by SAFRING, Animal Demography Unit at the University of Cape Town Afring News accepts papers containing ringing information about birds. This includes interesting

More information

Inheritance of Livershunt in Irish Wolfhounds By Maura Lyons PhD

Inheritance of Livershunt in Irish Wolfhounds By Maura Lyons PhD Inheritance of Livershunt in Irish Wolfhounds By Maura Lyons PhD Glossary Gene = A piece of DNA that provides the 'recipe' for an enzyme or a protein. Gene locus = The position of a gene on a chromosome.

More information

New Mexico Avian Protection (NMAP) Feather Identification Guide

New Mexico Avian Protection (NMAP) Feather Identification Guide New Mexico Avian Protection (NMAP) Feather Identification Guide It is very common to find only feathers as remains beneath a power line due to predation, length of elapsed time since the mortality, weather,

More information

Biology 2108 Laboratory Exercises: Variation in Natural Systems. LABORATORY 2 Evolution: Genetic Variation within Species

Biology 2108 Laboratory Exercises: Variation in Natural Systems. LABORATORY 2 Evolution: Genetic Variation within Species Biology 2108 Laboratory Exercises: Variation in Natural Systems Ed Bostick Don Davis Marcus C. Davis Joe Dirnberger Bill Ensign Ben Golden Lynelle Golden Paula Jackson Ron Matson R.C. Paul Pam Rhyne Gail

More information

Blue is the New Black How genes can influence appearance.

Blue is the New Black How genes can influence appearance. Blue is the New Black How genes can influence appearance. Backstory Humans have selectively bred plants and animals for thousands of years in order to create variations most useful to our purposes. This

More information

BEYOND MENDEL. Incomplete Dominance: Blue (BB) Red (RR) F 1 hybrids have appearance in between 2 parents Purple (BR)

BEYOND MENDEL. Incomplete Dominance: Blue (BB) Red (RR) F 1 hybrids have appearance in between 2 parents Purple (BR) AP BIOLOGY EVOLUTION/HEREDITY UNIT Unit 1 Part 4 Chapter 14 Activity #5 NAME DATE PERIOD BEYOND MENDEL INCOMPLETE DOMINANCE Incomplete Dominance: Blue (BB) Red (RR) F 1 hybrids have appearance in between

More information

Page 2. Explain what is meant by codominant alleles (1) Male cats with a tortoiseshell phenotype do not usually occur. Explain why. ...

Page 2. Explain what is meant by codominant alleles (1) Male cats with a tortoiseshell phenotype do not usually occur. Explain why. ... Q1.In cats, males are XY and females are XX. A gene on the X chromosome controls fur colour in cats. The allele G codes for ginger fur and the allele B codes for black fur. These alleles are codominant.

More information

Worksheet for Morgan/Carter Laboratory #9 Mendelian Genetics II: Drosophila

Worksheet for Morgan/Carter Laboratory #9 Mendelian Genetics II: Drosophila Worksheet for Morgan/Carter Laboratory #9 Mendelian Genetics II: Drosophila Ex. 9-1: ESTABLISHING THE ENZYME REACTION CONTROLS Propose a hypothesis about AO activity in flies from vial 1a and flies from

More information

No tail (Manx) is a dominant trait and its allele is represented by M The presence of a tail is recessive and its allele is represented by m

No tail (Manx) is a dominant trait and its allele is represented by M The presence of a tail is recessive and its allele is represented by m Lab #4: Extensions to Mendelian Genetics Exercise #1 In this exercise you will be working with the Manx phenotype. This phenotype involves the presence or absence of a tail. The Manx phenotype is controlled

More information

UNIT 6 Genes and Inheritance sciencepeek.com

UNIT 6 Genes and Inheritance sciencepeek.com Part 1 - Inheritance of Genes Name Date Period 1. Fill in the charts below on the inheritance of genes. 2. In a diploid cell, there are copies of each chromosome present. 3. Each human diploid cell has

More information

Cow Exercise 1 Answer Key

Cow Exercise 1 Answer Key Name Cow Exercise 1 Key Goal In this exercise, you will use StarGenetics, a software tool that simulates mating experiments, to analyze the nature and mode of inheritance of specific genetic traits. Learning

More information

Incomplete Dominance, Co-Dominance, and Sex-linked dominance NON-MENDELIAN GENETICS

