Aberrant Colouration in the Atlantic Puffin ( Fratercula arctica ), the Common Murre ( Uria aalge ), and the Thick-billed Murre ( U. lomvia ) from Atlantic Canada

ALEXANDER L. B OND 1, 2, 4 and ANTONY W. D IAMOND 3

1Department of Biology, University of Saskatchewan, and Environment Canada, 11 Innovation Boulevard, Saskatoon, Saskat - chewan S7N 3H5 Canada 2Present address: RSPB Centre for Conservation Science, Royal Society for the Protection of Birds, The Lodge, Sandy, Bedfordshire SG19 2DL United Kingdom 3Atlantic Laboratory for Avian Research, University of New Brunswick, PO Box 4400, Fredericton, New Brunswick E3B 5A3 Canada 4Corresponding author: [email protected]

Bond, Alexander L., and Antony W. Diamond. 2016. Aberrant colouration in the Atlantic Puffin ( Fratercula arctica ), the Common Murre ( Uria aalge ), and the Thick-billed Murre ( U. lomvia ) from Atlantic Canada. Canadian Field-Natu - ralist 130(2): 140–145. The colour of birds’ plumage and bare parts is an important feature in choice of mate, camouflage, thermoregulation, species recognition, and flight mechanics. Abnormalities in colouration occur in a variety of species and can have important consequences for an individual’s survival and fitness. We present 7 new cases of colouration abnormalities in 3 species of auk (Alcidae) and review previous reports to correctly assign the proper form of abnormality to specimens in museums or photographs. Of the 53 reported colouration abnormalities, we reclassified 42, progressive greying being the most common (18 or 19 cases out of 42 , 43–45%), followed by brown (10/42 cases, 24%), in addition to 6 cases of melanism, 4 of dilution, 2 of partial leucism, and 1 likely somatic mutation. Properly describing the form of colour abnormality improves our understanding of the frequency, causes, and consequences of aberrant colouration. Key Words: Atlantic Puffin; Fratercula arctica ; Common Murre; Uria aalge ; Thick-billed Murre; Uria lomvia ; plumage; colouration; colour aberration; Atlantic Canada Introduction dark patches, to an overall dilution of pigment (van The colour of birds’ plumage and bare parts (bill and Grouw et al. 2011; Jakubas and Wojczulanis-Jakubas legs) plays many important functional roles in flight, 2012; van Grouw 2013). Cases of extreme melanism, thermoregulation, mate choice, camouflage from pred - where the bird appears entirely black, are rare (Clarke ators, and species recognition (Hamilton and Heppner 1913; Loomis 1918). 1967; Barrowclough and Sibley 1980; Cairns 1986; There has been a fair degree of interest and a long Jones 1990; Andersson et al. 1998; Murphy and Pham history of documenting plumage aberrations in seabirds 2012). However, a variety of colouration abnormalities (e.g., Newton 1877), including the Procellariiformes of feathers and bare parts occur, albeit at very low fre - (Mancini et al. 2010), and Sphenisciformes (Forrest quencies (Sage 1962, 1963; Ross 1964; Gross 1965; and Naveen 2000). In the auks (Charadriiformes, fami - Forrest and Naveen 2000; McCardle 2012). There is a ly Alcidae), plumage abnormalities have been described long history of documenting abnormal colouration in for many species (Deane 1876; Newton 1877; Arnold a variety of birds (Krüper 1857; Deane 1876; Newton 1950; Lockley 1953; Tuck 1961; Sealy 1969; Reinsch 1877), but only recently has the physiological basis 1983; Leopold et al. 2010; van Grouw et al. 2011; for the varying conditions been understood (Summers Jakubas and Wojczulanis-Jakubas 2012). Here, we 2009). Historically, many individuals with any sort of describe five additional cases of plumage abnormali - lack of plumage colouration were termed “albino,” ties in 3 species of Atlantic alcid: Thick-billed Murre des pite this term having a very specific meaning and (Uria lomvia ), Common Murre ( U. aalge ), and Atlantic the condition being very rare in wild populations (van Puffin ( Fratercula arctica ), and we use current termi - Grouw 2010, 2013). nology to clarify the particular mutation in several mu - The most extreme lack of colouration, albinism, is a seum specimens described previously. complete lack of eumelanin and phaeomelanin and is extremely rare (McCardle 2012). More common are Methods cases of partial under-expression of pigments (leucism, We used the system described by van Grouw (2013) progressive greying, brown, dilution, and ino), and these for assigning colour aberrations in 7 categories (Table can exhibit a wide range of phenotypes from light or 1). We made regular notes on abnormal plumages dur -

