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Calculating food production in the subsistence harvest of birds and eggs.

ABSTRACT. Subsistence harvest studies use number-to-mass conversion factors (CFn-m) to transform numbers of animals harvested into food production (CFn-m = body mass x recovery rate; where recovery rate is the percentage of the body mass represented by the processed carcass). Also, if egg harvest was reported as volume (e.g., a bucket), volume- to-number conversion factors (CFv-n) are needed to calculate the number of eggs taken. Conversion factors (CF) for subsistence harvest of birds and eggs have been based on unclear assumptions. We calculated a mean recovery rate (65%) by weighing and processing wild birds, compiled data on bird and egg mass, developed an egg CFv-n equation, and presented CF for 88 bird species, 13 subspecies or populations, and 25 species categories likely to be harvested in Alaska. We also made recommendations on how to apply and adjust CF according to study objectives. We recommend that subsistence harvest studies (1) collect egg harvest data as egg numbers (not volume); (2) clearly explain considerations and assumptions used in CF; (3) report recovery rates and mass of birds and eggs; and (4) cite original sources when referring to CF from previous studies. Attention to these points of method will improve the accuracy of food production estimates and the validity of food production comparisons across time and geographic areas.

Key words: bird; egg; subsistence harvest; subsistence hunt; harvest survey; food production; edible mass; recovery rate; number-to-mass conversion factor; volume-to-number conversion factor; Alaska

RESUME. Les etudes sur la recolte de subsistance utilisent des facteurs de conversion nombre-masse (CFn-m) pour transformer le nombre d'animaux chasses en production alimentaire (CFn-m = masse corporelle x taux de recuperation; le taux de recuperation etant le pourcentage de la masse corporelle representee par la carcasse transformee). De plus, si la recolte des ceufs etait rapportee en volume (p. ex. un seau), les facteurs de conversion volume/nombre (CFv-n) s'averent necessaires pour calculer le nombre d'ceufs preleves. Les facteurs de conversion (FC) pour la recolte de subsistance d'oiseaux et d'ceufs s'appuient sur des hypotheses floues. Nous avons calcule une moyenne du taux de recuperation (65 %) en pesant et en transformant des oiseaux sauvages, recueilli des donnees sur la masse des oiseaux et des ceufs, trouve une equation pour les facteurs de conversion volume/nombre pour les ceufs et presente des FC pour 88 especes d'oiseaux, 13 sous-especes ou populations et 25 categories d'especes susceptibles d'etre chassees en Alaska. Nous avons egalement formule des recommandations sur la fafon d'appliquer et d'ajuster les FC selon les objectifs de I'etude. Nous recommandons que les etudes sur la recolte de subsistance (1) recueillent les donnees sur la recolte des ceufs en nombre d'ceufs (et non en volume); (2) expliquent clairement les considerations et les hypotheses utilisees pour les FC; (3) rendent compte des taux de recuperation et de la masse des oiseaux et des ceufs; et (4) citent les sources originales quand elles font reference aux FC d'etudes precedentes. L'attention portee a ces elements methodologiques ameliorera la precision des estimations de la production alimentaire et la validite des comparaisons en matiere de production alimentaire en fonction des periodes et des regions geographiques.

Mots cles : oiseau; ceuf; recolte de subsistance; chasse de subsistance; enquete sur les recoltes; production alimentaire; masse comestible; taux de recuperation; facteur de conversion nombre/masse; facteur de conversion volume/nombre; Alaska

Traduit pour la revue Arctic par Nicole Giguere.

INTRODUCTION

Number-to-Mass Conversion Factors for Birds and Eggs

Studies of subsistence uses of wild resources report harvest as the number of animals taken and as the amount of food produced (edible mass). Estimates of the number of animals taken are used to document subsistence activities, to assess harvest impacts on fish and wildlife populations, and to allocate harvestable amounts among user groups (Usher and Wenzel, 1987). Food production data are used to depict the relative importance of resources (e.g., moose, salmon, geese) and their role in subsistence economies (Brown and Burch, 1992), to assess exposure to contaminants derived from wild foods (Usher, 2000), and to estimate the monetary (replacement) value of harvest. Food production estimates do not account for the nutritional and cultural importance of different resources (Usher, 1976; Behnke, 1982). But these data are also important in adjudicating disputes among stakeholders, quantifying ecological services provided by resources and ecosystems, assessing food security, and prioritizing human activities on the basis of their socioeconomic contribution to communities' well-being (Jones, 1997; Magdanz et al., 2011; Hoover et al., 2013).

Food production is calculated by multiplying the number of animals taken by a number-to-mass conversion factor (CFn-m). A CFn-m integrates two variables: the live (whole, round) body mass of individual species or multi-species categories and the recovery (yield) rate, which is the percentage of the live mass represented by the processed carcass. Studies have commonly failed to explain assumptions used in CFn-m and to report body mass and recovery rates (Table 1). Over the decades, CFn-m developed in earlier studies have often been reused without critical evaluation or clear reference to original sources. Thus, it is difficult to evaluate discrepancies in CFn-m and food production estimates across studies. For instance, conversion factors (CF) used for Sandhill Crane Grus canadensis harvested in Alaska (CFn-m = 10-15 lb/bird) (4536-6804 g/bird) (Patterson, 1974; Wolfe, 1981; Ikuta et al., 2014) appear overestimated when compared to body mass of the subspecies occurring in Alaska (8.17-9.82 lb) (3705-4455 g) (Rodewald, 2015).

To inform stakeholders who rely on accurate food production data, studies need to clearly report considerations and variables used to derive CF. From subsistence users' perspectives, biased-low recovery rates conflict with the non-waste principle that is intrinsic to their cultural values (Zavaleta, 1999). Biased-low recovery rates also lead to underestimating the importance of wild resources in subsistence economies. On the other hand, biased-high recovery rates can discredit food production assessments and their use in mitigation and litigation.

Birds and eggs are a small proportion of the subsistence harvest, but data on their food production help address complex management issues (Fienup-Riordan, 1999; Zavaleta, 1999). In Alaska, the subsistence harvest (about 34 million edible pounds per year) is composed of fish (53%), land and marine mammals (23% and 14%), plants (4%), shellfish (3%), and birds and eggs (3%) (Fall, 2016). Developing CF for each of these resource categories involves diverse challenges. Previous efforts to consolidate information and clarify CF have addressed all resource categories (JBNQHRC, 1982; Usher, 2000; Ashley, 2002), and because this task is immense, some issues pertaining to bird and egg CF remained unresolved.

For wild birds, body mass depends on species, subspecies, population, sex, and age. Within these categories, body mass also varies because of ecological conditions and breeding, migration, and feather molting cycles (Piersma and Lindstrom, 1997). Because of difficulties in species identification, harvest surveys use species categories, which also complicate CFn-m because the species within a category may differ in body mass. Some studies have defined CFn-m for categories that include species with considerable size difference (Patterson, 1974). For instance, surveys have used one category for gull eggs, but eggs of large gulls are at least twice the size of those of small gulls (Rodewald, 2015). Social science researchers and other staff working on harvest surveys are often unfamiliar with the identification (including size), distribution ranges, and relative abundance of the dozens of bird species that may be harvested in a region. Thus, it is often difficult for them to critically evaluate bird and egg CF used in previous studies and to generate new CF.

Recovery rates in subsistence harvests depend on harvesting and processing conditions, cultural practices, species, and food preferences (Burch, 1985; Usher, 2000). Assumptions underlying recovery rates used for subsistence bird harvest are sometimes unclear, and rates have ranged from 40% to 75% (Table 1). Although in Alaska Native cultures birds have not been widely used as dog food, recovery rates in some earlier studies considered harvest for such use (Usher, 1970) and likely differ from rates that consider human consumption only. In many studies, recovery rates for egg harvest do not indicate whether the shell mass was included as edible mass.

Volume-to-Number Conversion Factors far Eggs

To facilitate accurate recall of harvest events and minimize burden on respondents, harvest surveys may use reporting units that are meaningful to harvesters (fish tub, truckload of wood, bucket of eggs) (Tobias and Kay, 1994). Even when respondents are asked to report their harvest in number of eggs, some values may instead be recorded in volume. For such cases, volume-to-number conversion factors (CFv-n) allow calculating the number of eggs taken. A method to estimate egg CFv-n involves comparing the mass (as a proxy for volume) of wild bird eggs to that of chicken eggs (J. Magdanz, pers. comm. in Naves, 2010). But CFv-n estimated in this way seemed high compared to CFv-n based on researcher or key respondent information (Burch, 1985; Fall et al., 1995). Use of padding material (e.g., grass, moss) to protect eggs reduces the total volume of eggs in a given container (Hunn et al., 2003). To clarify assumptions and refine this estimation method, we considered the use of padding material and the fact that full containers may not be filled to the brim to prevent egg damage during harvesting and transporting.

