Examination of the Bill and Buccal Cavity of an Adult Razorbill
(Alca torda) Specimen from the Dry Tortugas, December 18, 2012
Rostrolateral view of an Alca torda bill and buccal organs, sketched from direct examination of a corpse found in Dry Tortugas National Park on 12/18/2012. CLICK IMAGE TO ENLARGE.
During the winter of 2012-2013, hundreds of Razorbills were seen off the coasts of South Florida; an unprecedented event in recorded history. Most of the documented birds that engaged in this ‘invasion’ event appeared to be young or immature. The distinctive bill of the species has received significant attention in several studies, and has been suggested as an aid in determining the maturity of individual birds. Considering the relative few adult individuals documented on Florida waters this winter, the finding of an adult Alca torda corpse in the Dry Tortugas (DRTO) during the Christmas Bird Count (113th) expedition offered a unique opportunity to closely examine a mature bill. The fresh specimen also presented an opportunity to study the uncorrupted interior of the mouth and the buccal organs, which are adapted to catching and retaining fish prey and are bright yellow in adult birds.
The bill of the Alca torda – or Razorbill – is perhaps the most distinctive physical trait of the species. With maturity, the bill may grow to be a relatively large and laterally compressed organ with a steeply curved culmen and a complex sculpting of grooves and ridges. Birds develop an arching white “band” across the upper and lower mandibles as they age, and rarely two bands. The bill undergoes many stresses while foraging and during breeding disputes, and may develop a buttressed sculpt to reinforce the narrow organ. The mouth interior and tongue of adults are an intense warm yellow. These are used in breeding displays and to incite feeding behavior.
By contrast, the bills of younger birds have flatter culmens, and may be entirely devoid of grooves. At all ages, the bill is primarily black and glossy, with a hooked tip and sharp distal tomia that flatten and broaden towards the gape. The interior of the juvenile mouth is pale yellow.
Fig. 1. Lateral view of Alca torda bill, sketched from direct observation, DRTO 12/18/2012 specimen. CLICK IMAGE TO ENLARGE.
After breeding, Razorbills will shed the outermost layer of the bill, reducing its overall size, and dulling its gloss and color. This shedding of the external sheath of the bill is true also of other alcids, particularly the puffins (Fratercula). With the arrival of the breeding season, an outer keratin layer develops; in the Alcidae this may be manifested in the form of plates, knobs or horny projections that serve as ornamentation for courtship purposes. In Razorbills, the sculpting of the bill’s grooves becomes pronounced, the ridges smoother and rounder, and the black darker and glossier, contrasting a whiter band when present.
Studies at breeding colonies have explored the potential for aging individual Razorbills based on the number and characteristic of bill grooves (Jones 1988). However, the shape and sculpture of bills is variable individually, geographically and possibly even sexually in Razorbills. It is unclear how sexual maturity relates to bill development. Some Razorbills have been known to breed earlier in Nearctic colonies, compared to the Palearctic (Lavers et al. 2008). In general, the bill develops slowly in the species, and may continually become deeper and more grooved in some birds as they age.
R. A. Galvez. Adapted from Jones 1988, Lloyd and Perrins 1977, Lavers et al. 2008. CLICK CHART TO ENLARGE.
The distinctive white band across the bill does not generally begin developing until approximately 15 months of age, so it is unlikely that birds in their first winter show any sign of this trait. Rarely, old birds with two bands have been documented, but this may be associated with geographical tendencies. As Razorbills age and the bill becomes deeper, one proximal groove, and up to three grooves distal of the white band may develop.
The ageing of Razorbills based on the development of their bills cannot be done with accuracy, in part because sexual maturity in the species may be attained anytime between the third and sixth year of life. Some birds may never develop more than a single distal groove their entire lifespan, and this may influence their success as breeders (De Wijs 1983). A 1988 study by Jones demonstrated through the trapping and measuring of breeding birds in the Palearctic that a small margin of birds (9%) had developed a third groove distal of the white band. Only birds past their third summer had more than two distal grooves. That study also showed a tendency for bill depth and wing length to increase with age.
The impromptu situation during this Christmas Bird Count expedition to the Dry Tortugas (December 18-20, 2012) did not allow for the proper measuring of the found corpse, therefore it is hoped that a prepared specimen will supply future details. On the other hand, this might have been one of the only opportunities to examine the remarkable yellow buccal cavity of the bird, since internal tissue deterioration had likely commenced, and frozen birds may present challenges in this camp.