140 ©The Ottawa Field-Naturalists’ Club (2016) 2016 BOND AND DIAMOND : A BERRANT COLOURATION IN AUKS 141

TABLE 1. The physiological and genetic basis for colour aberrations and their phenotypic effect. Colour aberration Physiological or genetic basis Phenotypic effect Albinism Complete absence of eumelanin and phaeomelanin All white feathers, red eyes, pink feet in all tissues, caused by inherited lack of tyrosinase and bill Brown Reduction in the expression of eumelanin molecules Typically black tissues become caused by incomplete oxidation brown, and rufous/yellow/brown tissues are unaffected Dilution – isabel Reduced number of eumelanin molecules (but not Typically black tissues become their expression) silver/grey, and rufous/yellow/brown tissues are unaffected Dilution – pastel Reduced number of eumelanin and phaeomelanin Typically black tissues become molecules (but not their expression) silver/grey, and rufous/yellow/brown tissues become buff/cream Ino – dark Strong reduction in the expression of both eumelanin Typically black tissues are light and phaeomelanin caused by incomplete oxidation brown, and rufous/yellow/brown tissues are buff/cream; bill and feet pink Ino – light Mild reduction in the expression of both eumelanin Typically black tissues are very pale and phaeomelanin caused by incomplete oxidation brown/cream, and rufous/yellow/brown tones are hardly present; eyes, bill, and feet pink Leucism Partial or total lack of eumelanin and phaeomelanin Plumage all white or all-white in feathers/skin caused by inherited lack of feathers mixed with normally pigment cells coloured feathers. Bill and feet can be pink or unaffected; normally coloured eyes Melanism Increased melanin deposits Increase in black or darker red/brown pigments Progressive greying Partial or total loss of eumelanin and phaeomelanin Plumage all white or all-white in feathers/skin caused by gradual loss of pigment feathers mixed with normally cells with age coloured feathers. Bill and feet can be pink or unaffected; normally coloured eyes Source: Adapted from van Grouw (2013). ing our research in eastern Canada and, where possi - Thick-billed Murre, CMNAV 38151 ble, documented cases of abnormal plumage with pho - This is a melanistic bird described by Tuck (1961) as tographs. We also searched published and unpublished being shot in Trinity Bay, Newfoundland and Labrador, articles, including government reports, monographs, on 31 January 1952 (Figure 2); its destination was list - and journal articles for mention of plumage or coloura - ed only as the “National Museum of Canada” (now the tion abnormalities in our 3 species of interest. Finally, Canadian Museum of Nature). It is uniformly dark we contacted curators at several museums known to brown/black above, and dark grey below, rather than house specimens with abnormal colouration (see Ack - white. nowledgements) and obtained photographs of the spec - Common Murre, Cabot Island, Newfoundland and imens in question to allow further examination and re - Labrador classification where possible. On 2 August 2011, 1 bird among 100 adult breeding Common Murres banded on Cabot Island (49°17'N, Results 53°36'W) lacked pigment in parts of its bill, and in its Thick-billed Murre, CMNAV 37719 feet (a form of progressive greying; Figure 3). On the This is a previously undescribed case of dilution in Isle of May, Scotland, this occurs in about 1 in 1000 a bird collected by L. M. Tuck in Hants Harbour, New - Common Murres (M. P. Harris, personal communica - foundland and Labrador, on 12 March 1951 (Figure 1), tion). A single Thick-billed Murre with yellow feet was now in the Canadian Museum of Nature (CMNAV). seen on Coats Island in the 1980s (A. J. Gaston, per - The normally dark brown back, neck, head, and wings sonal communication). are a silvery grey. Tuck (1961) also described 3 “full Atlantic Puffin, Gull Island, Witless Bay, Newfound - albinos” and 3 “partial albinos.” This is almost cer - land and Labrador tainly incorrect, and the latter, whose black feathers In May and June 2012, 91 adult Atlantic Puffins were appeared grey were likely cases of dilution (van Grouw banded on Gull Island (47°15'N, 52°12'W). One bird’s 2013) and could include CMNAV 38151 (see below); black head feathers were grey, flecked with white (Fig - the true status of the 3 “albinos” is unknown (Table S1 ). ure 4). The bird was a breeding adult, with more than 142 THE CANADIAN FIELD -N ATURALIST Vol. 130

FIGURE 1. Thick-billed Murre (CMNAV 37719) showing dilution. Photo: M. Gosselin.

FIGURE 2. Melanistic Thick-billed Murre (CMNAV 38151). Photo: M. Gosselin.