Study Objectives

It is impractical to account precisely for all sources of variation in recovery rate and wild bird body and egg mass that may affect CF (Usher, 1976; Burch, 1985). Thus, rather than defining highly precise and detailed CF, the objectives of this study were (a) to develop CFn-m and CFv-n equations based on clear variables and assumptions that can be easily adjusted depending on the study objectives and context and (b) to provide recommendations on the development and use of CF that will increase the accuracy of food production estimates and the validity of food production comparisons across time and geographic areas.

To achieve these objectives, we first collected ethnographic information from key respondents on subsistence practices in bird processing and egg harvesting. Information on bird parts usually consumed was needed to clarify which parts should be included as edible mass when calculating a recovery rate reflecting subsistence practices. Information on use of containers and padding material in egg harvesting was needed to refine the CFv-n estimation method. Second, we processed and weighed wild birds to calculate a recovery rate. Third, we compiled data on bird and egg mass, as well as distribution ranges and population sizes, for species likely to be harvested in Alaska. We integrated these social science and biological data to develop CFn-m and CFv-n equations and calculated CF for use in harvest studies. Although we addressed bird species composition and subsistence practices in Alaska, our approach and recommendations also apply to food production studies of other resources and regions.

METHODS

Ethnographic Information on Bird Processing and Egg Harvesting

To calculate the bird recovery rate, the first step was to determine which bird parts should be included as edible mass, depending on how birds are processed and which parts are usually consumed. Also, refining the CFv-n equation required information on egg harvesting (see below). To gather this ethnographic information, we designed 14 questions on bird and egg harvesting and processing related to CF (online Appendix S1). The questions asked for information about general harvesting and processing in a region, as opposed to individual practices. We identified 27 Alaska Native people as key respondents who could provide information on the subsistence harvest and culture in their region of origin. Rather than a random selection of individuals within a sampling universe, key respondents are particularly knowledgeable people who can provide expert opinion on a domain (Huntington, 1998; Bernard, 2011). Participation in the Alaska Migratory Bird Co-Management Council (AMBCC, 2016) and in the U.S. Fish and Wildlife Service Refuge Information Technician Program were the main criteria used to identify such individuals. Participation in these programs served as an index of key respondents' extensive experience as subsistence resource users and community leaders, including their engagement in harvest management.

In April-May 2015, printed copies of the questionnaire were distributed in person or via postal mail to the potential respondents. Pre-stamped, pre-addressed envelopes were provided for return of completed questionnaires. We followed ethical principles for social science research, including informed consent and voluntary participation (ARCUS, 1999). One month after the questionnaire was first distributed, a reminder was mailed to people who had not yet returned responses. We received 16 completed questionnaires (a 59% response rate). In results pertaining to the questionnaire, "n" refers to the number of responses to individual questions or the number of times respondents indicated a categorical answer (yes, no, sometimes).

Respondents were instructed to leave fields for answers blank if they did not know the answer or if some species categories did not occur or were not used in their region. Most responses referred to species categories commonly harvested across Alaska (ducks, geese, grouse, and ptarmigan). Fewer responses were obtained to questions related to egg harvest than to those about bird harvest.

Definition of Edible Mass

On the basis of key respondent information (see Results), we defined the edible mass as including the carcass mass (meat, bones, skin, fat, and other tissues remaining after removal of feathers, feet, head, and viscera), as well as the heart and gizzard mass because these parts were also usually consumed by subsistence users. Although some responses indicated that other parts are sometimes consumed (e.g., liver, blood, intestine, stomach; see Results), these responses were infrequent, and the exclusion of these parts from the edible mass was inconsequential for the calculation of the recovery rate. While the exclusion of these parts may have resulted in a minor underestimation of the recovery rate, such underestimation was likely offset by the inclusion of skin and bones, which are sometimes not consumed. In Alaska, wild birds and eggs cannot be bought of sold and therefore have no defined market value. To facilitate assessments of the monetary value of wild foods, the definition of edible mass must be comparable to likely replacement products available in grocery stores. For bird eggs, we used a recovery rate of 100% (whole egg including the shell). Although the shell is not consumed, chicken eggs are a likely replacement product and are sold whole and by the dozen (not directly by weight).

Processing and Weighing Wild Birds to Calculate Bird Recovery Rate

To calculate the mean bird recovery rate according to our definition of edible mass, we weighed and processed wild birds harvested for home use in September-October 2015 and September 2016. This sample included ducks (n = 18), geese (n = 9), and ptarmigan (n = 2) harvested at several locations in south-central Alaska and the Alaska Peninsula. Mass measurements were obtained using an electronic scale with precision of one gram. We weighed the whole body mass of freshly killed birds. We plucked and singed the birds, removed the head, wing tips (cut at the metacarpus and ulna-radius joint), feet (cut at the tarsometatarsus and tibia-fibula joint), and all viscera. The skin-on, bone-in mass of birds thus processed constituted the carcass mass. After the carcass mass was recorded, we cut out and weighed the breast fillets (boneless, skin-on outer and inner fillets, or pectoralis and supracoracoideus muscles) and the whole leg (bone-in, skin-on thigh and drumstick, or tibia-fibula and tarsus sections). We also weighed the heart and clean gizzard (opened to remove food remains and its tough inner lining) to be included as edible mass. Weights were presented as arithmetic means of proportions of the live mass including all species. The bird recovery rate was calculated as the mean proportion of the carcass, heart, and gizzard mass relative to the live mass.

Breast fillets and whole legs are common cuts in sport hunting and poultry processing. Mass data for these cuts are useful to gauge recovery rates used in previous subsistence harvest studies, to generate alternative recovery rates based on different processing methods, and to allow comparisons with potential replacement poultry products.

Bird and Egg Mass Data

We compiled body and egg mass data for bird species, subspecies, and populations occurring in Alaska from Rodewald (2015) unless otherwise noted. Data on sex and age composition of Alaska subsistence bird harvest were unavailable. Thus, it was impossible to account for variation in body mass among sex and age categories in the harvest. We calculated the arithmetic mean body mass including data for males, females, adults, and immatures (as available) to represent sex and age categories potentially harvested. Mass data referred to Alaska-breeding populations in spring (as available) because at least 51% of the annual bird subsistence harvest occurs in spring (Paige and Wolfe, 1997). We used mass of freshly laid eggs because water loss during incubation decreases egg mass.

Mean body and egg mass were calculated based on all items (species, subspecies, populations) within categories. Population size data were used to weight mass means. Population size data were sometimes unavailable because (a) populations have not been monitored; (b) surveys have not differentiated among species (goldeneyes, mergansers, scoters, scaups) (Stehn et al., 2013; Platte and Stehn, 2015); and (c) estimates of abundance were not directly comparable among items within categories. If mass data were unavailable for one or more items within a category, means or reference values were defined by considering data for similar species and species distributions (online Appendix S2).

Body and egg mass data were reported both in grams (as rounded numbers with no decimal places) and in pounds (body mass data with two decimal places and egg mass data with three decimal places). Rather than displaying excessive precision, this level of detail when dealing with mass data in pounds was needed to properly represent mass of small birds and eggs.

Number-to-Mass Conversion Factors for Birds and Eggs

Using the recovery rates defined in this study and the bird and egg mass data compiled, we calculated CFn-m as:

Bird CFn-m = 0.65 x body mass (see Results). (1)

Egg CFn-m = 1.00 x egg mass (shell included). (2)

Volume-to-Number Conversion Factors for Eggs

To refine the CFv-n estimation method based on comparison of chicken and wild bird egg mass, we considered how the volume of eggs in a container is affected by (1) use of padding material and (2) not filling the container to the brim. First, to assess whether these considerations reflect egg harvesting practices, the key respondent questionnaire included questions on characteristics of containers used, frequency of use of padding material, and whether containers are only partially filled to avoid egg loss and damage during transport (questions 8-11, online Appendix S1). On the basis of information from key respondents (see Results), we assumed that padding material is always used and that full containers are filled to 80% of their capacity.

To quantify the reduction of the volume of eggs in a container resulting from use of padding material, we packed large chicken eggs (24 ounces or 680 g per dozen) (U.S. Department of Agriculture, 2000) in a one-gallon (3.8 L) bucket, filling it without any padding material and then adding dry grass between egg layers. We repeated packing with and without grass three times and counted the number of eggs needed to fill the bucket to the brim. With grass, the number of eggs needed to fill the bucket in the three measurements was 37, 36, and 33 eggs (mean = 35.3). Without grass, in each of the three measurements, 48 large chicken eggs were needed to fill the bucket.