In addition to ornamental bill parts, the Alcidae are also known for colorful mouth interiors, used during threat displays and courtship. These range from light blue in Ancient Murrelet (Synthliboramphus antiquus) to vermilion in guillemots (Cepphus), and are yellow-orange in breeding Razorbills. Additionally, their mouths and tongues are equipped with specialized projections and gripping accessories that allow for the trapping of slippery prey.
During the breeding season, Razorbills may carry multiple fish from several fishing bouts crosswise on their bills. Prey such as capelin or sandlace are carried by breeding adults from the ocean back to nest sites on cliff ledges, and fed to the young. The Razorbill’s buccal organs are well adapted for continually foraging and catching prey while securely retaining earlier catches. After fish are caught, they are slipped back towards the gape with the aid of the smooth anterior end of the tongue. The fish are gripped tightly against the horny palate by projections on the posterior of the tongue. The knobbed and flexible inner lining of the mouth helps secure prey. In this manner, a Razorbill is able to repeatedly open its bill to continue foraging while as many as half a dozen fish remain in place.
Working from the Dead Specimen
I found this freshly deceased Razorbill washed up on the southeastern-most shoreline of Garden Key, in Dry Tortugas National Park on the evening of December 18, the night before the annual Christmas Bird Count. During these occasions, our count team stays in the barracks of Fort Jefferson, which offer modest accommodations but no suitable work space. I spent much of that night awake, sketching this Razorbill under flashlights and my headlamp.
The renderings of the bill and buccal cavity of this specimen were drafted from direct observation, aiming to depict proportionate relationships in the form and shapes of each organ. No calipers were used, nor was there an opportunity to make drawings based on measurements. The specimen was surrendered to the National Park Service and transported back to the mainland in the freezer of the NPS M/V Fort Jefferson vessel. I was informed that it would be taken to a research facility for preparation and possible eventual submission to the Florida Museum of Natural History at UF.
References:
Barrett, R.T., Ankier-Nilssen, T. & Krasnov, Y.V. 1997. Can Norwegian and Russian Razorbills Alca torda be identified by their measurements? Marine Ornithology 25: 5–8.
Chapman, Frank M. 1966. Handbook of Birds, Eastern North America. Dover Publications, New York.
De Wijs, W. J. R. 1983. Proposed ageing system for Common Guillemot and Razorbill. Nieuwsbrief Nederlands Stook. Omler. 16: 123-132.
Feducia, Alan. 1999. The Origin and Evolution of Birds. Yale University Press, New Haven.
Johnsgard, Paul A., “Diving Birds of North America: Species Accounts — Auks (Alcidae)” (1987). Diving Birds of North America, by Paul Johnsgard. Paper 11.
Jones, P. Hope (1988): Post‐fledging wing and bill development in the Razorbill Alca torda islandica, Ringing & Migration, 9:1, 11-17.
Lavers, Jennifer, Mark Hipfner, Gilles Chapdelaine and J. Mark Hipfner. 2009. Razorbill (Alca torda), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/635 doi:10.2173/bna.635
Lavers, J.L., I.L. Jones and A.W. Diamond. 2008. Age at First Return and Breeding of Razorbills (Alca torda) on the Gannet Islands, Labrador and Machias Seal Island, New Brunswick. Waterbirds 31(1): 30-34, 2008.
Lloyd, C. S. and C. M. Perrins. 1977. Survival and age at first breeding in the Razorbill. Bird-Banding 48: 239-252.
Lofgren, Lars. 1987. Ocean Birds. Crescent Books, New York.
Proctor, Noble S. and Pratrick J. Lynch. Manual of Ornithology. Avian Structure & Function. Yale University Press, New Haven.
Pyle, P. 2008. Identification guide to North American birds. Part II: Anatidae to Alcidae. Slate Creek Press, Point Reyes Station, CA.
Welty, Joel C. and Luis Baptista. 1988. The Life of Birds. Fourth Edition. Saunders College Publishing, New York.
When I visited Andalucía during the spring of 2002, the Red-necked Nightjar (Chotacabras Pardo o Cuellirojo – Caprimulgus ruficollis) had for long been a fascination of mine. It was a bird I had not yet experienced in the Western Palearctic, and my love of Caprimulgiformes had me searching for the species with increased enthusiasm.
