Data to inform population assessment of the Interior subspecies of band-tailed pigeon, Patagioenas fasciata fasciata (breeding range from Colorado and Utah south into Sierra Madre Occidental of Mexico), have been lacking despite substantial past banding efforts. We used a data set of more than 26,000 bandings from Colorado, with 3,500 live recaptures and 780 recoveries from the harvest of banded individuals to estimate annual survival, fidelity, and harvest rates. Most birds were harvested in Colorado (62%) followed by Mexico (18%); New Mexico (16%); Arizona (3%); and 1% or less each in California, Washington, and Utah. On average, each year 15% (range 0–30%) of surviving band-tailed pigeons did not return to Colorado. From 1969 to 1981 mean annual survival was 0.633 (standard error [SE] = 0.031) for hatch-year and 0.719 (SE = 0.016) for after-hatch-year birds, with a mean annual recovery rate of 0.015 (SE = 0.002) for hatch-year and 0.011 (SE = 0.001) for after-hatch-year birds. From 1970 to 1974, mean annual abundance of band-tailed pigeons in Colorado on 1 September was 59,911–88,290. These data provide a baseline for additional data collection for band-tailed pigeons in the range of the Interior subspecies.

The band-tailed pigeon, Patagioenas fasciata, is a migratory game bird in western North America. Two subspecies occupy two distinct geographic areas in western North America: Coastal P. f. monilis (British Columbia south into Washington, Oregon, California, Nevada, and into Baja California) and Interior P. f. fasciata (Colorado, Utah, Arizona, New Mexico, west Texas, and into central Mexico; Keppie and Braun 2000). Harvest management decisions are made independently for each subspecies (Pacific Flyway Study Committee and Central Flyway Webless Migratory Game Bird Technical Committee 2001). A survey of band-tailed pigeons at mineral sites is used to inform harvest management decisions for the Coastal subspecies (Casazza et al. 2005). The North American Breeding Bird Survey has been used to assess the status of the Interior subspecies (Sauer et al. 2003). However, the usefulness of the Breeding Bird Survey is questionable because the Interior subspecies is detected on few routes and indices of abundance have high variance (Sanders 2014). Thus, there is a pressing need for baseline demographic data for Interior band-tailed pigeons.

More than 26,000 band-tailed pigeons were banded in Colorado from 1969 through 1981 as part of a large demographic and harvest management study (Braun 1972; Braun et al. 1975). This store of banding, live recapture, and dead recovery data is unmatched for this species. Although some of these data (1969–1974) have been summarized previously (Braun et al. 1975; Kautz and Braun 1981), we believe more information can be gleaned using additional capture–recapture analytical techniques (e.g., Burnham et al. 1987; Lebreton et al. 1992; Williams et al. 2002).

Our objectives were to estimate annual survival, fidelity, and harvest rate of band-tailed pigeons from this historic data set. These data, although collected more than 30 y ago, are extremely valuable because knowledge of the population biology for this species is limited. In addition, annual decisions regarding the regulation of harvest for band-tailed pigeons are based on limited information (Braun 1994; Keppie and Braun 2000).

Field methods

Band-tailed pigeons were captured at foraging areas in Colorado. Foraging locations were solicited through surveys of field personnel of the Colorado Division of Wildlife and U.S. Forest Service, contacts with landowners who were either feeding band-tailed pigeons at home sites or at harvesting (grain fields) or grain storage areas, and searches by project personnel starting in 1969 (Braun 1972, 1976). Reported foraging areas were visited to determine whether band-tailed pigeons were present and locate possible capture sites. Sites were baited with grain (depending upon what was available to band-tailed pigeons), and band-tailed pigeons were trapped mostly using cannon nets, but also with funnel and drop traps (Braun 1976). Techniques were developed to temporarily hold (in damp burlap sacks, portable wood cages) band-tailed pigeons for banding. All protocols for banding followed those of the Bird Banding Laboratory (U.S. Fish and Wildlife Service 1991, and earlier versions). Age and sex identification techniques were developed early (June 1969) in the banding process (Braun 1976), and sex was verified through collections. All band-tailed pigeons were released at the capture site. Band numbers as well as age and sex were recorded for all recaptured band-tailed pigeons.

Encounter data consisted of both live recaptures and dead recoveries. Live recaptures occurred only at banding sites within Colorado, whereas dead recoveries could come from anywhere in the species range. Individuals recovered from outside the study area may have been permanent or temporary emigrants, or individuals that died during migration or in wintering areas. We did not use individuals banded after 30 August because the hunting season began at this time. All data were archived and are publicly available from the Bird Banding Laboratory (administered by the U.S. Geological Survey). We present data arrays that summarize all capture, dead recovery, and live recapture (Table S1).