We then developed a CFv-n equation including four variables: (a) number of chicken eggs needed to fill a 1-gallon (3.8 L) bucket; (b) proportion of container volume filled; (c) mass of a chicken egg; and (d) mass of a wild bird egg [CFv-n = (a x b) x (c / d)]. Considering our assumptions:

Number of eggs/gallon: CFv-n = (35.3 x 0.8) x 0.126 / mass of wild bird egg, in pounds). (3)

Number of eggs/L: CFv-n = (9.3 x 0.8) x (57.0 / mass of wild bird egg, in grams). (4)

RESULTS

Number-to-Mass Conversion Factors for Birds and Eggs

The following ethnographic information was used to identify bird parts that should be included as edible mass when calculating the recovery rate. Key respondents reported that Alaska subsistence users consumed wild birds as bone-in, skin-on preparations, usually as a roast or soup (see also Mishler, 2003; Unger, 2014). Birds were consumed fresh or preserved by freezing, drying, or canning. Bird processing involved plucking, singeing, and gutting birds.

Meat from the breast, legs, neck, head, back, and wings was usually consumed, as well as skin, fat, heart, and gizzard (Table 2). The liver was indicated as consumed in more than half of responses. Other parts were identified as not usually consumed, but some respondents indicated consumption of blood, intestine (ptarmigan, ducks, and geese), stomach (ducks and geese), kidney, and tongue. Bones were boiled to render broth, and bone marrow was sometimes consumed. We did not ask about non-food uses, but respondents reported that sometimes goose down was used and that the viscera and wings of harvested birds were used as bait in traps for fur animals.

Plucking seemed a preferred processing method among subsistence users, although skinning was sometimes used as a quicker option. To facilitate plucking, birds whose feathers are difficult to remove (swan, crane, seabirds, sea ducks) may be dipped in hot water. Such birds were sometimes skinned. The thin skin of grouse and ptarmigan often tears off during plucking, so these birds were commonly skinned. Plucking allowed consumption of the skin and associated fat. We asked respondents what proportion of the bird's body weight they thought is usually consumed when birds are plucked or skinned (the recovery rate) (questions 5 and 6, online Appendix S1). Responses ranged from 50% to 100%, but some seemed to refer to total mass after processing [recovery rate = 100% (n = 2) and "90% minus guts and bones"]. Because this question seemed unclear to respondents, we based the recovery rate solely on the data from wild birds processed in this study.

Using our data from processed wild birds (n = 29), the mean carcass mass was 60% of the live mass (range = 54%-66%), the heart was 1% (range = 0.5%-1.2%), and the gizzard, 4% (range = l%-7%), resulting in a mean bird recovery rate of 65% (range = 56%-70%) (Table 3). Breast fillets were 22% (range = 18%-28%) and the legs were 10% (range = 7%-13%) of the live mass.

Using a recovery rate of 65% for birds and 100% for eggs and the mass data compiled, we calculated bird and egg CFn-m for 88 bird species, 13 subspecies or populations, and 25 species categories (Table 4 and online Appendix S2).

Volume-to-Number Conversion Factors for Eggs

The key respondent questionnaire indicated that five-gallon (19 L; n = 7) and one-gallon (3.8 L; n = 5) buckets were commonly used for egg harvesting, but that any available container may be used (basket, tea pot, bag, cooler; n = 8). In areas where eggs were commonly harvested, padding material was almost always used (question 9.a, online Appendix SI: "every time" n = 7, "three out of four times" n = 2). Padding was sometimes not used in murre egg harvesting because murre eggs are sturdy. Responses indicating infrequent use of padding material occurred for regions where eggs are harvested occasionally and in small numbers ("two out of four times" n = 2, "one out of four times" n = 1, "do not use moss, grass" n = 1).

Some responses to the question on whether containers are only partially filled to avoid egg loss and damage during transport considered (1) the volume of padding material separately from the volume of eggs; (2) whether eggs were abundant enough to fill containers; or (3) the number of eggs that people needed and intended to harvest (question 10.a: "yes" n = 7, "no" n = 2; question 10.b: "yes" n = 6, "no" n = 7; question 10.c: mean = 69%, range = 30%-100%). Although these questions may have been understood differently by some respondents, responses indicated that, even if reports refer to full containers (e.g., two buckets), these were often not filled to the brim.

We asked the number of eggs packed in a five-gallon bucket (question 11, online Appendix S1). Only one respondent provided direct information on the number of eggs per gallon, and the answer indicated a range (36-60 large gull eggs in a five-gallon bucket). Three respondents indicated proportions of volume, which suggested that this question was not clear for them. Two respondents specified that they did not know the answer (e.g., "1 never count them"). This question was left blank in the remaining 10 completed questionnaires.

DISCUSSION

Bird Recovery Rate

Although the composition of our wild bird sample reflected species availability at a limited set of locations and time of the year, our results were consistent with diverse data sources, including previous subsistence harvest studies, poultry production, and biological data on the relative mass of bird body parts. Considering the range of recovery rates used in previous subsistence harvest studies (40%-75%; Table 1), 40% was likely an underestimate, because it was little more than the percentage (32%) that we measured for only the breast fillets and legs. A recovery rate of 75% was likely an overestimate, because (1) it would involve including as edible mass bird parts other than those identified in this study as commonly consumed, and (2) it is higher than our recommended recovery rate (65%), which included the skin and bones, although the skin is sometimes removed during processing. Our recommended recovery rate (65%) agreed with several subsistence harvest studies in which the rate was based on assumptions by researchers. Even if not explained, recovery rates in some studies were likely based on information from local, Native experts and from researchers with wide experience in ethnographic work involving participant observation and residency in subsistence communities.

Selective breeding and commercial production conditions may result in differences in body composition between poultry and wild birds, but recovery rates in wild birds and poultry were similar. For poultry, the recovery rate for a carcass processed for removal of blood, feathers, head, feet, and all viscera was 65% (range = 58%-72%) of the body mass (Watt and Merrill, 1963; Fanatico, 2003; Lessler et al., 2007; Poltowicz and Doktor, 2011). The breast and legs were about 38% of the body mass (Solomon et al., 2006; Haslinger et al., 2007). In wild birds, the breast and legs were 32% of the body mass in our sample and 28% in other sources (Raveling, 1979; Thompson and Drobney, 1996; Jacobs et al., 2011).

Both our study and other sources reported the heart as 1% of the body mass of wild birds (Thompson and Drobney, 1996; Piersma and Gill, 1998; Jacobs et al., 2011). The gizzard was 1%-2% of the body mass in seabirds and shorebirds (Piersma and Gill, 1998; Jacobs et al., 2011), 5%-7% in geese (Raveling, 1979; Barnes and Thomas, 1987), and 2%-5% in ducks (Barnes and Thomas, 1987; Goudie and Ryan, 1991; Thompson and Drobney, 1996). The relative gizzard mass we obtained (4%) was at the mean for ducks and geese. Using this mean was appropriate because it was in accordance with the overall species composition of subsistence harvest in Alaska (ducks were 58% and geese were 31% of the number of birds annually taken; Paige and Wolfe, 1997).

Using the allometric equation of Prange et al. (1979), bone mass for wild bird species likely harvested in Alaska accounted for 7%-9% of the body mass (results not presented here). Because some bone mass is removed during processing (head, feet, wing tips), the lower end of this range could be used to adjust the recovery rate when exclusion of bones is appropriate.

Recovery rates must reflect prevailing processing practices, which may differ among hunting traditions. A characterization of bird processing by sport users was beyond the scope of this study. In Alaska, bird sport hunting generally applies to harvest in non-subsistence areas as defined by the Joint Board of Fisheries and Game, which are primarily urban areas (State of Alaska, 2015:5 AAC 99.015). Sport hunters pluck birds for bone-in, skin-on preparations, and a recovery rate of 60% is likely adequate for this use (if the heart and gizzard are not usually consumed). But sport hunters also commonly skin birds, and only the breast (recovery rate = 22% for skin-on processing) or the breast and legs (recovery rate = 32%) may be consumed (Shaw, 2013). These three values could be combined to generate a recovery rate for sport hunting. In contrast to subsistence harvest studies, sport hunting economic valuations have focused on hunting activities and expenditures rather than food production (Gan and Luzar, 1993; ECONorthwest, 2014). A better understanding of food production in bird sport hunting as well as other differences and similarities between sport and subsistence bird hunting traditions could help alleviate conflict between user groups and promote positive outcomes in management and conservation issues.