A watercolor study of a Red-necked Nightjar’s wing, done soon after a bird was found, killed by the roadside.
During the last two weeks of May, I crisscrossed the Costa del Sol several times from Ronda through Málaga, spending some time along the Río Guadalhorce, stopping at Macharaviaya (ancestral home from my father’s side), and eventually passing Almería to the east.
By the time I reached Cabo de Gata – the region’s largest coastal protected natural park – I had seen several excellent species including hundreds of Greater Flamingos, ducks and shorebirds in the salinas (salt flats) of the region, but no Red-necked Nightjars.
A pair of quick sketches on the gessoed rough side of a masonite board. These are relatively underdeveloped, so I hesitated in including them here, but they tell part of the story. Left is Macharaviaya – a whitewashed town near Málaga. Right is a view of the Mediterranean from the coastal cliffs of Cabo de Gata.
During my first night at Cabo de Gata, I camped out near coastal crags that dropped steeply into the Mediterranean, taking drives and hikes up the arid roads and rocky cliffs north of the cape. I had heard an Eagle Owl from this location, so was intent in pushing my investigation well into the night. However, it was when I unexpectedly flushed nightjars from a sandy path that I was truly captivated.
I stopped the car at the sight of a Red-necked Nightjar on the ground, perfectly lit by my headlights, and managed to step out with my watercolor and paper to sketch the bird under the harsh artificial lights.
Unfortunately, several minutes into my sketch a police vehicle drove up and the bird flew off. I was immediately put under scrutiny and suspicion. The two officers were pleasant gentlemen not much older than me, and were very curious as to what I was doing. I showed them my sketches and they listened to my description of the nocturnal bird with interest. After an awkward moment of silence during which I continued painting under my headlamp, the officers simply excused themselves and drove away.
About a week later, while coiling down mountainous highways from Grazalema towards Málaga late at night, I saw a vehicle in front of me hit a nightjar. I pulled over to inspect the victim, and found a perfectly intact and fresh C. ruficollis specimen. Arriving at Tarifa – the southernmost coast of southern Spain – I spent it sleepless, doing the sketches included in this post.
Watercolor and ink pen study of the upper side of the open wing of a Red-necked Nightjar. Sketched in Tarifa, Spain soon after the bird was found dead.
A watercolor study of a Red-necked Nightjar wing, done soon after the bird was found, killed by the roadside.
Study of the open mouth of a Red-necked Nightjar, done with watercolor, ink pens and color pencil. While the bills of nightjars are relatively small, their open mouths are very large. Note the stiff bristle feathers lining the mouth, which act as nets to help bring insect prey into the buccal cavity.
Dorsal study of a Red-necked Nightjar done with ink and color pencils.
The Process
This slideshow requires JavaScript.
1. I typically start most field sketches with a pencil drawing. If the shapes, posture and placement of body parts are not generally accurate in the pencil gesture, then the painting runs the risk of turning out awkward. If the bird flies before I can start applying watercolor, then I’m left with a sketch I can use for future field paintings or as reference for a longer study.
2. In this case, I had the time to put down a substantial amount of watercolor before the bird was flushed. Unfortunately, the nightjar was left half-painted, and I was without a life reference to confidently finish the rendering.
3. However, when I found a dead Red-necked Nightjar days after, and created detailed studies of its plumage, I was able to use my specimen renderings to bring the field sketch to a more complete place.
4. Over the years, I often wondered if I could add just a few strokes to further develop this sketch without ruining it. I kept the hope of returning to Andalucía during spring, and re-encountering a Red-necked Nightjar under similar circumstances, but this has not happened yet. As time went by, the more I looked at the watercolor studies of the deceased bird, the more I felt I should add dabs of color to the field sketch, to give it the desired density. One summer night, as I drove through the Everglades and came upon a Chuck-will’s Widow with my lights, I immediately thought of the nightjar. Both the Red-necked Nightjar and the “Chuck” were until recently classified in the same genus – Caprimulgus (the latter has been recently changed to Antrostomus). Both species are roughly the same size, and both tend to exhibit overall rustier plumages than their congeners. As I carefully observed the Chuck-will’s-widow through my binoculars, it occurred to me that I was gaining useful information to help me develop the field sketch from Andalucía – not in the detail of the plumage or in the shape of the bird, but in the way a nightjar appears under the contrasting lights of car, and how this affects its volume and color.
galvezbirds.com 