Incomplete Dominance, Co-Dominance, and Sex-linked dominance NON-MENDELIAN GENETICS Incomplete Dominance, Co-Dominance, and Sex-linked dominance NON-MENDELIAN GENETICS INCOMPLETE DOMINANCE INCOMPLETE DOMINANCE Two alleles dominant and recessive Genotypes are the same as simple Mendelian

More information

Here are some ground rules that you should ALWAYS follow when tackling an Inheritance Problem:

Here are some ground rules that you should ALWAYS follow when tackling an Inheritance Problem: E p is od e T h r e e : N o n - M ed ellian Inheritance Here are some ground rules that you should ALWAYS follow when tackling an Inheritance Problem: 1. Define the Alleles in question - you must state

More information

SBI3U: Exploring Modes of Inheritance. Purpose

SBI3U: Exploring Modes of Inheritance. Purpose SBI3U: Exploring Modes of Inheritance Assigned: Purpose Name: Due: To master understanding of various modes of inheritance by creating original creatures with various traits that are passed on by each

More information

HEREDITY BEYOND MENDEL INCOMPLETE DOMINANCE CODOMINANCE: Heredity Activity #3 page 1

HEREDITY BEYOND MENDEL INCOMPLETE DOMINANCE CODOMINANCE: Heredity Activity #3 page 1 AP BIOLOGY HEREDITY ACTIVITY #3 NAME DATE HOUR BEYOND MENDEL INCOMPLETE DOMINANCE CODOMINANCE: Heredity Activity #3 page 1 ABO BLOOD GROUPS Blood Type A B AB O Genotype RBC Antigen Plasma Antibodies In

More information

The purpose of this lab was to examine inheritance patters in cats through a

The purpose of this lab was to examine inheritance patters in cats through a Abstract The purpose of this lab was to examine inheritance patters in cats through a computer program called Catlab. Two specific questions were asked. What is the inheritance mechanism for a black verses

More information

1 - Black 2 Gold (Light) 3 - Gold. 4 - Gold (Rich Red) 5 - Black and Tan (Light gold) 6 - Black and Tan

1 - Black 2 Gold (Light) 3 - Gold. 4 - Gold (Rich Red) 5 - Black and Tan (Light gold) 6 - Black and Tan 1 - Black 2 Gold (Light) 3 - Gold 4 - Gold (Rich Red) 5 - Black and Tan (Light gold) 6 - Black and Tan 7 - Black and Tan (Rich Red) 8 - Blue/Grey 9 - Blue/Grey and Tan 10 - Chocolate/Brown 11 - Chocolate/Brown

More information

Beyond Mendel. Extending Mendelian Genetics. Incomplete Dominance. Think about this. Beyond Mendel. Chapter 12

Beyond Mendel. Extending Mendelian Genetics. Incomplete Dominance. Think about this. Beyond Mendel. Chapter 12 Beyond Mendel Extending Mendelian Genetics Chapter 12 Mendel s work did, however, provide a basis for discovering the passing of traits in other ways including: Incomplete Dominance Codominance Polygenic

More information

Part One: Introduction to Pedigree teaches students how to use Pedigree tools to create and analyze pedigrees.

Part One: Introduction to Pedigree teaches students how to use Pedigree tools to create and analyze pedigrees. Genetics Monohybrid Teacher s Guide 1.0 Summary The Monohybrid activity is the fifth core activity to be completed after Mutations. This activity contains four sections and the suggested time to complete

More information

We are learning to analyze data to solve basic genetic problems

We are learning to analyze data to solve basic genetic problems Gene 3 We are learning to analyze data to solve basic genetic problems Success Criteria: I can - use Punnett squares to solve basic genetic problems involving monohybrid crosses, incomplete dominance,

More information

What is Genetics? Genetics is the scientific study of heredity

What is Genetics? Genetics is the scientific study of heredity What is Genetics? Genetics is the scientific study of heredity What is a Trait? A trait is a specific characteristic that varies from one individual to another. Examples: Brown hair, blue eyes, tall, curly

More information

Seed color is either. that Studies Heredity. = Any Characteristic that can be passed from parents to offspring

Seed color is either. that Studies Heredity. = Any Characteristic that can be passed from parents to offspring Class Notes Genetic Definitions Trait = Any Characteristic that can be passed from parents to offspring Heredity The passing of traits from parent to offspring - Blood Type - Color of our Hair - Round

