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"‘5‘ "hu-SI’” ‘1“.‘39 «.95 . .- 5...“ “gs ‘ ‘ ' ' .— '- - '1'. ‘ .- '4 O '.I."'I‘ ‘ - .‘n. .I. um «’55. . 52 -- - 1”."15- 51,5.“ 5‘ 3‘3? ~~_.- 9'. ..uM. ‘ L: 2-1 .w.‘ "5%- llllllllllllllllll “2818 ( {a «‘1 r; , This is to certify that the thesis entitled Predictive Models for the Fall Flight of Selected Waterfowl Species in Michigan presented by Karen Tay Cleveland has been accepted towards fulfillment of the requirements for M.S. Jegree in Fisheries & Wildlife 3663752 ‘4) omzwtiw Major professor Date ’L/ 8/97 0.7639 MS U is an Affirmative Action/Equal Opportunity Institution LIERARY Michigan State University PLACE ll RETURN BOXtoromavothbdnckomtmmywm TO AVOID FINES Mum anorbdondatoduo. DATE DUE DATE DUE DATE DUE u.“ i——‘| - - IIEI III I I MSU icAnAfiinn‘iwAanEqud Opportunity intuition 1 PREDICTIVE MODELS FOR THE FALL FLIGHT OF SELECTED WATERFOWL SPECIES IN MICHIGAN BY Karen Tay Cleveland A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Fisheries and Wildlife 1994 ABSTRACT PREDICTIVE MODELS FOR THE FALL FLIGHT OF SELECTED WATERFOWL SPECIES IN MICHIGAN BY Karen Tay Cleveland Michigan has traditionally lacked accurate estimates of its populations of breeding waterfowl and fall flight. Models of the production of mallard, black duck, Canada goose, and wood duck were constructed using data from the Michigan Breeding Waterfowl Survey (MBWS). The resultant models take the form of a distribution of values of young produced per adult, and point estimates of young produced to flight stage, adults at migration, newly fledged birds at migration, and total fall flight. Production rates were assessed for their impact on population size. The mallard and black duck models accurately predict the number of young produced annually. The wood duck model overestimates production by loo-300%. The Canada goose model incorporates age specific production and underestimates the fall flight in Michigan. Recommendations to improve the NEWS were made including changes in stratification, modifications of the calculation of visibility correction factors, and reduction of observer bias. ACKNOWLEDGMENTS I am indebted to Dr. Scott Winterstein, my major professor, for his guidance throughout the duration of my study at Michigan State University and for his capable editing of this thesis. Also, I would like to thank Dr. Harold Prince and Dr. Donald Beaver, committee members, for their advice in refining arguments and structure in the final draft. Gratitude is extended to the personnel of the Michigan Department of Natural Resources, specifically to Mr. Gerald Martz and Mr. Nate Levitte for their assistance in analyzing the Michigan Breeding Waterfowl Survey and Mr. Al Stewart and Mr. Barry Loper for their insights into waterfowl banding efforts in Michigan. Also, the personnel of the Patuxent Wildlife Research Center are thanked for providing information on federal waterfowl survey techniques and results. Finally, I would like to thank my parents, Dick and Johanna, for providing financial, emotional, and technical support to my endeavors. iii TABLE OF CONTENTS Page List of Tables ............................................. v List of Figures ........................................... vi Introduction ............................................... 1 Michigan Breeding Waterfowl Survey .................... 1 Waterfowl Modelling ................................... 3 Objectives ................................................. 7 Methods .................................................... 8 Michigan Breeding Waterfowl Survey and Visibility Correction Factors ........................ 8 Species of Interest .................................. 12 Model Framework ...................................... 14 Model Development .................................... 15 Results & Discussion ...................................... 21 Mallard .............................................. 21 Black duck ........................................... 29 Canada goose ......................................... 37 Wood duck ............................................ 45 General Summary ...................................... 52 Recommendations for the Michigan Breeding Waterfowl Survey ............................... 54 Strata Designation ................................... 54 Visibility Correction Factor Calculation ............. 56 Observer Bias ........................................ 60 Sample Size .......................................... 64 Glossary .................................................. 68 Appendix .................................................. 70 Literature Cited .......................................... 85 iv Table Table Table Table Table Table Table Table LIST OF TABLES gage . Mallard model input values ....................... 22 . Mallard model output values ...................... 27 Black duck model input values .................... 30 Black duck model output values ................... 35 Canada goose model input values .................. 38 Canada goose model output values ................. 43 . Wood duck model input values ..................... 46 . Wood duck model output values .................... 50 Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure 10. 11. 12. LIST OF FIGURES Flowchart based waterfowl model (Johnson et a1. 1986) ...................... . Mechanistic model equations to estimate fall flight by species (Walters et a1. 1974) ...................... Michigan Breeding Waterfowl Survey transect routes ............................ Model development flowchart ................ Distribution of mallard production rate values ................................ Projected population values for mallards in Michigan ....................... Distribution of black duck production rate values ................................ Projected population values for black ducks in Michigan .......................... Distribution of Canada goose optimal production rate values ..................... Distribution of Canada goose realized production rate values .................... Distribution of wood duck production rate values ............................... Projected population values for wood ducks in Michigan ........................ vi Page ...... 5 ...... 6 ...... 9 ..... l6 ..... 24 ..... 25 ..... 31 ..... 32 ..... 39 ..... 41 ..... 47 ...... 48 Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure l3. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. Stratum variance across changing survey sample sizes for the Southern Lower Peninsula ...................................... 65 Stratum variance across changing survey sample sizes for the Northern Lower Peninsula ...................................... 66 Stratum variance across changing survey sample sizes for the Upper Peninsula ...................................... 66 Production rate distribution calculated from mallard literature values ................. 70 Result of 10,000 random draws from the constructed distribution of mallard eggs per adult values ............................... 71 Result of 10,000 random draws from the constructed distribution of mallard nest success values ................................. 72 Result of 10,000 random draws from the constructed distribution of mallard survival from laying to flight values .......... 73 Production rate distribution calculated from black duck literature values .............. 74 Result of 10,000 random draws from the constructed distribution of black duck eggs per adult values .......................... 75 Result of 10,000 random draws from the constructed distribution of black duck nest success values ............................ 76 Result of 10,000 random draws from the constructed distribution of black duck egg success in successful nests values ......... 77 Result of 10,000 random draws from the constructed distribution of black duck survival from hatching to flight values ........ 78 vii Figure Figure Figure Figure Figure Figure 25. 26. 27. 28. 29. 30. Result of 10,000 random draws from the constructed distribution of Canada goose eggs per adult values .......................... 79 Result of 10,000 random draws from the constructed distribution of Canada goose survival rate of eggs to hatching values ....... 80 Result of 10,000 random draws from the constructed distribution of Canada goose survival rate of goslings to flight values ..... 81 Result of 10,000 random draws from the constructed distribution of wood duck clutch size values ............................. 82 Result of 10,000 random draws from the constructed distribution of wood duck survival from laying to hatching values ........ 83 Result of 10,000 random draws from the constructed distribution of wood duck survival from hatching to flight values ........ 84 viii INTRODUCTION In recent years, Michigan has gathered more attention as a source of waterfowl production. This is due in great part to the declining populations of breeding waterfowl in the prairie pothole region of North America, which produces about 50% of the ducks in North America, though it contains only 10% of the continental breeding range (Klett et al. 1988). It has been suggested that this decline may be due to more intensive agricultural practices or hunting pressure (Johnson and Shaffer 1987, Nudds and Cole 1991). Michigan Breeding Waterfowl Survey: During the late 1940's and early 1950's, the United States Fish and Wildlife Service developed and implemented the aerial Waterfowl Breedinginpulation and Habitat Survey (WBPHS), which is conducted in the spring to estimate the number of breeding birds and the abundance of suitable breeding habitat, mostly in the form of potholes. This was done in large part to aid in the determination of 2 appropriate management options to comply with the Migratory Bird Treaty Act of 1918. The aerial Waterfowl Production and Habitat Survey (WPHS), established experimentally in 1950 and operational in 1956, has also been used to estimate population sizes and production rates. It is flown in July to estimate the total production of waterfowl and the number of late nesting birds. Since the mid-1950's, both of these surveys have been conducted annually. Beginning in 1959, air-ground comparisons were made in the counts to provide visibility correction factors for all species counted. These adjustments only apply to the WBPHS, however, as problems caused the discontinuance of air—ground sampling on the WPHS. Current efforts in the United States and Canada account for the monitoring of nearly 1.4 million square miles every year (USFWS 1987) In 1991, efforts to estimate waterfowl production and the size of the breeding population in Michigan were instituted. The initial aim of the project was to estimate the size of the adult spring population using visibility correction factors measured in Ontario and a survey design which was designed for use in the prairie pothole region. It has developed into a program to assess the number of spring breeders and estimate the expected fall flight. The Michigan Department of Natural Resources (MDNR) conducts annual 3 aerial surveys of the entire state during the spring which consist of fixed-wing and helicopter flights to determine the number of breeding waterfowl. The fixed-wing flights establish a rough count which is modified by counts from the helicopter flights. Due to differences in land use and ground cover in the southern Lower Peninsula, northern Lower Peninsula and Upper Peninsula, the assumption was made that sightability of birds will not be constant for the entire state. A subset of the segments of fixed-wing flights is flown with a helicopter to quantify differences in sightability. As of 1993, visibility correction factors (VCF's) for three species, mallard (Anas platyrhynchos), Canada goose (Branta canadensis), and wood duck (Aix sponsa), were calculated using only data collected on flights in Michigan. Visibility correction factors for all other species have been calculated from flights and ground surveys in Ontario. Waterfowl Modelling: Estimating the fall flight of waterfowl from the prairie pothole region of North America has been a goal of the United States Fish and Wildlife Service and the Canadian Wildlife Service since the inception of the WBPHS and the 4 WPHS. Several models of mallard production have been developed for this region (Walters et al. 1974, Cowardin and Johnson 1979, Martin et al. 1979, Dudderar 1985, Johnson et al. 1986, Johnson et al. 1987, Cowardin et al. 1988, Johnson et al. 1988). No other species on the prairie potholes received this attention, and almost no work has been done to model species outside the prairie pothole region. Waterfowl production models have never been developed for Michigan. Waterfowl production models have taken two common forms: flowchart based models which determine success of individual animals and mechanistic models which employ survival and reproduction rates at the population level. The