FIGURE 3. Common Murre on Cabot Island, Newfoundland, showing progressive greying in the legs and bill. Photo: A. Lang. 2016 BOND AND DIAMOND : A BERRANT COLOURATION IN AUKS 143

FIGURE 4. Atlantic Puffin on Gull Island, Newfoundland, show - ing a likely somatic mutation, as other dark plumage ap peared normal. Photo: A. Bond.

2 bill grooves (Harris 1981), and all other aspects of plumage and colouration of bare parts were typical, including the black feathers on the back and proximal FIGURE 5. Atlantic Puffin on Machias Seal Island, New Bruns- region of the breast. Although this might appear to be wick, with progressive greying expressed as white a case of dilution, the genetic mechanism (a heritable feathers around the neck. Photo: T. Einfeldt. mutation affecting the number of pigment molecules, but not their colour) would result in all black plumage appearing grey. Therefore, this is likely the result of a Atlantic Puffin, Machias Seal Island, and Grand somatic mutation, rather than a heritable plumage Manan basin, New Brunswick abnormality. In 2002, 2004, and 2009, a “white puffin” was Atlantic Puffin, Machias Seal Island, New Brunswick observed at sea around Machias Seal Island or in the In May 2013, an adult Atlantic Puffin with more than Grand Manan basin (Figure 6). The bare parts were 2 bill grooves was photographed on Machias Seal Is - unaffected, as were portions of the feathers (e.g., tips land (44°3'N, 67°06'W). It had white feathers speckled of primary feathers), making this an extreme case of throughout the black contour feathers on its neck and progressive greying. Other “white puffins” have been back (Figure 5), showing characteristics of progressive recorded at the Isles of Scilly, United Kingdom, in greying. Colouration of all other aspects of plumage May 2009 (Harris and Wanless 2011) and at the Faroe and bare parts appeared normal. Islands in the late 1800s (Lockley 1953; see below).

FIGURE 6. A “white puffin” in the Grand Manan basin, New Brunswick, in 2009, showing an extreme case of progressive greying. Photo: New England Aquarium. 144 THE CANADIAN FIELD -N ATURALIST Vol. 130