More information

1 This question is about the evolution, genetics, behaviour and physiology of cats.

1 This question is about the evolution, genetics, behaviour and physiology of cats. 1 This question is about the evolution, genetics, behaviour and physiology of cats. Fig. 1.1 (on the insert) shows a Scottish wildcat, Felis sylvestris. Modern domestic cats evolved from a wild ancestor

More information

Monohybrid Cross Video Review

Monohybrid Cross Video Review Name: Period: Monohybrid Cross Video Review 1. What is the name of the little boxes used in order to predict offspring without having to breed? 2. Define Punnett Square: 3. Define a monohybrid cross: 4.

More information

Understandings, Applications and Skills (This is what you maybe assessed on)

Understandings, Applications and Skills (This is what you maybe assessed on) 3. Genetics 3.4 Inheritance Name: Understandings, Applications and Skills (This is what you maybe assessed on) Statement Guidance 3.4.U1 3.4.U2 3.4.U3 3.4.U4 3.4.U5 3.4.U6 3.4.U7 3.4.U8 3.4.U9 Mendel discovered

More information

Mendel s Laws: Their Application to Solving Genetics Problem

Mendel s Laws: Their Application to Solving Genetics Problem Solving Genetics Problems Page 1 Mendel s Laws: Their Application to Solving Genetics Problem Objectives This lab activity is designed to teach students how to solve classic genetics problems using Mendel

More information

DO NOT WRITE ON THIS TEST Unit 6 Assessment Genetics Objective 3.2.2

DO NOT WRITE ON THIS TEST Unit 6 Assessment Genetics Objective 3.2.2 DO NOT WRITE ON THIS TEST Unit 6 Assessment Objective 3.2.2 Vocabulary Matching + 1 point each 1. dominant 2. recessive 3. genotype 4. phenotype 5. heterozygous 6. homozygous 7. incomplete dominance 8.

More information

Biology 3201 Sex Linked Review Mr.Gillam Name:

Biology 3201 Sex Linked Review Mr.Gillam Name: Biology 3201 Sex Linked Review Mr.Gillam Name: A female has the chromosomes XX, while a male has the chromosomes XY. In sex-linked inheritance the genes are carried on the chromosome and as a rule and

More information

Name: Period: Student Exploration: Mouse Genetics (One Trait)

Name: Period: Student Exploration: Mouse Genetics (One Trait) Directions: 1) Go to Explorelearning.com; 2) Login using your assigned user name and password. USER NAME: 1C772 PASSWORD: RAIN515 3) Find the MOUSE GENETICS ONE TRAIT Gizmo and click Launch Gizmo Name:

More information

Basic Terminology and Eyeband Colors

Basic Terminology and Eyeband Colors Color Genetics of the Dwarf Hotot Amy Hinkle, M.S. Printed in the 2011 ADHRC Guidebook, Free to use and distribute WITH ATTRIBUTION: Hinkle, Amy. "Color Genetics of the Dwarf Hotot." American Dwarf Hotot

More information

Questions from last week. You have a mouse with red eyes and a mouse with blue eyes. How could you determine which is the dominant trait?

Questions from last week. You have a mouse with red eyes and a mouse with blue eyes. How could you determine which is the dominant trait? Questions from last week You have a mouse with red eyes and a mouse with blue eyes. How could you determine which is the dominant trait? Mouse Eyes Without knowing anything about the parents you ll need

More information

Biology 100. ALE #8. Mendelian Genetics and Inheritance Practice Problems

Biology 100. ALE #8. Mendelian Genetics and Inheritance Practice Problems Biology 100 Instructor: K. Marr Name Lab Section Group No. Quarter ALE #8. Mendelian Genetics and Inheritance Practice Problems Answer the following questions neatly and fully in the spaces provided. References:

More information

What would explain the clinical incidence of PSS being lower than the presumed percentage of carriers should be producing?

What would explain the clinical incidence of PSS being lower than the presumed percentage of carriers should be producing? Many of the data sources seem to have a HUGE margin of error (e.g., mean age of 7.26 +/- 3.3 years). Is that a bad thing? How does this impact drawing conclusions from this data? What would need to be

More information

Independent Practice: Red throated booby bird R = red throat r = white throat. 1. Cross RR with rr. 2. Cross Rr with RR.