flowchart based model (Figure 1) uses a tally of the individual birds which survive the season (@>in.Figure 1). and the birds which do not (C>in Figure 1). The mechanistic model is composed of a series of equations which apply population derived reproduction and survival rates to individuals, which are then used to estimate end values for the population as a whole (Figure 2). Estimates from the Michigan Breeding Waterfowl Survey suggest that Michigan accounts for approximately 400,000 breeding mallards, 150,000 breeding Canada geese and 5,500 breeding black ducks (Anas rubripes) each year. All of these birds and their young which survive to migrate comprise the DummMMnghnuml “huwawummu BMIGNMMUW Dmuflsanmn} wauduum/ Cunnafluflmmn egg-um Dmnkwmnmn ufiwhdomnr Aammmmmpncmn OnnGNsmNWdrmn} Pnfldfimrflehi } Bh!awflfia1 [hwctmwhmm 'Bmodsmvivalrlto} omnqunKMbmynna} Off-nutmomiitymte} hMmhvof bankcamn F_L_VI i l Numberot funniesdead Figure 1: Flowchart based waterfowl model (Johnson et al., 1986) 6 fall flight, which provide recreation through sport hunting and viewing. An accurate assessment of production within the state is necessary to set appropriate bag limits and season lengths. Production rate =(eggs produced per adult)* (survival rate of eggs to hatching)* (survival rate of chicks to fledging)* (survival rate through early flight period) Production =(new adults in spring * production rate for first breeding) + (old adults in spring * adult production rate) Fall adult population = (new & old spring adults)* (adult summer survival rate) Fall juvenile population = (production)*, (juvenile summer survival rate) Fall flight = fall adult population + fall juvenile population Figure 2: Mechanistic model equations to estimate fall flight by species (Walters et al. 1974) OBJECTIVES The objectives of this study were to develop statistically and biologically sound models to predict the fall flight of mallards, black ducks, Canada geese, and wood ducks from Michigan and to provide recommendations for refinement of the Michigan Breeding Waterfowl Survey. METHODS Michigan Breeding Waterfowl Survey And Visibility Correction Factors: Michigan's Breeding Waterfowl Survey (MEWS) was conducted in late April and early May of 1991, 1992, 1993, and 1994. Yearly timing depends on the weather since the survey needs to start after spring migrants have returned and end before complete leafout. Because of these factors, the state is censused earliest in the south and latest in the Upper Peninsula. Differences in land use and cover types in the state have necessitated the division of the state into two strata for the purposes of the survey (Figure 3). Stratum A, the Forest Stratum, is composed of the Upper Peninsula and the northern Lower Peninsula. The Upper Peninsula consists mainly of forests of conifers and northern hardwoods (Omernik and Gallant 1988). Forest and vegetation cover is very thick with little in the way of cleared areas for either agricultural or residential use. The northern Lower Peninsula is a fairly equal mixture of forests of conifers and northern hardwoods, hardwood stands with moderate agricultural development, and agricultural and urban areas — Stratum A - — Stratum B gym/mag; Areas omitted from the survey "-:« -'.-;‘.v.:x-. my _‘ ' Rmmm&fl Figure 3: Michigan Breeding Waterfowl Survey transect routes 10 with woodlots (Omernik and Gallant 1988). Transects flown inthis stratum are 28 miles apart and contain a total of 59 18—mile long segments. Stratum B, the Farm-Urban Stratum, encompasses the remainder of the Lower Peninsula and is characterized by heavy agricultural and urban development with woodlots and swamps dotting the landscape (Omernik and Gallant 1988). At the time of the survey, cover over most of the region is sparse with the exception of flooded swamps and woodlots. The transects in the stratum are 14 miles apart and contain 93 18-mile segments. The difference in survey coverage is due to the greater amount of vegetative cover in the Forest Stratum and the traditional higher density of breeding birds in the Farm—Urban Stratum. Much of the procedure used in the MEWS follows the techniques set forth in the Standard Operating Procedures for the Aerial Waterfowl Breeding Ground Population and Habitat Surveys (USFWS 1987). The few differences include the use of helicopter flown segments in Michigan for calculation of VCF's, rather than ground walked segments, and the addition of an observer on fixed-wing flights so that the pilot does not act as an observer. The MDNR counts ponds (i.e. "water areas"; e.g. rivers, streams, ditches, marshes, swamps, ponds, lakes) on all segments, as set forth in the Standard Operating Procedures (SOP) from the USFWS. 11 The fixed-wing flights follow transects across the state, recording the number and species of waterfowl seen. The transects are flown 100—150 feet above the ground with 10-15 minutes being spent on each 18-mile segment. The area observed along each transect is % mile wide, with an observer on each side of the plane being responsible for half of that width. The observer in the front right seat of the plane (next to the pilot) counts birds and acts as the navigator; the rear seat observer (behind the pilot) counts birds and water areas visible on the left side of the plane. Birds are recorded by species and gender; gender is noted by separation into the categories of lone drakes, flocked drakes, and pairs. All unidentified birds are also recorded as such. Individual transects are flown in the same direction (east-to-west or west-to-east) every year. A subset of the segments of the total survey are flown with a helicopter as well. This allows for some quantification of the visibility bias of the fixed—wing flights. These flights are made with the observers seated as in the fixed-wing flights and performing the same duties. The same amount of area is surveyed on helicopter segments as fixed—wing segments. Unlike the fixed-wing segments, however, on helicopter flights, travel north and south along the segment to search for birds is allowed. The segments are 12 flown at 150 feet or lower; often the helicopter will descend to 5-10 feet to better search for birds. It takes from 34 to 185 minutes to fly each 18—mile segment. Visibility correction factors in Michigan-are calculated using the ratio of birds counted on helicopter flights to birds counted on fixed-wing flights as set forth by Martin et al.(1979). 