Egg Recovery Rate

Studies have often assumed an egg recovery rate of 100% (e.g., Georgette and Loon, 1993), although this assumption may not be clearly stated. The eggshell is 8%-14% of the total egg mass (Williams et al., 1982). Across species, larger eggs have proportionally thicker shells and higher shell mass (Rahn and Paganelli, 1989). Murre eggs are an important subsistence resource, and their shells are about 14% of the total egg mass (Williams et al., 1982). Whether to include eggshell mass within edible mass depends on the study objectives. In replacement cost evaluation, eggshell should be included as edible mass (recovery rate = 100%) because a likely store-bought replacement product (chicken eggs) would include shells. When assessing exposure to contaminants, eggshells should be excluded from the edible mass because they are not consumed. If discounting shell mass, we recommend a recovery rate of 90% for all egg harvest.

Volume-to-Number Conversion Factors for Eggs

It is possible that the previous attempt to calculate CFv-n based on 48 chicken eggs/gallon (12.6 eggs/L) (J. Magdanz, pers. comm. in Naves, 2010) assumed that padding material was not used and that containers were filled to the brim. Estimates based on these assumptions were likely too high and resulted in numbers of eggs about 40% higher than ours. For murre eggs, the CFv-n calculated using our equation (16.0 eggs/gallon) (4.0 eggs/L) was half of that estimated by Burch (1985) (32 eggs/gallon) (8.4 eggs/L). Considerations used by Burch (1985) were unknown, but this difference may be related to the fact that we assumed use of padding material. For large gull eggs, our CFv-n (17.5 eggs/gallon) (4.3 eggs/L) was higher than (a) empirical data in Hunn et al. (2003) (12 eggs/gallon) (3.2 eggs/L); (b) the range provided by a key respondent in this study (7-12 eggs/gallon) (1.8-3.2 eggs/L); and (c) the value for "gull (unidentified)" provided by a key respondent in Fall et al. (1995) (10 eggs/gallon) (2.6 eggs/L). Although our CFv-n equation relied on simple assumptions, these were clearly stated and their variables can be easily adjusted to suit different study objectives and contexts. For example, if it is known that padding was not used, the equation could consider 35 chicken eggs/gallon (9.2 eggs/L).

In harvest survey interviews, considering individual harvest events, respondents can provide the best data on the number of eggs harvested. If respondents report eggs as volume, surveyors can assist respondents by sequentially asking (1) the kind of eggs harvested (species); (2) the size of containers used; and (3) whether padding material was used. Then, respondents may be asked to estimate how many eggs were harvested. For egg harvest reported as volume, the unit used in the original report must be reported so that standard CFv-n can be applied. Undocumented conversions of egg volume to number make it difficult to compare results among studies.

Species Categories, Regional, and Seasonal Conversion Factors

Mean body and egg mass used in CFn-m should approximate the harvest composition in a given geographic area and season of the year. In this study, we calculated mean mass for species categories weighted by Alaska-wide populations. Using the same principle, the mean mass for species categories may be adjusted for smaller geographic scales. Use of means weighted by population size is relevant for categories that include species of very different sizes. However, to simplify the application of CF and facilitate comparison among studies, whenever appropriate, CF should refer to relatively large geographic areas and encompass all seasons of the year.

Regardless of the level of analytical complexity researchers can implement when using CF, we offer four recommendations for this method. First, surveyors must be prepared to assist respondents in accurately reporting the number of eggs harvested, instead of volume. Second, considerations and assumptions used in CF must be clearly explained. Third, recovery rates and mass of birds and eggs used to generate CF must be reported (with citation of the source) so that users can assess which of these two variables accounts for potential differences from CF in other studies. Fourth, if using CF from previous studies, citations must refer to original sources, avoiding second-hand citations. Attention to these points will improve the accuracy of food production estimates and our ability to compare them across time and geographic areas.

ACKNOWLEDGEMENTS

This study was funded by the Alaska Department of Fish and Game, Division of Wildlife Conservation (RSA-1155353), as a component of the Harvest Assessment Program of the Alaska Migratory Bird Co-Management Council. We thank the key respondents for sharing information on bird processing and egg harvesting; Andrew Ramey and Txai-the Tundra Drummer for providing wild birds for weighing and processing; and Christian Dau (U.S. Fish and Wildlife Service) and Craig Ely (U.S. Geological Service, Alaska Science Center) for providing unpublished data on swan body mass. We also thank Jim Magdanz, Tom Rothe, Julian Fischer, and two anonymous reviewers for suggesting improvements to manuscript drafts.

APPENDICES

The following appendices are available as supplementary files to the online version of this article at: https://arctic.journalhosting.ucalgary.ca/arctic/index.php/ arctic/rt/suppFiles/4630/0

APPENDIX S1. Key respondent questionnaire to collect ethnographic information on subsistence harvesting and processing of birds and eggs in Alaska.

APPENDIX S2. Notes to accompany TABLE 4. Conversion factors to estimate food production in subsistence harvest of birds and eggs in Alaska.

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Liliana C. Naves (1,2) and James A. Fall (1)

(1) Alaska Department of Fish and Game, Division of Subsistence, 333 Raspberry Road, Anchorage, Alaska 99502, USA

(2) Corresponding author: [email protected]

(Received 31 March 2016; accepted in revisedform 4 September 2016)
TABLE 1. Recovery rates used in conversion factors for
subsistence bird harvest.

Recovery    Information used to define      Study (2)
rate1       the recovery rate or
            reference cited

70%         Proportion of live mass of      White, 1953
            domesticated pigs (heavy-
            bodied, short-legged
            animals)

70%         White (1953)                    Foote, 1965

70%         White (1953) cited in           Usher, 1970
            Foote (1965)

70%         Carcass mass determined         Patterson, 1974
            in consultation with village
            representatives

70%         Not explained                   Wolfe, 1981; Behnke,
                                            1982; Georgette and
                                            Loon, 1993

60%-70%     White (1953) and poultry        JBNQNHRC, 1982
            carcass yield (Watt and
            Merrill, 1963), including
            meat, edible organs, half
            of the bone mass, and two-
            thirds of the mass of
            blood and feathers

60%-70%     JBNQNHRC (1982)                 Berkes et al., 1994;
                                            Tobias and Kay, 1994

70%         Poultry carcass yield,          Usher, 2000
            White (1953); JBNQNHRC
            (1982); Georgette and
            Loon (1993)

60%         Poultry carcass yield           Gambell, 1984

40%         Researcher estimate             Fall and Morris,
                                            1987; Fall et
                                            al., 1995

65%         Researcher estimate             Kristensen, 2011

68%         Poultry carcass yield           Goldstein et
                                            al., 2014

75%         Not explained                   Wolfe et al., 1990;
                                            Wentworth, 2007

Unknown     Wolfe (1981); Braund &          Fuller and George,
            Associates (1993); CSIS         1997; Brower et
            (2016a)                         al., 2000

Unknown     Wolfe (1981); CSIS              Braund & Associates,
            (2016a)                         1993; Ahmasuk et
                                            al., 2008

Unknown     Not explained                   Whiting, 2006

(1) Percentage of live mass.

(2) This table summarizes our review of literature on the
development and use of conversion factors for birds and other
subsistence resources. It does not present all documents we
consulted. It includes examples of application of conversion
factors and issues related to these factors.

TABLE 2. Consumption of bird parts by subsistence
users in Alaska. (1)

                            Breast   Legs   Neck   Head   Back

Usually consumed:

Ducks                         15      14     13     11     14
Geese                         15      13     13     9      14
Swans                         10      9      8      7      10
Crane                         9       8      7      6      9
Gulls, murres, puffins        1       1      1      1      1
Loons                         2       2      1      1      2
Snipe, godwit, whimbrel       2       2      1      1      2
Grouse, ptarmigan             12      11     10     8      11

Sometimes consumed:

Ducks                         --      1      1      --     --

Geese                         --      --     --     1      --
Swans                         --      --     1      --     --
Crane                         --      --     1      1      --
Gulls, murres, puffins        --      --            --     --
Loons                         --      --     1      1      --
Snipe, godwit, whimbrel       --      --     --     --     --
Grouse, ptarmigan             --      --     --     --     --

Usually not consumed:

Ducks                         --      --     2      --     2
Geese                         --      --     --     3      --
Swans                         --      --     1      3      --
Crane                         --      --     1      3      --
Gulls, murres, puffins        --      --     --     --     --
Loons                         --      --     --     --     --
Snipe, godwit, whimbrel       --      --     1      1      --
Grouse, ptarmigan             --      --     2      4      1

                            Wings   Skin   Fat   Heart

Usually consumed:

Ducks                        12      14    14     11
Geese                        12      13    13     10
Swans                         8      7      9      8
Crane                         8      6      7      7
Gulls, murres, puffins       --      --    --      1
Loons                         1      --    --      1
Snipe, godwit, whimbrel       2      1      2      2
Grouse, ptarmigan            11      10    11      9