Independent Practice: Red throated booby bird R = red throat r = white throat. 1. Cross RR with rr. 2. Cross Rr with RR. Using Punnett Squares (Use with the Weblink Baby Steps Through Punnett Squares. ) Guided Practice: T = tall t = short Independent Practice: Red throated booby bird R = red throat r = white throat 1. Cross

More information

River Private Nature Reserve, Limpopo Province, South Africa. Hatfield 0028, Pretoria, South Africa. *Corresponding author,

River Private Nature Reserve, Limpopo Province, South Africa. Hatfield 0028, Pretoria, South Africa. *Corresponding author, Hooded Vulture (Necrosyrtes monachus) and African White-backed (Gyps africanus) nesting at the Olifants River Private Nature Reserve, Limpopo Province, South Africa Ara Monadjem 1,2*, Kerri Wolter 3, and

More information

Studying Gene Frequencies in a Population of Domestic Cats

Studying Gene Frequencies in a Population of Domestic Cats Studying Gene Frequencies in a Population of Domestic Cats Linda K. Ellis Department of Biology Monmouth University Edison Hall, 400 Cedar Avenue, W. Long Branch, NJ 07764 USA lellis@monmouth.edu Description:

More information

7. Describe the following with words and give an example: Heterozygous, homozygous recessive, homozygous dominant

7. Describe the following with words and give an example: Heterozygous, homozygous recessive, homozygous dominant Name: Genetics UNIT EXAM Review Below are review questions for each of the 5 learning goals we have addressed during this unit. This is the majority of the science content we covered. However, as a disclaimer

More information

Today: Mendel s Technique: What Mendel Observes: Mendelian Genetics: Consider this. Mendelian Genetics and Problems (In-Class 6)

Today: Mendel s Technique: What Mendel Observes: Mendelian Genetics: Consider this. Mendelian Genetics and Problems (In-Class 6) Today: Mendelian Genetics and Problems (In-Class 6) Mendelian Genetics: Consider this. 8 million possible chromosome combinations in each egg, and each sperm = >70 trillion possibilities! How are we able

More information

Genetics & Punnett Square Notes

Genetics & Punnett Square Notes Genetics & Punnett Square Notes Essential Question What is Genetics and how are punnett squares used? History of Genetics Gregor Mendel Father of modern genetics Studied pea plants Found that plants that

More information

Name period date assigned date due date returned. The Genetics of Garden Peas

Name period date assigned date due date returned. The Genetics of Garden Peas Name period date assigned date due date returned ollow instructions 1-4. ross 1. Place the parents genotypes in the Punnett Square and fill in the offspring s genotypes. Parent 2 Parent 1 Genotype Results

More information

7.013 Spring 2005 Problem Set 2

7.013 Spring 2005 Problem Set 2 MIT Department of Biology 7.013: Introductory Biology - Spring 2005 Instructors: Professor Hazel Sive, Professor Tyler Jacks, Dr. Claudette Gardel NAME TA 7.013 Spring 2005 Problem Set 2 FRIDAY February

More information

Studying Mechanisms of Inheritance using Drosophila melanogaster

Studying Mechanisms of Inheritance using Drosophila melanogaster Revised Fall 2018 Studying Mechanisms of Inheritance using Drosophila melanogaster Learning Objectives: 1. Describe the life cycle of Drospohilia. 2. Explain why the fruit fly is an ideal model organism

More information

EVOLUTIONARY GENETICS (Genome 453) Midterm Exam Name KEY

EVOLUTIONARY GENETICS (Genome 453) Midterm Exam Name KEY PLEASE: Put your name on every page and SHOW YOUR WORK. Also, lots of space is provided, but you do not have to fill it all! Note that the details of these problems are fictional, for exam purposes only.

More information

RECESSIVE BUDGIES: A BEGINNERS INTRODUCTION TO RECESSIVES IN BUDGERIGARS.