2 indicated birds seen by helicopter on helicopter flown segments VCF - 2 indicated birds seen by fixed-wing on helicopter flown segments where indicated birds - (ZXEsingle drakes)o(2)<2pairs of birds)+(2flocked drakes) Variance formulas for VCF's are also set forth by Martin et al. (1979). Estimates of the total adult spring population for each species are calculated using the number of indicated birds from the fixed-wing flights, the relevant VCF's, and an expansion factor to convert from area in the transects to area in the state as follows: adult spring _ ( total indicated birds )x(VCF)x( total area of stratum population counted on fixed-wing flights area of stratum surveyed Visibility correction factors are calculated separately for each species, each stratum, and for each year. l3 Species Of Interest: Species for which the MDNR expressed interest in production models included mallard, Canada goose, wood duck, blue—winged teal (Anas discors), black duck, ring—necked duck (Aythya collaris), and mergansers (MErgus spp.). The scientific literature was searched for estimates of the various inputs to the model. Mergansers, for which very little reproduction research had been conducted, were eliminated as potential model candidates. Blue-winged teal, for which some information was available, were eliminated as potential model candidates, since no information was available on breeding in Michigan, and their VCF's are very high, suggesting that population size estimates are likely to be highly inaccurate. The ring-necked duck, for which there is information on breeding success in Michigan and which has a low VCF, was eliminated since the individuals seen on the NEWS were not on their traditional breeding grounds, suggesting that the estimates of breeding birds include migrants as well as breeders (Reeves 1991). Three of the remaining species, mallard, black duck, and Canada goose, fit the requirements of ample literature values, available Michigan production values, sightings on confirmed breeding grounds, and low VCF's moderately well. A wood duck 14 model was constructed due to current management needs of the MDNR, despite a poor estimate of adult spring population. Model Framework: The framework for the four models is the mechanistic model developed by Walters et al. (1974) for mallards in the prairie pothole region as adapted to available information. This model consists of five equations: survival rate of survival rate )x( chicks through )x(through early) fledging flight stage Production eggs survival rate Rate ' (produced/admit) x (until hatching Production - (adult spring population)X(production rate) Juvenile Fall Population - (production)X(juvenile summer survival rate) adult breeding season survival rate Adult Fall Population - (adult spring population) X( ) Fall Flight - (adult fall population)+(juvenile fall population) All inputs, except for the adult spring population, are from the scientific literature. Preference was given to values calculated from research conducted in Michigan and to recent values. Where Michigan based research or recent values were not available, values from the northern end of the Mississippi and Atlantic Flyways since 1960 were used preferentially. Values outside these areas, such as clutch 15 sizes in Canada geese, were used when local values were not available or to support local values when few were available. The adult spring population is calculated using the counts of indicated birds and VCF's from the NEWS. Model Development: The methodology used to construct the models is outlined in Figure 4. The first model output generated is the production rate, which was calculated using the following process: 1. An equation for production rate was built that incorporated the endpoints available in the literature for each species (e.g. clutch size, nest success, survival from hatching to flight, survival from laying to flight, etc.) The purpose of this equation was to account for all mortality from the point at which eggs are laid until the young reach early flight stage. 2. The available literature values were input into the equations in all possible combinations. 3. The results of this initial calculation were examined for extreme values (Figure 13, Figure 17). Values were judged to be extreme when a single input 16 Specre' s of interest from MDNR -rnallard -black duck | Collection of Breeding .g-ingnecig‘iick Waterfowl Sum data from blue-winged teal -merganser -Canada oose \. Map distribution of birds during spring survey l v Review USFWSICWS standard operatin procedures and MD R techniques Calculation of visibility correction factors and their 95% confidence intervals Examine pond number and bird number relationship 7 Calculate number of Construction of framework breeding birds of mechanistic model Use literature values and MDNR population estimates Recommendations for the in the model framework Aerial Waterfowl Bmding Ground Population Survey f l 3 Construct probability distributions for appropriate model elements Y Run repeated iterations of ecom mm the models using remit)” {uglier want:rtowl breefging drawn values - - ' cted distributions research in Michigan Generate estimate offall flight and its vanance' Figure 4: Model development flowchart 5. 17 was responsible for several very high or very low production rate values. The only example of this found was the value of 9.2% for the survival of black ducks from hatching to fledging (Stotts and Davis 1960). This value alone is responsible for the production rate values in the range 0.15 to 0.40 (Figure 17), therefore, this value was excluded from further model building efforts. After examination of these results, probability distributions were constructed for each production rate component to reflect patterns which were observed in the literature values. An example of this is the high frequency of values in the range 0.5-0.565 for nest success of black ducks. The majority of the area under the curve for the nest success distribution constructed is in the range 0.5—0.565 (Figure 19). When only two or three literature values were available, a uniform distribution between the minimum and maximum was used. An estimate of production rate was then calculated by randomly drawing one value from the distribution of each production rate component. These were input into the model equation of production rate, 18 generating a value in the units of young produced per breeding adult. This was repeated ten thousand times to generate ten thousand production rate estimates. 