These 3 records may or may not pertain to a single increase the chances of birds with abnormal colouration individual. coming to light and provide us with a better sense of the Reclassification of museum specimens and previous frequency and consequences of such abnormalities. records We located 10 reported cases of plumage abnormal - Acknowledgements ities in Atlantic Puffins, 23 in Common Murres, and 18 K. Hobson, G. Robertson, S. Wilhelm, Environment in Thick-billed Murres. Of these, we determined the Canada, and the Bold Coast Charter Company (Cutler, precise form of colouration abnormality from the des - Maine, USA) provided logistic support for our studies cription, original specimen, or photographs and reclas - in Newfoundland and New Brunswick. Work in the sified 10, 16, and 12 specimens of each species, re - Witless Bay Ecological Reserve was done with the per - spectively (Table S1). Most of these were originally mission of the Parks and Natural Areas Division of the described as “albino,” “partial albino,” or “melanistic,” Newfoundland and Labrador Department of Environ - the most extreme forms of plumage colouration anom - ment and Conservation, and research on Machias Seal alies, and all were reclassified as a milder form. Among Island was done with the permission of the Canadian live and museum specimens combined, progressive Wildlife Service and Canadian Coast Guard. We thank greying was the most common abnormality (16 or 17 J. Araney, A. Lang, D. Pirie-Hay, G. Robertson, and S. cases out of 41), followed by brown (10 of 41 cases), Wilhelm for assistance in the field. C. Cicero (Museum with 6 cases each of dilution and melanism (Table S1). of Vertebrate Zoology), M. Gosselin (Canadian Muse - Partial leucism was the least common abnormality and um of Nature), J. Hinshaw (University of Michigan occurred in only 2 Common Murres (Table S1). Museum of Zoology), P. Sweet, P. Campianolo, and T. Trombone (American Museum of Natural History), Discussion and H. van Grouw (Natural History Museum, United Of the 52 cases of colour abnormalities in these 3 Kingdom) provided information on museum speci - species, we were able to reclassify 40 (80%) and add mens, and M. Harris discussed plumage abnormalities an additional 4 cases, including rarer abnormalities in puffins. A. Patterson, P. Wilcox, L. Murison, and M. af fecting bare parts (Table S1). Previous classifications Zani provided details on sight and photographic rec - tended to adopt one of the extremes (albinism/mela n - ords. This manuscript was improved by comments from ism) rather than the underlying mechanistic explana - A. J. Gaston, H. van Grouw, and S. G. Sealy. tion, which masks the root cause for the colouration abnormality and artificially groups abnormalities that Literature Cited Andersson, S., J. Örnborg, and M. Andersson. 1998. Ultra - have no common mechanism. This greater understand - violet sexual dimorphism and assortative mating in blue ing of the mechanisms of colour aberrations has re- tits. Proceedings of the Royal Society of London B Bio - ceived more attention only recently (van Grouw 2010 , logical Sciences 265: 445–450. 2013). Abnormalities of bare parts are less commonly Arnold, J. R. 1950. An albino murre. Condor 50: 141. reported than those of plumage, despite being noted Barrowclough, G. F., and F. C. Sibley. 1980. Feather pigmen - in auks since the 19th century (Krüper 1857; Newton tation and abrasion: test of a hypothesis. Auk 97: 881–883. 1877). This may be because birds with abnormal plu- Cairns, D. K. 1986. Plumage colour in pursuit-diving sea- mage are easier to identify in large colonies, such as birds: why do penguins wear tuxedos? Bird Behaviour 6: cliff-nesting murres, whereas birds with abnormally 58–65. Clarke, W. J. 1913. Notes for 1912 from Yorkshire. British coloured bare parts would not stand out so clearly. 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Jakubas, D., and K. Wojczulanis-Jakubas. 2012. Not always Ross, C. C. 1964. Albinism among North American birds. black and white: colour aberrations in the Dovekie. Arctic Cassina 47: 2–21. 65: 229–232. Roulin, A. 2004. The evolution, maintenance and adaptive Jones, I. L. 1990. Plumage variability functions for status sig - function of genetic colour polymorphism in birds. Biolog - nalling in least auklets. Animal Behaviour 39: 967–975. ical Reviews 79: 815–848. Kelly, T. C. 1980. Leucism in a guillemot. Irish Birds 1: 532– Sage, B. L. 1962. Albinism and melanism in birds. British 535. Birds 55: 201–225. Krüper, T. 1857. Ornithologische miscellen. Naumannia 7: Sage, B. L. 1963. The incidence of albinism and melanism 436–438. in British birds. British Birds 56: 409 –416. Leopold, M. F., R. S. A. van Bremmelen, and H. Verdaat. Sealy, S. G. 1969. Color aberrations in some alcids on St. 2010. Grijze alk bij de bruine bank. Sula 23: 38–39. Lawrence Island, Alaska. Wilson Bulletin 81: 213–214. Lockley, R. M. 1953. Puffins. J. M. Dent and Sons, London, Shaughnessy, P. D. 1970. The genetics of plumage phase United Kingdom. dimorphism of the southern giant petrel Macronectes gigan - Loomis, L. M. 1918. Expedition of the California Academy teus . Heredity 25: 501–506. of Sciences to the Galapagos Islands, 1905–1906. XII. A Storer, R. W. 1952. A comparison of variation, behavior and review of the albatrosses, petrels, and diving petrels. Pro - evolution in the sea bird genera Uria and Cepphus . Univer - ceedings of the California Academy of Sciences 2: 1–187. sity of California Publications in Zoology 52: 121–222. Mancini, P. L., S. Jiménez, T. Neves, and L. Bugoni. 2010 . Summers, C. G. 2009. Albinism: classification, clinical char - Records of leucism in albatrosses and petrels (Procellari - acteristics, and recent findings. Optometry and Vision Sci - iformes) in the South Atlantic Ocean. Revista Brasileira ence 86: 659–662. de Ornitologia 18: 245–248. Tuck, L. M. 1961. The murres. Monograph series 1. Canadi - McCardle, H. 2012. Albinism in wild vertebrates. M.Sc. the - an Wildlife Service, Ottawa, Ontario, Canada. sis, Texas State University-San Marcos, San Marcos, Texas, van Grouw, H. J. 2010. How to recognize colour aberrations USA. in birds (in museum collections). Journal of Afrotropical Murphy, T. G., and T. T. Pham. 2012. Condition and bright - Zoology Special Issue: 53–59. ness of structural blue-green: motmot tail-racket brightness van Grouw, H. J. 2013. What colour is that bird? The causes is related to speed of feather growth in males, but not in and recognition of common colour aberrations in birds. fe males. Biological Journal of the Linnean Society 106: British Birds 106: 17–29. 673–681. van Grouw, H. J., S. Russell, and O. J. Merne. 2011. Notes Newton, A. 1877. A variety of the guillemot ( Alca troile ). on colour aberrations in Common Guillemot Uria aalge Proceedings of the Zoological Society of London 1877: 2. and Northern Gannet Morus bassanus . Seabird 24: 33–41. Reinsch, H. H. 1983. Albinismus bei Dickschnabellumme, Uria lomvia , und Eichelhäher, Garrulus glandarius . Bei - Received 23 January 2015 traege zur Vogelkunde 29: 318–319. Accepted 28 February 2016

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