RECESSIVE BUDGIES: A BEGINNERS INTRODUCTION TO RECESSIVES IN BUDGERIGARS. RECESSIVE BUDGIES: A BEGINNERS INTRODUCTION TO RECESSIVES IN BUDGERIGARS. Published on the AWEBSA webpage with the kind permission of the author: Robert Manvell. Please visit his page and view photos of

More information

Level 2 Biology, 2015

Level 2 Biology, 2015 91157 911570 2SUPERVISOR S Level 2 Biology, 2015 91157 Demonstrate understanding of genetic variation and change 9.30 a.m. Monday 16 November 2015 Credits: Four Achievement Achievement with Merit Achievement

More information

Student Exploration: Mouse Genetics (One Trait)

Student Exploration: Mouse Genetics (One Trait) Name: Date: Student Exploration: Mouse Genetics (One Trait) Vocabulary: allele, DNA, dominant allele, gene, genotype, heredity, heterozygous, homozygous, hybrid, inheritance, phenotype, Punnett square,

More information

6. The lifetime Darwinian fitness of one organism is greater than that of another organism if: A. it lives longer than the other B. it is able to outc

6. The lifetime Darwinian fitness of one organism is greater than that of another organism if: A. it lives longer than the other B. it is able to outc 1. The money in the kingdom of Florin consists of bills with the value written on the front, and pictures of members of the royal family on the back. To test the hypothesis that all of the Florinese $5

More information

Genetics Problems. Character Dominant Recessive

Genetics Problems. Character Dominant Recessive Genetics Problems 1. A rooster with gray feathers is mated with a hen of the same phenotype. Among their offspring, 15 chicks are gray, 6 are black, and 8 are white. What is the simplest explanation for

More information

Mendelian Genetics Problem Set

Mendelian Genetics Problem Set Mendelian Genetics Problem Set Name: Biology 105 Principles of Biology Fall 2003 These problem sets are due at the beginning of your lab class the week of 11/10/03 Before beginning the assigned problem

More information

Bio homework #5. Biology Homework #5

Bio homework #5. Biology Homework #5 Biology Homework #5 Bio homework #5 The information presented during the first five weeks of INS is very important and will be useful to know in the future (next quarter and beyond).the purpose of this

More information

Page 1 of 7. Name: A. Preliminary Assessment #3. You may need a calculator for numbers 2&3.

Page 1 of 7. Name: A. Preliminary Assessment #3. You may need a calculator for numbers 2&3. Page 1 of 7 Name: 03-121-A Preliminary Assessment #3 You may need a calculator for numbers 2&3. You may bring one 3 inch by 5 inch card or paper with anything handwritten on it (front and back). You have

More information

17 Inherited change Exam-style questions. AQA Biology

17 Inherited change Exam-style questions. AQA Biology 1 Two genes in a mouse interact to control three possible coat colours: grey, black and brown. The two genes are located on separate chromosomes. Each gene has two alleles: A is dominant to a and B is

More information

Mendelian Genetics SI

Mendelian Genetics SI Name Mendelian Genetics SI Date 1. In sheep, eye color is controlled by a single gene with two alleles. When a homozygous brown-eyed sheep is crossed with a homozygous green-eyed sheep, blue-eyed offspring

More information

Great Horned Owl (Bubo virginianus) Productivity and Home Range Characteristics in a Shortgrass Prairie. Rosemary A. Frank and R.

Great Horned Owl (Bubo virginianus) Productivity and Home Range Characteristics in a Shortgrass Prairie. Rosemary A. Frank and R. Great Horned Owl (Bubo virginianus) Productivity and Home Range Characteristics in a Shortgrass Prairie Rosemary A. Frank and R. Scott Lutz 1 Abstract. We studied movements and breeding success of resident

More information

GENETIC DRIFT Carol Beuchat PhD ( 2013)

GENETIC DRIFT Carol Beuchat PhD ( 2013) GENETIC DRIFT Carol Beuchat PhD ( 2013) By now you should be very comfortable with the notion that for every gene location - a locus - an animal has two alleles, one that came from the sire and one from

More information

Bell Ringer. Which features do you have that match your mother? Your father? Which of the following features do you have?

Bell Ringer. Which features do you have that match your mother? Your father? Which of the following features do you have? Bell Ringer Which features do you have that match your mother? Your father? Which of the following features do you have? Widow s Peak? Ability to roll your tongue? Attached earlobes? Simple Genetics Exploring

More information

Genetics. What s Genetics? An organism s heredity is the set of characteristics it receives from its parents.