6. Production rate frequencies from these calculations serve as the production rate probabilities for the models. The mean and variance of this distribution served as the production rate component of the models. The remainder of the model consists of point estimates, rather than distributions, and is of a simpler construction. The adult breeding season survival rate is the mean of available literature values. Due to a lack of research on survival of young from fledging to migration, the juvenile summer survival rate for all species except wood ducks, for which literature values are available, is assumed to be equal to 1, with a variance of 0. A point estimate and variance was calculated for each model output value. Renesting values are not included in the model due to a paucity of available literature values for renesting. To judge the degree to which model values reflect actual conditions, projections of the number of spring breeders in Michigan were produced. Assumptions were made that all birds home to their natal breeding grounds and that 19 the sex ratio in eggs is 1:1. The projections were calculated using a reproduce-then—survive life table approach, and the projections were run for at least 80 years to allow low production rates to drive the population to 0. When low production rates, in units of young produced per breeding adult, were applied to the projections, the population continued to produce young after all the females in the population had died. To avoid this, production rates, in the units of young produced per breeding female, were applied only to the females in each population. These results can be compared to model production rate results, for mallards and black ducks, by dividing projection production rates by 2 since these rates are in the units of young produced per breeding female, and model production rates are in the units of young produced per breeding adult. Wood duck rates, however, are in the units of young produced per breeding female for both the model and the projections as this made it easier to model dump nesting. These projections allow for the determination of which production rates yield declining, stable, or increasing population sizes. Model production rates were also compared to ratios of immatures per adult in the harvests from Michigan for 1987— 1991 (Padding et al. 1992). These numbers do not accurately 20 estimate the number of young produced per adult, but, after modification, they have been used as an approximation of that value for mallards (Fred Johnson, USFWS, pers. com.). As the numbers are calculated using wing survey data, the higher vulnerability of immature birds is reflected in values which are higher than the actual production rate. To adjust for this, immature per adult ratios were divided by two. This will not isolate Michigan's production from overall flyway production as some fraction of the birds harvested in Michigan are a result of production north of Michigan, however, it is the only large scale estimate available which may approximate Michigan's production rate. These numbers were then compared to production rate values for all species except Canada goose. At the time of the harvest, it is difficult to differentiate between adult and juvenile Canada geese based solely on wing plumage, resulting in an highly inaccurate estimate of immatures per adult in the harvest. RESULTS & DISCUSSION Mallard: All of the available appropriate literature values (Table l) for the mallard were used, though some nest success values were given low probabilities of occurrence, and the adult female survival rate values were supplemented with black duck values. While there are several very low values for nest success (Bilogan 1992), these are the daily survival rates as calculated using the Mayfield method (Mayfield 1960) and cannot be directly compared to the other values, which are simply the ratio of successful nests to total initiated nests. For the purposes of this model, simplistic survival rates were considered to be suitable as they do not assume that days are independent, as Mayfield does, and the survival rates required by the model are in terms of survival to a specific biological endpoint rather than over a set period of time. Due to the sparsity of adult summer survival rates, the female survival rates of the mallard and black duck were combined for a mean value of 0.685. Both the adult male and adult female summer survival rates were assumed to have a variance of 0. Annual survival rates from Anderson (1975) were used in the population 21 22 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII-I-I-I-I-I-I-I-I-I-I- Table 1: Mallard model input values model value sourcea input clutch 9.6 Coulter and Miller 1968 size 9.0 Bilogan 1992 nest 0.20 Bilogan 1992 success 0.01 Bilogan 1992 0.04 Bilogan 1992 0.01 Bilogan 1992 0.63 Orthmeyer and Ball 1990 0.78 Krapu and Luna 1991 999 success in 0.784 Talent 1980 successful nests survival from 0.35 Talent et al. 1983 hatching 0.33 Krapu and Luna 1991 to flight survival from 0.3951 Orthmeyer and Ball 1990 laying to 0.44 Ball at al. 1975 flight adult 0.713 (9) Kirby and Cowardin 1986 summer 0.603 (9) Blohm et al. 1987 survival 0.998 (d) Blohm et al. 1987 rate juvenile summer 1.0 (var.=0) assumed survival rate adult 1991: 277,729 (var=8.5*10% MBWS spring 1992: 371,288 (var=2.3*lOU MBWS population 1993: 412,104 (var=2.0*10U MBWS 1994: 716,918 (var=2.7*10m) MBWS a:values not found in the literature were set to values designated by “assumed" 23 projections. Calculated production rates (Figure 5) compare favorably to those estimated to be necessary to maintain a constant population size (Figure 6) and to immature per adult ratios in Michigan's harvests (Padding et al. 1992). Corrected values of immatures per adult in the harvest range from 0.75 to 0.95 (1987—0.75, 1988—0.95, 1989-0.9, 1990—0.8, 1991—0.8). According to values yielded by the projections, a production rate near 0.79 young produced per breeding adult is necessary to maintain a stable population, and this value lies within the range of production rate values generated by the model. These runs of the model use the means and variances of the production rate distributions as the model production rates, though this method does not allow for annual variation in breeding conditions. An equally valid method would use the peak of the production rate distribution (0.40) as the model rate of an average year; higher or lower values could then be selected for years during which breeding conditions are better or worse. This choice can be influenced by the timing of spring thaw, number of cloudy days in the spring, amount of available breeding habitat, quality of breeding habitat, amount of available food, and the condition of the birds returning from their wintering 24 mmSHc> when cofluospoum UHMHHME mo cofiusnflwumflo .m mwsmflm 38 gazes—coca me... mm... moé mad mod med mud mod 0 - -13 33 com 3 ---- o9q - 1313-, 333; com ---------- 33-3-3- -- -- -3333 com gape Lon 80:35 950» u 2.2 5:335 .