Sometimes consumed:

Ducks                        --      --    --      1
Geese                        --      --    --     --
Swans                        --      2     --     --
Crane                        --      2      2     --
Gulls, murres, puffins       --      1      1     --
Loons                        --      --    --     --
Snipe, godwit, whimbrel      --      1     --     --
Grouse, ptarmigan             --     2     1--     1

Usually not consumed:

Ducks                         1      --     3      7
Geese                         1      --    --      3
Swans                         2      2      1      2
Crane                         1      1     --      2
Gulls, murres, puffins        1      1     --     --
Loons                         1      3      1     --
Snipe, godwit, whimbrel      --      --    --     --
Grouse, ptarmigan             1      --    --      2

                            Feet   Gizzard   Liver    Bone
                                                     marrow

Usually consumed:

Ducks                        4       13        9       4
Geese                        3       12        8       4
Swans                        2        9        7       5
Crane                        2        7        6       3
Gulls, murres, puffins       --      --        1       --
Loons                        --      --       --       --
Snipe, godwit, whimbrel      1        1        2       1
Grouse, ptarmigan            3       10        8       3

Sometimes consumed:

Ducks                        --      --       --       1
Geese                        1       --       --       --
Swans                        2       --       --       --
Crane                        --      --       --       --
Gulls, murres, puffins       --      --       --       --
Loons                        --      --       --       --
Snipe, godwit, whimbrel      --      --       --       --
Grouse, ptarmigan            --      --        1       1

Usually not consumed:

Ducks                        1        5        7       7
Geese                        7        1        5       7
Swans                        6        1        3       4
Crane                        6        2        3       5
Gulls, murres, puffins       1        1       --       1
Loons                        1        1        1       1
Snipe, godwit, whimbrel      --      --       --       --
Grouse, ptarmigan            7        2        3       4

                            Blood   Intestines   Pancreas   Stomach

Usually consumed:

Ducks                         2         1           --         1
Geese                         2         1           --         1
Swans                         2         --          --        --
Crane                         2         --          --        --
Gulls, murres, puffins       --         --          --        --
Loons                        --         --          --        --
Snipe, godwit, whimbrel       1         --          --        --
Grouse, ptarmigan             2         1           --        --

Sometimes consumed:

Ducks                         1         --          --        --
Geese                         1         1           --        --
Swans                         1         1           --        --
Crane                         1         1           --        --
Gulls, murres, puffins       --         --          --        --
Loons                        --         --          --        --
Snipe, godwit, whimbrel      --         --          --        --
Grouse, ptarmigan             1         --          "          ~

Usually not consumed:

Ducks                         9         11          10         8
Geese                         6         8           10         9
Swans                         4         6           7          7
Crane                         5         6           7          7
Gulls, murres, puffins        1         1           1          1
Loons                         1         1           1          1
Snipe, godwit, whimbrel      --         --          --        --
Grouse, ptarmigan             5         8           11        10

                            Kidneys   Lungs   Bones   Tongue

Usually consumed:

Ducks                          2       --      --       1
Geese                          2       --      --       1
Swans                          1       --      --       1
Crane                          1       --      --       1
Gulls, murres, puffins        --       --      --       --
Loons                         --       --      --       --
Snipe, godwit, whimbrel       --       --      --       --
Grouse, ptarmigan              2       --      --       1

Sometimes consumed:

Ducks                         --       --      --       --
Geese                         --       --      --       --
Swans                         --       --      --       --
Crane                         --       --      --       --
Gulls, murres, puffins        --       --      --       --
Loons                         --       --      --       --
Snipe, godwit, whimbrel       --       --      --       --
Grouse, ptarmigan

Usually not consumed:

Ducks                         11        2      --       --
Geese                          7        8       2       --
Swans                          5        7       2       --
Crane                          5        7       2       --
Gulls, murres, puffins        --       --      --       --
Loons                         --       --      --       --
Snipe, godwit, whimbrel       --        1      --       --
Grouse, ptarmigan              7       10       2       --

(1) Values are number of key respondents that indicated bird parts as
usually, "sometimes," and "usually not" consumed in this study's
questionnaire (online Appendix SI).

-- : Consumption not indicated by key respondents.

TABLE 3. Mass (g) of wild birds and common cuts in harvest processing
of birds harvested in south-central Alaska and the Alaska
Peninsula in September--October 2015 and September 2016.

Species                 Live mass   Carcass     Heart     Gizzard
                                      (1)

American                   922        563         6         22
Wigeon Anas                919        492         8         25
americana                  738        423         8         31
                           901        533         7         31
                           766        437         7         43
                           566        307         3         35
                           777        426         7         53
                           598        349         6         39

Mallard A.                1157        721        11         45
platyrhynchos             1080        679         9         54
                          1307        814        11         57
                          1055        609         9         47
                          1149        669         9         49

Northern Pintail          1167        752         9         32
A. acuta                   848        527         8         44
                           948        627         8         28
                           686        424         8         30
Green-winged               386        243         2          3
Teal A. crecca

Black Brant               1952       1153        16         89
Branta bernicla

Canada/Cackling           1485        882        13         82
Goose Branta spp.         1732       1000        14         84
                          1677       1010        13         64
                          2549       1491        17         110
                          1685       1097        12         75
                          1471        851        10         71
                          1602        919         9         87
                          2493       1494        16         129

Willow Ptarmigan           590        378         7         14
Lagopus lagopus            611        372         7         16

Mean proportion           100%        60%        1%         4%
of live mass

Range of proportions                54%-66%   0.5%-1.2%    l%-7%
of live mass

Species                   Total        Breast       Whole
                        edible (2)   fillets (3)   legs (4)

American                   591           213          84
Wigeon Anas                525           211          71
americana                  462           160          56
                           571           193          71
                           487           167          57
                           345           113          39
                           486           162          58
                           394           119          45

Mallard A.                 777           285         102
platyrhynchos              742           267          95
                           882           305         141
                           665           214          92
                           727           251         106

Northern Pintail           793           256          95
A. acuta                   579           206          76
                           663           228          97
                           462           164          59
Green-winged               248           95           45
Teal A. crecca

Black Brant                1258          412         202
Branta bernicla

Canada/Cackling            977           301         179
Goose Branta spp.          1098          336         186
                           1087          342         204
                           1618          507         312
                           1184          350         220
                           932           287         155
                           1015          305         186
                           1639          452         287

Willow Ptarmigan           399           166          69
Lagopus lagopus            395           163          71

Mean proportion            65%           22%         10%
of live mass

Range of proportions     56%-70%       18%-28%      7%-13%
of live mass

(1) Bone-in, skin-on. Feathers, wing tips, feet,
head, and viscera removed (see Methods).

(2) Edible mass included the carcass, heart, and gizzard.

(3) Boneless, skin-on, outer and inner fillets, right and left sides.

(4) Bone-in, skin-on thigh and drumstick, right and left sides.

TABLE 4. Conversion factors to estimate food production in subsistence
harvest of birds and eggs in Alaska. Source for body and egg mass was
Rodewald (2015) unless otherwise noted. To calculate CF for
multi-species or multi-population categories, we weighted body and
egg mass by population size whenever possible. Asterisks indicate
species or categories for which further information is available in
online Appendix S2- continued:

                                           Bird

                                 Body     Relative   CFn-m (2)
                               mass (g)    mass1        (g)

American Wigeon                  735                    478
Anas americana

Gadwall Anas strepera            859                    558

Teal (unidentified) *            328                    213

  Green-winged Teal              328        12%         213
  A. crecca

  Blue-winged Teal A.            371                    241
  discors

Mallard A. platyrhynchos         1122                   729

Northern Pintail A. acuta        820                    533

Northern Shoveler                603                    392
A. clypeata

Black Scoter                     1052                   684
Melanitta nigra

Surf Scoter M.                   1022                   664
perspicillata

White-winged                     1825                  1186
Scoter M. fusca

Bufflehead                       397                    258
Bucephala albeola

Goldeneye                        887                    577
(unidentified) *

  Common Goldeneye               863         5%         561
  Bucephala clangula

  Barrow's Goldeneye             910                    592
  B. islandica

Canvasback Aythya                1210                   787
valisineria

Scaup (unidentified) *           943                    613

  Greater Scaup Aythya           943                    613
  marila

  Lesser Scaup A. affinis        751        20%         488

Common Eider Somateria           2288                  1487
moliissima v-nigrum *

King Eider 5.                    1570                  1021
spectabilis *

Spectacled Eider S.              1466                   953
fischeri *

Steller's Eider                  833                    541
Polysticta stelleri *

Harlequin Duck                   588                    382
Histrionicus histrionicus