Genetics. What s Genetics? An organism s heredity is the set of characteristics it receives from its parents. Genetics Why don t you look exactly like your parents? Pull How are traits passed to the next generation? Pull What s Genetics? An organism s heredity is the set of characteristics it receives from its

More information

Naked Bunny Evolution

Naked Bunny Evolution Naked Bunny Evolution In this activity, you will examine natural selection in a small population of wild rabbits. Evolution, on a genetic level, is a change in the frequency of alleles in a population

More information

Virtual Lab: Sex-Linked Traits Worksheet. 1. Please make sure you have read through all of the information in the

Virtual Lab: Sex-Linked Traits Worksheet. 1. Please make sure you have read through all of the information in the Virtual Lab: Sex-Linked Traits Worksheet 1. Please make sure you have read through all of the information in the Questions and Information areas. If you come upon terms that are unfamiliar to you, please

More information

Mendelian Inheritance Practice Problems

Mendelian Inheritance Practice Problems Name: Period: Mendelian Inheritance Practice Problems Team Problem 1 2. 3. Team Problem 2 2. Team Problem 3 Team Problem 4 Mendelian Inheritance Monohybrid Practice Problems In cats, long hair is recessive

More information

Determining the Inheritance Patterns of Purple Eye, Lobe Eye, and Yellow Body Traits of. Drosophilia Flies. Introduction

Determining the Inheritance Patterns of Purple Eye, Lobe Eye, and Yellow Body Traits of. Drosophilia Flies. Introduction Karen Jacques and Audrey Puleio Mrs. Lajoie Honors Biology April 30, 2012 Determining the Inheritance Patterns of Purple Eye, Lobe Eye, and Yellow Body Traits of Drosophilia Flies Introduction This experiment

More information

The color and patterning of pigmentation in cats, dogs, mice horses and other mammals results from the interaction of several different genes

The color and patterning of pigmentation in cats, dogs, mice horses and other mammals results from the interaction of several different genes The color and patterning of pigmentation in cats, dogs, mice horses and other mammals results from the interaction of several different genes 1 Gene Interactions: Specific alleles of one gene mask or modify

More information

BIOL4. General Certificate of Education Advanced Level Examination June Unit 4 Populations and environment. Monday 13 June pm to 3.

BIOL4. General Certificate of Education Advanced Level Examination June Unit 4 Populations and environment. Monday 13 June pm to 3. Centre Number Surname Candidate Number For Examiner s Use Other Names Candidate Signature Examiner s Initials General Certificate of Education Advanced Level Examination June 2011 Question 1 2 Mark Biology

More information

3. Complete the Punnett square for heterozygous yellow (yellow is dominant): What is the genotype: and what is the phenotype:

3. Complete the Punnett square for heterozygous yellow (yellow is dominant): What is the genotype: and what is the phenotype: Name: Period: Video Review: Two Factor Crosses & Independent Assortment: 1. Mendel discovered many things about the characteristics of pea plants including the qualities of the peas themselves. What two

More information

The Genetics of Color In Labradors

The Genetics of Color In Labradors By Amy Frost Dahl, Ph.D. Oak Hill Kennel First published in The Retriever Journal, June/July 1998 Seeing that two of the dogs I brought in for CERF exams were black Labs, the vet's assistant started telling

More information

Sample Size Adapted from Schmidt, et al Life All Around Us.

Sample Size Adapted from Schmidt, et al Life All Around Us. Lab 9, Biol-1, C. Briggs, revised Spring 2018 Sample Size Adapted from Schmidt, et al. 2006. Life All Around Us. Name: Lab day of week: Objectives Observe the benefits of large sample sizes. Instructions

More information

HEREDITARY STUDENT PACKET # 5

HEREDITARY STUDENT PACKET # 5 HEREDITARY STUDENT PACKET # 5 Name: Date: Big Idea 16: Heredity and Reproduction Benchmark: SC.7.L.16.1: Understand and explain that every organism requires a set of instructions that specifies its traits,

More information

Name: Block: Date: Packet #12 Unit 6: Heredity

Name: Block: Date: Packet #12 Unit 6: Heredity Name: Block: Date: Packet #12 Unit 6: Heredity Objectives: By the conclusion of this unit, you should be able to: Topic 1: Simple Heredity 1. Define and relate the following terms: self-fertilization,

More information

Patterns of Inheritance. What are the different ways traits can be inherited?