----33-3-3-,- , ------------- i coo... com. F Fees—co: 25 semanofiz Ga mUHmHHmE Mom mwSHm> cofluoasmom Umuommowm tee> Pmmnmonmovvvmmmmtmp d4‘_..——«Aquuaq4—Aflqjl—qqqdddd—4_1a—a.__-.«___HJ.‘-.-_-.__qq—_—._d-___«#4441— I i ,th. ; £25¢3a§825a056>u82c26265 000.000 000.00»‘ 000.000 000.000 000.005 000.000 53%.... 3...: 0503.3 05.5.». gate .0 mwsmflm 26 grounds. The majority of the fall flight variance originates in the estimate of adult spring population (Table 2). The high value for the 1994 fall flight and its variance, almost twice that of 1992 or 1993, is due entirely to the MEWS estimates. All of the estimates of the adult spring population vastly exceed historic estimates of the breeding population in Michigan, making comparison of model estimates of the fall flight to historic values impossible. For example, Bellrose (1976) states that there are 47,000 breeding mallards in Michigan, a figure much lower than the 270,000 to 716,000 arrived at through the MEWS. 27 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII-I-I-I-I-I-I-I-I-I-II Table 2: Mallard model output values model output year estimate variance production rate all years 0.490744 0.057056 production 1991 136,294 4.6*109 1992 182,207 8.4*109 1993 202,238 l.0*10lo 1994 351,398 2.1*1010 adult fall population 1991 238,637 6.9*10a 1992 318,810 l.8*109 1993 357,786 1.6*109 1994 617,664 1.1*1010 juvenile fall population 1991 136,294 2.3*109 1992 182,207 2.1*109 1993 202,238 2.5*109 1994 351,398 1.l*1010 fall flight 1991 374,931 3.0*109 1992 501,017 3.9*109 1993 560,024 4.2*109 1994 969,062 2.1*1010 28 The following equations constitute the mallard model: Production Rate = E/A*Sn*SH where E/A C/2 Sl—f = Se*Sh—f or SM = literature values Var(Production Rate) = variance of production rate distribution Production = As*Production Rate stratum B var(Product.)- 2: [(Aflixvar( rustratum A product. rate product. rate )l:[( )’x (Inxxn) 2Warn/CF") 1 Adult Fall Population = Afs*Safis + Ams*Sams adult females stratum 8 var( ) ' ( fall-POP- .bmales instratum.A [(A,) inxVarisu) ,M is") ixlrhxxfl) ZxVarWCF) .1) Juvenile Fall Pop. = (0.5*Product.*(8”)flmfle)+(0.5*Product.*(Sfi)mflg 'uvenile females var(f7all pop.) ' l-mgles ( [ (0-5xproduct. )2XVar(SJ_)1]+[ ($1.):XO.ZSXVar(product. ) ] ) Fall Flight = Adult Fall Population +Juvenile Fall Population Var(Fall Flight) = Var(Adu1t Fall Pop.)+ Var(Juvenile Fall Pop.) where: E/A = eggs per adult Sn = nest success 81..f = survival from laying to flight C = clutch size Se = egg success in successful nests Smf = survival from hatching to flight Ag = adult spring population (APS = females; AHS = males) I = number of indicated birds (from MEWS) X = area expansion factor (from MEWS) VCF = visibility correction factor (from MEWS) Sas = adult summer survival rate Sjs = juvenile summer survival rate 29 Black duck: With the exception of one value for survival from hatching to fledging, all black duck literature values (Table 3) were used in the model. The value from Stotts and Davis (1960) was excluded since its presence caused the production rate distribution to have a peak at the low end of what would otherwise have been a fairly smooth curve (Figure 17). Values for adult summer survival rate were derived using black duck and mallard data. The female survival rate is the mean of the black duck and mallard values; the male survival rate is simply the male mallard survival rate. Due to the similarities of biology between these two species, the assumption of equivalent survival rates is reasonable. Annual survival rates used in the projections are from Blandin (1982). The mean of the production rate distribution (Figure 7) is considerably lower than what would be necessary, 1.14 young produced per breeding adult, to maintain a stable population (Figure 8). The values do, however, compare favorably to corrected immature per adult ratios in the wing survey (Padding et al. 1992) which range from 0.55 to 0.95 30 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII-I-I-I-I-I-I-I-I-I-Il Table 3. Black duck model input values model input value sourcea clutch size 9.2 Reed 1970 9.1 Stotts and Davis 1960 9.5 Coulter and Miller 1968 nest success 0.50+ Reed 1970 0.55 Coulter and Mendall 1968 0.67—0.84 Coulter and Mendall 1968 0.565 Stotts and Davis 1960 0.55 ‘ Laperle 1969 egg success in 0.911 Reed 1970 successful nests 0.848 Stotts and Davis 1960 survival from 0.4244 Ringelman and Longcore hatching to 0.092 1982 fledging Stotts and Davis 1960 survival from nest exodus to 0.29 Reed 1970 flight adult summer 0.74 (9) Ringelman and Longcore survival rate 0.998 (d) 1983 assumed juvenile summer 1.0 (var.=0) assumed survival rate adult spring 1991: 6,105 (var=l.1*10fi MEWS population 1992:12,988 (var=3.8*10U MEWS 1993: 5,856 (var=7.1*1OW MEWS 1994: 8,009 (var=3.1*10fi MEWS ‘ values not found in the literature were set to values designated by "assumed" 31 mmSHc> moon cofluospoum xosp xocan mo Coflusflflwumfla 3a.. 600300.:— 0mé 00;. NN.—. S; 00... 00.0 00.0 No.0 3.0 00.0 00.0 gape Log 082005 056» u 38 5.63005 o -:-:-oos 8N --;--.oom .:-:---oov .;--,--:-oom --::-.;--;-ooo 00x. Eco-.02“— .b mwzmflm 32 cmofinofiz ca mxodp xomHn no“ mosam> coaumHSQom Umuomflowm .0 madman teo> 02.00500030000th - «uq—~.-——-—..___—.—_a—__—_—_—_«—u———_._qqu—__~__—d—qq—10-———l——_——q.0_.—d—~_—q__—_« coo m 000.0 000.0 000.0 000.» 000.0 000.0 \.\. . 0.25.. tan 08:35 0:30» u S! concave... . .\ii.- - - - i .