Long-tailed Duck                 814                    529
Clangula hyemalis *

Merganser                        1209                   786
(unidentified) *

  Common Merganser               1472                   957
  Mergus merganser

  Red-breasted Merganser         946        36%         615
  M. serrator

Black Brant                      1321                   859
Branta bernicla

Canada/Cackling Goose            1972                  1282
(unidentified) *

  Lesser Canada Goose            3060        9%        1989
  Branta canadensis
  parvipes

  Dusky Canada Goose B.          2936       13%        1908
  c. occidentalis

  Vancouver Canada Goose         3366                  2188
  B. c. fulva

  Taverner's Cackling            2514       25%        1634
  Goose B. hutchinsii
  taverneri

  Aleutian Cackling Goose        1825       46%        1186
  B. h. leucopareia

  Cackling Cackling Goose        1429       58%         929
  B. h. minima

Greater White-fronted            2218                  1442
Goose Anser albifrons *

  Tundra Greater White-          2420        4%        1573
  fronted Goose A. a.
  gambelli

  Pacific Greater White-         2015       20%        1310
  fronted Goose A. a.
  sponsa

  Tule Greater White-            2510                  1632
  fronted Goose A.
  a. elgasi

Emperor Goose                    2148                  1396
Chen canagica

Lesser Snow Goose                1955                  1271
C. caerulescens

Swan (unidentified) *            7662                  4980

  Tundra Swan Cygnus             7111       34%        4622
  columbianus

    Eastern population           7014        3%        4559

    Western population           7207                  4685

  Trumpeter Swan C.             10 700                 6955
  buccinator

Sandhill Crane                   3763                  2446
Grus canadensis *

  G. c. canadensis               3705       17%        2408

  G. c. rowani                   4455                  2896

Ptarmigan                        542                    352
(unidentified) *

  White-tailed Ptarmigan         355        34%         231
  Lagopus leucura *

  Rock Ptarmigan L. muta *       420        23%         273

  Willow Ptarmigan L.            542                    352
  lagopus *

Grouse                           635                    413
(unidentified) *

  Ruffed Grouse Bonasa           591        41%         384
  umbellus *

  Spruce Grouse                  595        41%         387
  Falcipennis canadensis *

  Sooty Grouse                   1004                   653
  Dendragapus fuliginosus

  Sharp-tailed Grouse            720                    468
  Tympanuchusphasianellus *

Short-tailed Shearwater          527                    343
Puffinus tenuirostris *

Cormorant                        1985                  1290
(unidentified) *

  Double-crested Cormorant       2330                  1515
  Phalacrocorax auritus

  Red-faced Cormorant            2138        8%        1390
  P. urile

  Pelagic Cormorant              1868       20%        1214
  P. pelagicus

Bonaparte's/Sabine's             185                    120
Gull (unidentified) *

  Bonaparte's Gull               180         5%         117
  Chroicocephalus
  philadelphia

  Sabine's Gull Xema sabini      190                    124

Mew Gull Larus canus *           389                    253

Large gull                       1199                   779
(unidentified) *

  Herring Gull L.                1085       24%         705
  argentatus

  Glaucous-winged Gull           1077       25%         700
  L. glaucescens

  Glaucous Gull L.               1434                   932
  hyperboreus

Black-legged Kittiwake           429                    279
Rissa tridactyla

Red-legged Kittiwake             377                    245
R. brevirostris

Tern (unidentified) *            112                    73

  Arctic Tern Sterna             112         7%         73
  paradisea

  Aleutian Tern                  120                    78
  Onychoprion aleuticus

  Murre (unidentified) *         965                    627

  Common Murre Uria aalge        966                    628

  Thick-billed Murre V.          963        <1%         626
  lomvia

Guillemot                        505                    328
(unidentified) *

  Black Guillemot Cepphus        378        25%         246
  grille

  Pigeon Guillemot               507                    330
  C. columba

Auklet (unidentified) *          171                    111
  Least Auklet Aethia             84        83%         55
  pusilla

  Crested Auklet A.              255        50%         166
  cristatella

  Least/Crested Auklet           170                    111

  Cassin's Auklet                185        63%         120
  Ptychoramphus aleuticus

  Parakeet Auklet                262        48%         170
  Aethiapsittacula

  Whiskered Auklet               120        76%         78
  A. pygmaea

  Rhinoceros Auklet              507                    330
  Cerorhinca monocerata

Puffin (unidentified) *          707                    460

  Horned Puffin                  537        31%         349
  Fratercula corniculata

  Tufted Puffin F.               774                    503
  cirrhata

Black Oystercatcher              535                    348
Haematopus bachmani

Whimbrel/Curlew                  399                    259
(unidentified) *

  Whimbrel Numenius              391        10%         254
  phaeopus rufiventris

  Bristle-thighed Curlew         433                    281
  N. tahitiensis

Godwit (unidentified) *          421                    274

  Bar-tailed Godwit Limosa       464                    302
  tapponica baueri

  Hudsonian Godwit L.            241        48%         157
  haemastica

  Marbled Godwit L. fedoa        368        21%         239
  beringiae

Golden/Black-bellie              162                    105
Plover (unidentified) *

  American Golden Plover         133        39%         86
  Pluvialis dominica

  Pacific Golden Plover          148        32%         96
  P. fulva

  Black-bellied Plover           219                    142
  P. squatarola squatarola

Turnstone                        122                    79
(unidentified) *

  Ruddy Turnstone Arenaria       107        14%         70
  interpres interpres

  Black Turnstone A.             125                    81
  melanocephata

Phalarope                         42                    27
(unidentified) *

  Red-necked Phalarope            35        33%         23
  Phalaropus lobatus

  Red Phalarope P.                53                    34
  fulicarius

Small shorebird                   37                    24
(unidentified) *

  Western Sandpiper               28        60%         18
  Calidris mauri

  Dunlin C. alpina                65                    42
  arcticola

Wilson's Snipe                    99                    64
Gallinago delicata

Loon (unidentified) *            2513                  1633

  Red-throated Loon              1759       65%        1143
  Gavia stellata

  Arctic Loon G. arctica         3101       39%        2016

  Pacific Loon G. pacifica       2232       56%        1451

  Common Loon G. immer           5015        1%        3260

  Yellow-billed Loon G.          5056                  3286
  adamsii

Grebe (unidentified) *           799                    519

  Horned Grebe Podiceps          405        66%         263
  auritus

  Red-necked Grebe P.            1192                   775
  griseana

Snowy Owl Bubo                   1873                  1217
scandiacus

                                        Bird              Egg

                               Body mass   CFn-m (2)     CFn-m
                                (pound)     (pound)    = egg mass
                                                        (3) (g)

American Wigeon                  1.62        1.05          43
Anas americana

Gadwall Anas strepera            1.89        1.23          43

Teal (unidentified) *            0.72        0.47          28

  Green-winged Teal              0.72        0.47          --
  A. crecca

  Blue-winged Teal A.            0.82        0.53          28
  discors

Mallard A. platyrhynchos         2.47        1.61          52

Northern Pintail A. acuta        1.81        1.18          43

Northern Shoveler                1.33        0.86          38
A. clypeata

Black Scoter                     2.32        1.51          67
Melanitta nigra

Surf Scoter M.                   2.25        1.46          78
perspicillata

White-winged                     4.02        2.61          82
Scoter M. fusca

Bufflehead                       0.88        0.57          37
Bucephala albeola

Goldeneye                        1.96        1.27          66
(unidentified) *

  Common Goldeneye               1.90        1.24          64
  Bucephala clangula

  Barrow's Goldeneye             2.01        1.31          68
  B. islandica

Canvasback Aythya                2.67        1.74          68
valisineria

Scaup (unidentified) *           2.08        1.35          63

  Greater Scaup Aythya           2.08        1.35          63
  marila

  Lesser Scaup A. affinis        1.66        1.08          48

Common Eider Somateria           5.04        3.28         101
moliissima v-nigrum *

King Eider 5.                    3.46        2.25          69
spectabilis *

Spectacled Eider S.              3.23        2.10          71
fischeri *

Steller's Eider                  1.84        1.20          55
Polysticta stelleri *

Harlequin Duck                   1.30        0.85          53
Histrionicus histrionicus

Long-tailed Duck                 1.79        1.16          43
Clangula hyemalis *

Merganser                        2.67        1.74          69
(unidentified) *

  Common Merganser               3.25        2.11          70
  Mergus merganser

  Red-breasted Merganser         2.09        1.36          68
  M. serrator

Black Brant                      2.91        1.89         100
Branta bernicla

Canada/Cackling Goose            4.35        2.83         113
(unidentified) *

  Lesser Canada Goose            6.75        4.39          --
  Branta canadensis
  parvipes