Patterns of Inheritance. What are the different ways traits can be inherited? Patterns of Inheritance What are the different ways traits can be inherited? Review: Patterns of Inheritance we know already 1. Autosomal dominant: If an individual is heterozygous, only one allele is

More information

Complex Patterns of Inheritance Puzzle Stations Station #1: Multiple alleles, blood types

Complex Patterns of Inheritance Puzzle Stations Station #1: Multiple alleles, blood types Station #1: Multiple alleles, blood types (Remember, the possible multiple alleles for blood are written as I A, I B, i, with types A and B being codominant, and O being recessive.) 1. A man with blood

More information

Biol 160: Lab 7. Modeling Evolution

Biol 160: Lab 7. Modeling Evolution Name: Modeling Evolution OBJECTIVES Help you develop an understanding of important factors that affect evolution of a species. Demonstrate important biological and environmental selection factors that

More information

Incomplete Dominance and Codominance

Incomplete Dominance and Codominance Incomplete Dominance and Codominance Name Define incomplete dominance Incomplete dominance can be remembered in the form of Red flower X white flower = pink flower The trick is to recognize when you are

More information

BioSci 110, Fall 08 Exam 2

BioSci 110, Fall 08 Exam 2 1. is the cell division process that results in the production of a. mitosis; 2 gametes b. meiosis; 2 gametes c. meiosis; 2 somatic (body) cells d. mitosis; 4 somatic (body) cells e. *meiosis; 4 gametes

More information

INHERITANCE OF BODY WEIGHT IN DOMESTIC FOWL. Single Comb White Leghorn breeds of fowl and in their hybrids.

INHERITANCE OF BODY WEIGHT IN DOMESTIC FOWL. Single Comb White Leghorn breeds of fowl and in their hybrids. 440 GENETICS: N. F. WATERS PROC. N. A. S. and genetical behavior of this form is not incompatible with the segmental interchange theory of circle formation in Oenothera. Summary.-It is impossible for the

More information

Inheritance of the king coat colour pattern in cheetahs Acinonyx jubatus

Inheritance of the king coat colour pattern in cheetahs Acinonyx jubatus 1. Zool., Lond. (A) (1986) 209, 573-578 Inheritance of the king coat colour pattern in cheetahs Acinonyx jubatus R. 1. VAN AARDE* Mammal Research Institute, University of Pretoria, Pretoria 0002, South

More information

Biology 164 Laboratory

Biology 164 Laboratory Biology 164 Laboratory CATLAB: Computer Model for Inheritance of Coat and Tail Characteristics in Domestic Cats (Based on simulation developed by Judith Kinnear, University of Sydney, NSW, Australia) Introduction

More information

Mendelian Genetics and Punnett Squares 5/07 Integrated Science 2 Redwood High School Name: Period:

Mendelian Genetics and Punnett Squares 5/07 Integrated Science 2 Redwood High School Name: Period: Mendelian Genetics and Punnett Squares 5/07 Integrated Science 2 Redwood High School Name: Period: Background Monohybrid crosses are crosses in which only one characteristic/trait is considered. For example,

More information

GENETICS PRACTICE 1: BASIC MENDELIAN GENETICS

GENETICS PRACTICE 1: BASIC MENDELIAN GENETICS Period Date GENETICS PRACTICE 1: BASIC MENDELIAN GENETICS Solve these genetics problems. Be sure to complete the Punnett square to show how you derived your solution. 1. In humans the allele for albinism

More information

Chapter 11-2 Probability and Punnett Squares Notes

Chapter 11-2 Probability and Punnett Squares Notes Chapter 11-2 Probability and Punnett Squares Notes Every time Mendel performed a cross with his pea plants, he carefully counted the offspring (over 20,000 plants) his why he noticed there was a pattern!

More information

Station 1. Using the cards, match the vocabulary word with its definition. If there are any words you do not know, write them down if you have time!

Station 1. Using the cards, match the vocabulary word with its definition. If there are any words you do not know, write them down if you have time! Station 1 Using the cards, match the vocabulary word with its definition. If there are any words you do not know, write them down if you have time! Station 2 Answer the following questions on a separate

More information

Unit 5 Guided Notes Genetics

Unit 5 Guided Notes Genetics Gregor Mendel Modern genetics began in the mid-1800s in an abbey garden, where a monk named documented inheritance in peas Medel s Work What is inheritance: used good experimental design used analysis

More information

Co-operative breeding by Long-tailed Tits

Co-operative breeding by Long-tailed Tits Co-operative breeding by Long-tailed Tits v N. W. Glen and C. M. Perrins For most of this century, ornithologists have tended to believe that the majority of birds breed monogamously, with either the pair