- -. -- - 000 or 000. _. _. sass... not... 050er 05...? gave 33 (1987-0.55, 1988—0.95, 1989—0.85, 1990—0.7, 1991—0.95). While neither the projections nor the immature per adult ratios in Michigan's harvest provide accurate endpoints for production rate, these patterns imply that the model reflects production in Michigan, which is undergoing a decline in black duck numbers. As the model reflects the basic biology of the black duck during breeding, it would appear that there is little potential for growth of the population in Michigan without either improvement of breeding habitat, which would push the production rate toward the high end of the distribution, or reduction in annual or seasonal mortality, which would lower the production rate necessary to maintain a stable population in the projections. As with the mallard model, the majority of the variability in the model is due to the high variance of the MEWS (Table 4). The black duck VCF's used in Michigan, however, were derived from segments of the WBPHS in Ontario and have lower variances than some of the VCF's calculated in Michigan for other species. Also, this model may be adapted to perform under varying conditions. Before complete faith is placed in the model, however, verification of the model inputs must be made in Michigan. The values used in the model, at present, 34 were derived from research conducted on the East Coast of North America; none of the values come from Michigan, or anywhere else in the upper Midwest. 35 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII-I-I-I-I-I-I-I-I-I-II Table 4. Black duck model output values model output year estimate variance production rate all years 0.851276 0.013541 production 1991 5,197 l.3*106 1992 11,056 5.1*106 1993 4,985 9.8*10S 1994 6,818 2.9*106 adult fall population 1991 5,137 8.1*105 1992 11,489 3.3*106 1993 4,927 5.2*10S 1994 6,740 1.2*106 juvenile fall population 1991 5,197 6.5*10S 1992 11,056 l.3*106 1993 4,985 2.4*105 1994 6,818 2.9*106 fall flight 1991 10,334 2.1*106 1992 22,545 8.4*106 1993 9.912 l.5*106 1994 13,557 2.6*106 36 The following equations constitute the black duck model: Production Rate = iii/Al"Sn*Sen"'Sh.f where E/A = C/2 Var(Production Rate) = variance of production rate distribution Production = A3*Production Rate stratum B Var(Product.) - 2: [(As): rustratum A xVar(product. (product. rate 2 2 )]+[ rate ) x(Inxxnl xVar(VCF,,)] Adult Fall Population = Am*SM3 + AM*SMM females stratum B - 2 ( Z [(A,)§nxVar(Su)1]o[ (suijx(Ilnxxniszaz-(vcrini> l-males n-s tra tum A adult var(fall pop.) Juvenile Fall Population = (0.5*Product.*(SF)MMRJ+(0.5*Product.*(Sn)mhg juvenile females 2 2 var(fall pop.) - l-mgles ( [ (0.5>7 year olds - 94% Moser and Rusch 1989 age structure 1 year olds --—35% Kelley 1993 of females 2 year olds ---27% Kelley 1993 3 year olds —-—19% Kelley 1993 4 year olds ~-- 9% Kelley 1993 5+ year olds --10% Kelley 1993 adult summer 0.85 (9) assumed survival rate 0.85 (d) assumed juvenile summer 1.0 (var.=0) assumed survival rate adult spring 1991: 58,787 (var=1.8*10U MEWS population 1992: 101,587 (var=4.7*10U MEWS 1993: 177,999 (var=1.7*109) MEWS 1994: 308,083 (var=1.0*10m) MEWS a:values not found in the literature were set to values designated by "assumed“ 39 mmsam> much cofluodpoum HmEflqu mmoom mpmcmu mo coflusnfiuumflm .m mwdmflm 3s. 60269:. .2530 00 00.. 00.. 00N 520 - - - - - - OMN .0 Bo 83> o>c «EB too 08:09.. 0:33 u 28 cozoacoa EEnao 000 Eco—.02.— 40 estimate of the realized production rate. This is accomplished by multiplying each optimal production rate estimate by the percent of the population which is one year old females, then multiplying by the relative breeding rate of one year old females; this is done for all ages through five year olds and older. The sum of these values is the estimate of the realized production rate. Repeating this procedure results in ten thousand estimates of the realized production rate and a distribution which represents the production rate of the population corrected for age specific reproductive output (Figure 10). No projections were calculated as the results are highly dependent on the age structure of the population. At present, little information is available for accurate assessment of the age structure of Canada geese in Michigan. The age structure used here is the result of band returns of only a few hundred birds and is not likely to be representative of the actual values. As stated earlier, immatures per adult, as recorded using the wing survey, are not representative of immatures per adult in the harvest or immatures produced per adult in Michigan and cannot be used as an indicator of accuracy of the model. Anecdotal information on general population trends in Michigan from 41 mmdam> mumu cofluusponm UmNfiHmmH mmoom mpmcmu mo cowusflfluumfia .OH wusmflm £2 roses—.95 03:02 .00 00.0 50.0 00.0 0.0.0 3.0 00.0 50.0 00.0 00.0 «0.0 00.0 .-,;-i- com -- -- cow +----- coo --, --- com Eiiggucaguefizcoaesg 00332 000. _. 55:00...— 42 Gerald Martz of the MDNR (pers. com.) has proved to be the best index to actual production. Increasing nuisance goose complaints and annual harvests indicate a large and increasing population. This suggests that the realized production rate yielded by the model underestimates actual production. As with the other models, most of the fall flight variance is due to values from the MEWS (Table 6). Likewise, any improvements in the survey would improve the model. However, its greatest needs lie in the areas of accurate measurement of age specific reproductive output and the population's age structure. Until such time as these values are improved and a fall census is made, there will be no way to judge the accuracy of the estimates from the model. 43 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII-I-I-I-I-I-I-I-I-II-I Table 6. Canada goose model output values model output year estimate variance optimal production rate all years 1.510178 0.019617 realized production rate all years 0.385397 0.001278 production 1991 22,656 7.1*106 1992 39,152 2.0*107 1993 68,600 3.0*108 1994 118,734 1.7*109 adult fall population 1991 49,969 1.3*107 1992 86,349 3.4*107 1993 151,299 1.3*109 1994 261,871 4.2*109 juvenile fall population 1991 22,656 3.5*106 1992 39,152 5.0*106 1993 68,600 7.5*107 1994 118,734 8.4*108 fall flight 1991 72,625 2.0*107 1992 125,501 5.4*107 1993 219,900 1.6*109 1994 380.605 5.0*109 44 The equations of the Canada goose model are as follows: Optimal Production Rate = E/A*S,,"'Sg where E/A = C/2 Var(Opt. Prod. Rate)=variance of Optimal Production Rate distribution Realized Production Rate (RPR) = [(Optimal Prod. Rate*B31fi) for r = 1 year olds, 2 year olds, 3 year olds, 4 year olds, 5+ year olds Var(Realized Prod. Rate)=variance of the Realized Prod. Rate distribution Production = Ag*Realized Production Rate stratum B Var(Product.) - 2 [(A‘):XVar(RPR) ].[ (RPR)”x(InxanxVarivcrn)] rustratum A Adult Fall Population =.A“*S“m + Am*SmM adult females stratum B var( ) ' ( fall 909' 1.males n-stratum A [(A,)§nxVar(s,,) ,M (suljx (Imxxfl) ’xVar(VCF)nll Juvenile Fall Population = (0.5*Prod.