  Dusky Canada Goose B.          6.47        4.21          --
  c. occidentalis

  Vancouver Canada Goose         7.42        4.82          --
  B. c. fulva

  Taverner's Cackling            5.54        3.60         124
  Goose B. hutchinsii
  taverneri

  Aleutian Cackling Goose        4.02        2.61          --
  B. h. leucopareia

  Cackling Cackling Goose        3.15        2.05         101
  B. h. minima

Greater White-fronted            4.89        3.18         129
Goose Anser albifrons *

  Tundra Greater White-          5.34        3.47         129
  fronted Goose A. a.
  gambelli

  Pacific Greater White-         4.44        2.89         128
  fronted Goose A. a.
  sponsa

  Tule Greater White-            5.53        3.59          --
  fronted Goose A.
  a. elgasi

Emperor Goose                    4.74        3.08         122
Chen canagica

Lesser Snow Goose                4.31        2.80         122
C. caerulescens

Swan (unidentified) *            16.89       10.98        287

  Tundra Swan Cygnus             15.68       10.19        273
  columbianus

    Eastern population           15.46       10.05        273

    Western population           15.89       10.33         --

  Trumpeter Swan C.              23.59       15.33        363
  buccinator

Sandhill Crane                   8.30        5.40         151
Grus canadensis *

  G. c. canadensis               8.17        5.31         150

  G. c. rowani                   9.82        6.38         161

Ptarmigan                        1.19        0.77          22
(unidentified) *

  White-tailed Ptarmigan         0.78        0.51          19
  Lagopus leucura *

  Rock Ptarmigan L. muta *       0.93        0.60          21

  Willow Ptarmigan L.            1.19        0.77          22
  lagopus *

Grouse                           1.40        0.91          21
(unidentified) *

  Ruffed Grouse Bonasa           1.30        0.85          20
  umbellus *

  Spruce Grouse                  1.31        0.85          22
  Falcipennis canadensis *

  Sooty Grouse                   2.21        1.44          36
  Dendragapus fuliginosus

  Sharp-tailed Grouse            1.59        1.03          --
  Tympanuchusphasianellus *

Short-tailed Shearwater          1.16        0.75          NA
Puffinus tenuirostris *

Cormorant                        4.38        2.85          45
(unidentified) *

  Double-crested Cormorant       5.14        3.34          47
  Phalacrocorax auritus

  Red-faced Cormorant            4.71        3.06          --
  P. urile

  Pelagic Cormorant              4.12        2.68          45
  P. pelagicus

Bonaparte's/Sabine's             0.41        0.27          26
Gull (unidentified) *

  Bonaparte's Gull               0.40        0.26          --
  Chroicocephalus
  philadelphia

  Sabine's Gull Xema sabini      0.42        0.27          26

Mew Gull Larus canus *           0.86        0.56          52

Large gull                       2.64        1.72          97
(unidentified) *

  Herring Gull L.                2.39        1.55          95
  argentatus

  Glaucous-winged Gull           2.37        1.54          92
  L. glaucescens

  Glaucous Gull L.               3.16        2.05         105
  hyperboreus

Black-legged Kittiwake           0.95        0.62          52
Rissa tridactyla

Red-legged Kittiwake             0.83        0.54          49
R. brevirostris

Tern (unidentified) *            0.25        0.16          19

  Arctic Tern Sterna             0.25        0.16          19
  paradisea

  Aleutian Tern                  0.26        0.17          20
  Onychoprion aleuticus

  Murre (unidentified) *         2.13        1.38         105

  Common Murre Uria aalge        2.13        1.38         106

  Thick-billed Murre V.          2.12        1.38         103
  lomvia

Guillemot                        1.11        0.72          54
(unidentified) *

  Black Guillemot Cepphus        0.83        0.54          54
  grille

  Pigeon Guillemot               1.12        0.73          54
  C. columba

Auklet (unidentified) *          0.38        0.25          25
  Least Auklet Aethia            0.19        0.12          18
  pusilla

  Crested Auklet A.              0.56        0.36          36
  cristatella

  Least/Crested Auklet           0.37        0.24          27

  Cassin's Auklet                0.41        0.27          25
  Ptychoramphus aleuticus

  Parakeet Auklet                0.58        0.38          34
  Aethiapsittacula

  Whiskered Auklet               0.26        0.17          --
  A. pygmaea

  Rhinoceros Auklet              1.12        0.73          78
  Cerorhinca monocerata

Puffin (unidentified) *          1.56        1.01          87

  Horned Puffin                  1.18        0.77          76
  Fratercula corniculata

  Tufted Puffin F.               1.71        1.11          91
  cirrhata

Black Oystercatcher              1.18        0.77          46
Haematopus bachmani

Whimbrel/Curlew                  0.88        0.57          50
(unidentified) *

  Whimbrel Numenius              0.86        0.56          49
  phaeopus rufiventris

  Bristle-thighed Curlew         0.95        0.62          56
  N. tahitiensis

Godwit (unidentified) *          0.93        0.60          36

  Bar-tailed Godwit Limosa       1.02        0.66          37
  tapponica baueri

  Hudsonian Godwit L.            0.53        0.34          32
  haemastica

  Marbled Godwit L. fedoa        0.81        0.53          48
  beringiae

Golden/Black-bellie              0.36        0.23          30
Plover (unidentified) *

  American Golden Plover         0.29        0.19          27
  Pluvialis dominica

  Pacific Golden Plover          0.33        0.21          27
  P. fulva

  Black-bellied Plover           0.48        0.31          35
  P. squatarola squatarola

Turnstone                        0.27        0.18          18
(unidentified) *

  Ruddy Turnstone Arenaria       0.24        0.16          17
  interpres interpres

  Black Turnstone A.             0.28        0.18          18
  melanocephata

Phalarope                        0.09        0.06          7
(unidentified) *

  Red-necked Phalarope           0.08        0.05          6
  Phalaropus lobatus

  Red Phalarope P.               0.12        0.08          8
  fulicarius

Small shorebird                  0.08        0.05          8
(unidentified) *

  Western Sandpiper              0.06        0.04          7
  Calidris mauri

  Dunlin C. alpina               0.14        0.09          11
  arcticola

Wilson's Snipe                   0.22        0.14          15
Gallinago delicata

Loon (unidentified) *            5.54        3.60         103

  Red-throated Loon              3.88        2.52          77
  Gavia stellata

  Arctic Loon G. arctica         6.84        4.45          --

  Pacific Loon G. pacifica       4.92        3.20         101

  Common Loon G. immer           11.06       7.19         143

  Yellow-billed Loon G.          11.15       7.25         154
  adamsii

Grebe (unidentified) *           1.76        1.14          30

  Horned Grebe Podiceps          0.89        0.58          21
  auritus

  Red-necked Grebe P.            2.63        1.71          38
  griseana

Snowy Owl Bubo                   4.13        2.68          57
scandiacus

                                                Egg

                               CFv-n (4)   CFn-m = egg     CFv-n (4)
                               (eggs/L)     mass (3)     (eggs/gallon)
                                             (pound)

American Wigeon                   9.8         0.095          37.5
Anas americana

Gadwall Anas strepera             9.8         0.095          37.5

Teal (unidentified) *            15.1         0.062          57.4

  Green-winged Teal               --           --             --
  A. crecca

  Blue-winged Teal A.            15.1         0.062          57.4
  discors

Mallard A. platyrhynchos          8.1         0.115          30.9

Northern Pintail A. acuta         9.8         0.095          37.5

Northern Shoveler                11.1         0.084          42.4
A. clypeata

Black Scoter                      6.3         0.148          24.0
Melanitta nigra

Surf Scoter M.                    5.4         0.172          20.7
perspicillata

White-winged                      5.1         0.181          19.7
Scoter M. fusca

Bufflehead                       11.4         0.082          43.4
Bucephala albeola

Goldeneye                         6.4         0.146          24.4
(unidentified) *

  Common Goldeneye                6.6         0.141          25.2
  Bucephala clangula

  Barrow's Goldeneye              6.2         0.150          23.7
  B. islandica

Canvasback Aythya                 6.2         0.150          23.7
valisineria

Scaup (unidentified) *            6.7         0.139          25.6

  Greater Scaup Aythya            6.7         0.139          25.6
  marila

  Lesser Scaup A. affinis         8.8         0.106          33.6

Common Eider Somateria            4.2         0.223          16.0
moliissima v-nigrum *

King Eider 5.                     6.1         0.152          23.4
spectabilis *

Spectacled Eider S.               5.9         0.157          22.7
fischeri *

Steller's Eider                   7.7         0.121          29.4
Polysticta stelleri *