More information

Was the Spotted Horse an Imaginary Creature? g.org/sciencenow/2011/11/was-the-spotted-horse-an-imagina.html

Was the Spotted Horse an Imaginary Creature?   g.org/sciencenow/2011/11/was-the-spotted-horse-an-imagina.html Was the Spotted Horse an Imaginary Creature? http://news.sciencema g.org/sciencenow/2011/11/was-the-spotted-horse-an-imagina.html 1 Genotypes of predomestic horses match phenotypes painted in Paleolithic

More information

Genetics Intervention

Genetics Intervention Genetics Intervention Vocabulary: Define the following terms on a separate piece of paper. allele autosome chromosome codominance dihybrid diploid dominant gene gamete haploid heterozygous homozygous incomplete

More information

Different versions of a single gene are called allleles, and one can be dominant over the other(s).

Different versions of a single gene are called allleles, and one can be dominant over the other(s). Answer KEY 1 Different versions of a single gene are called allleles, and one can be dominant over the other(s). 2 Describe genotype and phenotype in your own words. A genotype is the genetic makeup of

More information

Science 10-Biology Activity 17 Worksheet on More Complex Genetics

Science 10-Biology Activity 17 Worksheet on More Complex Genetics Science 10-Biology Activity 17 Worksheet on More Complex Genetics 10 Name Due Date Show Me Hand In Correct and Hand In Again By NOTE: This worksheet is based on material from pages 398-404 in Science Probe.

More information

Level 2 Biology, 2017

Level 2 Biology, 2017 91157 911570 2SUPERVISOR S Level 2 Biology, 2017 91157 Demonstrate understanding of genetic variation and change 2.00 p.m. Wednesday 22 November 2017 Credits: Four Achievement Achievement with Merit Achievement

More information

Problem 1. What is the simplest explanation for the inheritance of these colors in chickens?

Problem 1. What is the simplest explanation for the inheritance of these colors in chickens? Problem 1 A rooster with gray feathers is mated with a hen of the same phenotype. Among their offspring, 15 chicks are gray, 6 are black, and 8 are white. What is the simplest explanation for the inheritance

More information

Problem 1. What is the simplest explanation for the inheritance of these colors in chickens?

Problem 1. What is the simplest explanation for the inheritance of these colors in chickens? Problem 1 A rooster with gray feathers is mated with a hen of the same phenotype. Among their offspring, 15 chicks are gray, 6 are black, and 8 are white. What is the simplest explanation for the inheritance

More information

The Pigeon Genetics Newsletter

The Pigeon Genetics Newsletter The Pigeon Genetics Newsletter News, Views, and Comments. Editor: R J Rodgers, Nova Scotia, Canada Co-Editor: Jith Peter, Palakkad, India March 2016, Volume 4, page 1 Section # (1) Beginner Text &Photos:

More information

Population dynamics of small game. Pekka Helle Natural Resources Institute Finland Luke Oulu

Population dynamics of small game. Pekka Helle Natural Resources Institute Finland Luke Oulu Population dynamics of small game Pekka Helle Natural Resources Institute Finland Luke Oulu Populations tend to vary in size temporally, some species show more variation than others Depends on degree of

More information

Immature Plumages of the Eastern Imperial Eagle Aquila heliaca

Immature Plumages of the Eastern Imperial Eagle Aquila heliaca Chancellor, R. D. & B.-U. Meyburg eds. 2004 Raptors Worldwide WWGBP/MME Immature Plumages of the Eastern Imperial Eagle Aquila heliaca William S. Clark ABSTRACT The Eastern Imperial Eagles, Aquila heliaca,

More information

Biology 201 (Genetics) Exam #1 120 points 22 September 2006

Biology 201 (Genetics) Exam #1 120 points 22 September 2006 Name KEY Section Biology 201 (Genetics) Exam #1 120 points 22 September 2006 Read the question carefully before answering. Think before you write. You will have up to 50 minutes to take this exam. After

More information

Lab 7. Evolution Lab. Name: General Introduction:

Lab 7. Evolution Lab. Name: General Introduction: Lab 7 Name: Evolution Lab OBJECTIVES: Help you develop an understanding of important factors that affect evolution of a species. Demonstrate important biological and environmental selection factors that

More information