*(8”)hmhm)+(0.5*Prod.*(Sn)mu5) juvenile females var(fall pop.) ' bangles(i(0.5xproduct.)ZXVar(Sj.)l]*[($1.)EXO.25xVar(product.)]) Fall Flight = Adult Fall Population+Juvenile Fall Population Var(Fall Flight) = Var(Adult Fall Pop.)+ Var(Juvenile Fall Pop.) where: /A = eggs per adult survival rate of eggs to hatching survival of goslings to flight clutch size relative breeding rate % of population adult spring population (An = females; Ams = males) number of indicated birds (from MEWS) area expansion factor (from MEWS) CF = visibility correction factor (from MEWS) ” = adult summer survival rate (Sfl,== females; Sams = males) Sjs = juvenile summer survival rate 0 O H aagnu10cn030 niiu 45 Wood duck: The input values for wood ducks (Table 7) differed from the other species in that two of the inputs already existed as distributions of raw data rather than point estimates. Distributions were given for clutch size and survival from laying to hatch. Clutch size values were broken down into natural nest values and nest box nest values. The ratio of natural nests to nest box nests in Michigan is not known and was assumed to be 1:1 for the purposes of the model (Figure 25). To simplify conversion of this data into a model input, it was retained in the form of clutch size rather than set to eggs per adult. This results in a model production rate value in the units of young produced to flight stage per breeding female, rather than per breeding adult. Similarly, survival from laying to hatch values were segregated between normal and dump nests; again, a 1:1 ratio was assumed (Figure 26). Otherwise, the model was constructed similarly to the mallard and black duck models. Annual survival rates for the projection were found in Bellrose and Holm (1994). Due to the poor quality of input values, most notably MEWS data, and the poor understanding of wood duck breeding biology in Michigan, the model value of production rate (Figure 11) and projection values (Figure 12) are 46 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII._.-I-I-I-I-I-I-I-I-I-II Table 7. Wood duck model input values model input value sourcea clutch size natural nest values (p.225) Bellrose and Holm 1994 nest box values (p.226) Bellrose and Holm 1994 survival from 'normal nest values (p.250) Bellrose and Holm 1994 laying to hatch dump nest values (p.250) Bellrose and Holm 1994 survival from 0.41 Klein 1955 hatch to flight 0.53 Grice and Rogers 1965 0.42 Grice and Rogers 1965 0.48 Grice and Rogers 1965 0.59 Prince 1965 0.38 Holloman 1967 0.53 McGilvrey 1969 0.50 McGilvrey 1969 0.59 McGilvrey 1969 0.53 McGilvrey 1969 0.56 Baker 1971 0.52 Baker 1971 0.52 ‘ Brown 1972 0.59 Haramis 1975 0.61 Haramis 1975 0.68 Hepp 1977 0.44 Cottrell 1979 0.58 Rothbart 1979 0.40 David 1986 0.56 Bellrose and Holm 1994 0.56 Bellrose and Holm 1994 0.48 Bellrose and Holm 1994 0.55 Bellrose and Holm 1994 0.53 Bellrose and Holm 1994 adult summer 0.8 (9) (var.=0) assumed survival rate 0.8 (d) (var.=0) assumed juvenile summer 0.91 (9) Kirby 1990 survival rate 0.89 (6) Kirby 1990 adult spring 1991: 57,588 (var=l.1*10% MEWS population 1992: 485,008 (var=l.0*10“) MEWS 1993: 206,182 (var=1.4*10”) MEWS 1994: 99,759 (var=1.8*109) MEWS a:values not found in the literature were set to values designated by "assumed" mmzam> mumu cofluusponm x030 0003 00 Goflusnfluumwn .HH musmflm 32 5.85020 0 0.\. n 0.0 0 0.0 0 0.0 v 0.0 0 0.0 N 0.9 _. 0.0 0 47 adjadajaj_fi.1..1.l_-..I. «ll. , , _ o w- - _ e e 2: e 8m 8m 2: - ----- ................................ Sm -. ------------------------------------- - 000 00» 35:02.— 48 Gmmflnoflz G0 mxozp 0003 How mmSHm> coflumHsmom Umuowhoum .NH mnnmflm use» 0s. 00 N0 00 S. 00 00 t. 0 .. .«quaud—_-..ufi‘._.ujfi—jA—ju—j—uAud—Hd~—d__—.fifiA—«_—qdq4-__.d«—_u‘—_.qdqaqfi \\ 12505598555055>u82028255 0 000.00 000.00.. 000.00 .. 000.00N 000.00N «050 0503.5 055» 5:00 49 considerably higher than the corrected ratios of immatures per adult in the wing survey. The corrected wing survey fall in the range of 0.6 to 1.25 (corrected wing survey ratios: 1987—0.85, 1988—0.9, 1989—1.25, 1990-0.6, 1991-0.7) (Padding et al. 1992), while the model value and projection value required for a stable population range from 2.23 to 2.65. This suggests that the production rate specified by the model overestimates the actual production rate by 100% to 300%. Any of a number of steps could be taken to improve the model (Table 8). The VCF's for wood ducks have been, for the most part, at least a factor of ten higher than the VCF's for other species; the variances of these VCF's are also very high. This is a result of surveying wood ducks with fixed—wing aircraft, which cannot be used to effectively observe all wood ducks in an area. Helicopter surveying of wood ducks is more effective than fixed—wing surveying, as can be seen on the segments used for the visibility correction factors. Michigan-based estimates of clutch size and ratios of natural nests to nest box nests, survival from laying to hatch and ratios of normal to dump nests, and survival from hatching to flight should go a long way toward reducing the difference between model outputs and external measures of production. 50 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII-I-I-I-I-I-I-I-I-I-II Table 8. Wood duck model output values model output year estimate variance production rate all years 2.23414 0.714675 production 1991 128,660 7.6*109 1992 1,083,576 6.8*1011 1993 460,638 9.8*1010 1994 222,876 1.3*1010 adult fall population 1991 46,070 6.7*108 1992 388,007 6.6*1010 1993 164,945 8.7*109 1994 79,807 5.9*108 juvenile fall population 1991 115,794 3.5*109 1992 975,219 1.7*1011 1993 414,575 2.5*1010 1994 200,589 5.2*109 fall flight 1991 167,623 4.0*109 1992 1,411,725 3.6*1O11 1993 600,137 5.1*1010 1994 280,396 5.8*109 51 The wood duck model equations are as follows: Production Rate = C*Sl__h*Sh_f Var(Production Rate) = variance of production rate distribution Production = A3*Production Rate stratum B r Var(Product.) - 2 [(A')EXVar(p rustratum A oduct. rate oduct. pr rate )]'[( )Zx (InXXn) ZXVar(VCFn)] Adult Fall Population = Afs*Safs + Ams*Sams adult females stratum 8 var( ) ‘ ( fall pap' 1.1113193 n-stratum A [(A_) inxVaMSul ,H is“) iMIlnxXn) 2Warn/C109) Juvenile Fall Population = (0.5*Prod.*(8”)fimnw)+(0.5*Prod.*(Sfi)mkm) juvenile females 2 2 Var(fall pop.) - l-m§1e5([(o.5xpr0du‘:t.) 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