Harlequin Duck                    8.0         0.117          30.4
Histrionicus histrionicus

Long-tailed Duck                  9.8         0.095          37.5
Clangula hyemalis *

Merganser                         6.1         0.152          23.4
(unidentified) *

  Common Merganser                6.0         0.154          23.1
  Mergus merganser

  Red-breasted Merganser          6.2         0.150          23.7
  M. serrator

Black Brant                       4.2         0.220          16.2
Branta bernicla

Canada/Cackling Goose             3.7         0.249          14.3
(unidentified) *

  Lesser Canada Goose             --           --             --
  Branta canadensis
  parvipes

  Dusky Canada Goose B.           --           --             --
  c. occidentalis

  Vancouver Canada Goose          --           --             --
  B. c. fulva

  Taverner's Cackling             3.4         0.273          13.0
  Goose B. hutchinsii
  taverneri

  Aleutian Cackling Goose         --           --             --
  B. h. leucopareia

  Cackling Cackling Goose         4.2         0.223          16.0
  B. h. minima

Greater White-fronted             3.3         0.284          12.5
Goose Anser albifrons *

  Tundra Greater White-           3.3         0.284          12.5
  fronted Goose A. a.
  gambelli

  Pacific Greater White-          3.3         0.282          12.6
  fronted Goose A. a.
  sponsa

  Tule Greater White-             --           --             --
  fronted Goose A.
  a. elgasi

Emperor Goose                     3.5         0.269          13.2
Chen canagica

Lesser Snow Goose                 3.5         0.269          13.2
C. caerulescens

Swan (unidentified) *             1.5         0.633           5.6

  Tundra Swan Cygnus              1.5         0.602           5.9
  columbianus

    Eastern population            1.5         0.602           5.9

    Western population            --           --             --

  Trumpeter Swan C.               1.2         0.800           4.4
  buccinator

Sandhill Crane                    2.8         0.333          10.7
Grus canadensis *

  G. c. canadensis                2.8         0.331          10.7

  G. c. rowani                    2.6         0.355          10.0

Ptarmigan                        19.2         0.049          72.6
(unidentified) *

  White-tailed Ptarmigan         22.2         0.042          84.7
  Lagopus leucura *

  Rock Ptarmigan L. muta *       20.1         0.046          77.4

  Willow Ptarmigan L.            19.2         0.049          72.6
  lagopus *

Grouse                           20.1         0.046          77.4
(unidentified) *

  Ruffed Grouse Bonasa           21.1         0.044          80.9
  umbellus *

  Spruce Grouse                  19.2         0.049          72.6
  Falcipennis canadensis *

  Sooty Grouse                   11.7         0.079          45.0
  Dendragapus fuliginosus

  Sharp-tailed Grouse             --           --             --
  Tympanuchusphasianellus *

Short-tailed Shearwater           NA           NA             NA
Puffinus tenuirostris *

Cormorant                         9.4         0.099          35.9
(unidentified) *

  Double-crested Cormorant        9.0         0.104          34.2
  Phalacrocorax auritus

  Red-faced Cormorant             --           --             --
  P. urile

  Pelagic Cormorant               9.4         0.099          35.9
  P. pelagicus

Bonaparte's/Sabine's             16.2         0.057          62.4
Gull (unidentified) *

  Bonaparte's Gull                --           --             --
  Chroicocephalus
  philadelphia

  Sabine's Gull Xema sabini      16.2         0.057          62.4

Mew Gull Larus canus *            8.1         0.115          30.9

Large gull                        4.3         0.214          16.6
(unidentified) *

  Herring Gull L.                 4.4         0.209          17.0
  argentatus

  Glaucous-winged Gull            4.6         0.203          17.5
  L. glaucescens

  Glaucous Gull L.                4.0         0.231          15.4
  hyperboreus

Black-legged Kittiwake            8.1         0.115          30.9
Rissa tridactyla

Red-legged Kittiwake              8.6         0.108          32.9
R. brevirostris

Tern (unidentified) *            22.2         0.042          84.7

  Arctic Tern Sterna             22.2         0.042          84.7
  paradisea

  Aleutian Tern                  21.1         0.044          80.9
  Onychoprion aleuticus

  Murre (unidentified) *          4.0         0.231          15.4

  Common Murre Uria aalge         4.0         0.234          15.2

  Thick-billed Murre V.           4.1         0.227          15.7
  lomvia

Guillemot                         7.8         0.119          29.9
(unidentified) *

  Black Guillemot Cepphus         7.8         0.119          29.9
  grille

  Pigeon Guillemot                7.8         0.119          29.9
  C. columba

Auklet (unidentified) *          16.9         0.055          64.7
  Least Auklet Aethia            23.4         0.040          89.0
  pusilla

  Crested Auklet A.              11.7         0.079          45.0
  cristatella

  Least/Crested Auklet           15.6         0.060          59.3

  Cassin's Auklet                16.9         0.055          64.7
  Ptychoramphus aleuticus

  Parakeet Auklet                12.4         0.075          47.4
  Aethiapsittacula

  Whiskered Auklet                --           --             --
  A. pygmaea

  Rhinoceros Auklet               5.4         0.172          20.7
  Cerorhinca monocerata

Puffin (unidentified) *           4.8         0.192          18.5

  Horned Puffin                   5.6         0.168          21.2
  Fratercula corniculata

  Tufted Puffin F.                4.6         0.201          17.7
  cirrhata

Black Oystercatcher               9.2         0.101          35.2
Haematopus bachmani

Whimbrel/Curlew                   8.4         0.110          32.3
(unidentified) *

  Whimbrel Numenius               8.6         0.108          32.9
  phaeopus rufiventris

  Bristle-thighed Curlew          7.5         0.123          28.9
  N. tahitiensis

Godwit (unidentified) *          11.7         0.079          45.0

  Bar-tailed Godwit Limosa       11.4         0.082          43.4
  tapponica baueri

  Hudsonian Godwit L.            13.2         0.071          50.1
  haemastica

  Marbled Godwit L. fedoa         8.8         0.106          33.6
  beringiae

Golden/Black-bellie              14.1         0.066          53.9
Plover (unidentified) *

  American Golden Plover         15.6         0.060          59.3
  Pluvialis dominica

  Pacific Golden Plover          15.6         0.060          59.3
  P. fulva

  Black-bellied Plover           12.1         0.077          46.2
  P. squatarola squatarola

Turnstone                        23.4         0.040          89.0
(unidentified) *

  Ruddy Turnstone Arenaria       24.8         0.037          96.2
  interpres interpres

  Black Turnstone A.             23.4         0.040          89.0
  melanocephata

Phalarope                        60.3         0.015          237.2
(unidentified) *

  Red-necked Phalarope           70.3         0.013          273.7
  Phalaropus lobatus

  Red Phalarope P.               52.7         0.018          197.7
  fulicarius

Small shorebird                  52.7         0.018          197.7
(unidentified) *

  Western Sandpiper              60.3         0.015          237.2
  Calidris mauri

  Dunlin C. alpina               38.3         0.024          148.3
  arcticola

Wilson's Snipe                   28.1         0.033          107.8
Gallinago delicata

Loon (unidentified) *             4.1         0.227          15.7

  Red-throated Loon               5.5         0.170          20.9
  Gavia stellata

  Arctic Loon G. arctica          --           --             --

  Pacific Loon G. pacifica        4.2         0.223          16.0

  Common Loon G. immer            2.9         0.315          11.3

  Yellow-billed Loon G.           2.7         0.340          10.5
  adamsii

Grebe (unidentified) *           14.1         0.066          53.9

  Horned Grebe Podiceps          20.1         0.046          77.4
  auritus

  Red-necked Grebe P.            11.1         0.084          42.4
  griseana

Snowy Owl Bubo                    7.4         0.126          28.2
scandiacus

(1) Relative mass is the percent difference in body mass (g)
between a given species and the heaviest species, subspecies, or
population within the same category. That is, relative mass = [1 -
(focal item heaviest item in category)] x 100.

(2) Bird CFn-m (number-to-mass conversion factor) = recovery rate
(65%) x body mass.

(3) Egg CFn-m (number-to-mass conversion factor) = recovery rate
(100%) x egg mass.

(4) Egg CFv-n: volume-to-number conversion factor.

Number of eggs/gallon: CFv-n = (35.3 x 0.8) * (0.126 / mass of wild
bird egg, in pounds).

Number of eggs/L: CFv-n = (9.3 x 0.8) x (57.0 / mass of wild bird
egg, in grams).

--: Data unavailable.
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Article Details
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Author:Naves, Liliana C.; Fall, James A.
Publication:Arctic
Article Type:Report
Geographic Code:1U9AK
Date:Mar 1, 2017
Words:12954
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