Estimating annual survival and fidelity

We used “live encounter-dead recovery” models (Burnham 1993) as implemented in program MARK to estimate annual survival and its variability (White and Burnham 1999). Model parameters were as follows: Si, probability that an individual survives from year i to i + 1; pi, probability of recapture of an individual in year i conditional on being alive and in the study area; ri, the probability of an individual being found dead in year i and having its band reported to the Bird Banding Laboratory, that is, reporting probability (Seber 1970); and Fi, the probability that an individual returns to the sampling area between year i and i + 1, given that it is alive, that is, fidelity. We hypothesized that annual survival and fidelity would vary by age (a: hatch-year [HY] vs. after-hatch-year [AHY]), sex (s), and time (t, independent annual estimates). We hypothesized that annual recapture and reporting probabilities would vary by age and time. There was no hunting season for band-tailed pigeons in Colorado from 1945 through 1969 because of low apparent numbers (Neff 1947; Braun et al. 1975). Thus, we hypothesized that reporting probability would be lower in 1969 than in other years and that reporting probability after 1969 would be constant. We used these competing hypotheses to formulate statistical models. Our most general model included all interactions between terms: Sa*s*t pa*t ra*t Fa*s*t. This model posits that survival varied by sex, age, and time. We used the general model with a median c-hat approach in program MARK to estimate overdispersion in the data set (White and Burnham 1999). This approach gave an estimate of the overdispersion parameter (ĉ) that we used as a variance inflation factor for all models. We used Akaike's Information Criterion adjusted for ĉ (AICc) to compare competing models (Burnham and Anderson 1998).

We began by modeling the first 6 y (1969–1974) because 89% of all data were from this time period. We used the top models (the top model and all models within 10 AICc units of the top model) from the analysis of the first 6 y to analyze the full 13-y data set. We first modeled age and time effects on p and r for the analysis of the 6-y data set, while leaving a general structure on S and F. We compared models for p and r with constant probability among ages and time periods: age only, time only, age by time, and age plus time. Age by time models allowed p (or r) to vary independently among age classes and years. Age plus time models allowed p (or r) to vary independently among years with an additive age effect (i.e., the difference in p or r between age-classes constant among years). We used the top ranked model structure for p and r as a basis for subsequent modeling of S and F. We compared models for S and F with constant probability among ages and time periods: age only, sex only, time only, age by time, sex by time, sex plus time, age plus time, and age by sex by time. We used a model with three categories when modeling sex (female, male, and unknown) because 2,312 individuals did not have sex assigned upon initial capture.

We followed this model building strategy with a few exceptions. Few HY individuals (n = 302) were banded after 1974, and it was unlikely that more complex models could be supported for this age-class during this time period. We created models with simpler structures (constant or age only survival) for the last 7 y for HY individuals when analyzing the 13-y data set. For example, we created models that allowed annual survival to vary among all years for AHYs, but only for the first 6 y for HYs. We did this for all parameters to make full use of these data.

Recovery rates

Inferences regarding harvest rate require an estimate of reporting rate (Nichols et al. 1991). The probability of a band being reported is a function of the banded animal being harvested, retrieved in the field, and having the band number reported to the Bird Banding Laboratory. If the retrieval and reporting rates are 100%, the proportion of banded birds reported is the harvest rate. Reward band studies are often used to estimate reporting rates (Bellrose 1955; Tomlinson 1968). To the best of our knowledge, reward band studies have never been conducted for band-tailed pigeons. Thus, we term our estimates “recovery rates,” and they are a product of harvest and reporting rates. Our estimates of annual recovery rates can be considered minimum estimates of harvest rate. In addition, inferences regarding differences among years or age-classes are valid as long as reporting rate did not differ among years or age-classes.

We used direct recoveries of birds banded from 1969 through 1974 as an index of annual harvest rates. A direct recovery occurs when a bird is harvested during the first hunting season after it was banded. We only used direct recoveries that were reported to the Bird Banding Laboratory as “shot.” We did not adjust the number of recoveries for reporting rate. We developed a small set of models to examine temporal and age-related patterns in direct recovery rate. The models were as follows: intercept only (constant recovery rate among years and age classes), time independent (recovery rate varied among years with no distinct pattern), age (HY recovery rate greater than AHY), and sex (female recovery rate different from male recovery rate). We also examined combinations of the above-mentioned models for age by time, sex by time, and age by sex by time. The basic model structure was logit(θ) = β0 + β1x, where θ is the probability that an individual was recovered given the value of x, x is a vector of explanatory variables (e.g., age of individuals), β0 is the intercept, and β1 is a slope parameter. We fit models using a logistic regression (Hosmer and Lemeshow 2000) with the LOGISTIC procedure in SAS version 9.2 of the SAS System for Windows. We ranked models based on AIC and used the top ranked model for inference.

Survival and fidelity

There were 26,490 band-tailed pigeons banded in Colorado during the course of this study. There were 3,586 recaptures (includes individuals recaptured more than once) and 780 recoveries of band-tailed pigeons due to harvest. We eliminated individuals banded after 30 August in a year, leaving 25,660 individuals (HY = 3,611; AHY = 22,044; unknown age = 5) that were used for the analysis of annual survival, fidelity, and harvest rates.

The best model structure (no other model within three AICc units) using only the first 6 y of data (Sa*t pt rt Ft) suggested survival varied between age-classes and years; and fidelity, recapture probability, and reporting probability varied among years. We kept the same general structure on p and r when using all years of data. Overdispersion within the data set was slight (ĉ = 1.11). The best model structure using all years of data suggested that survival varied between age-classes and among years for the first 6 y (Table 1; Figure 1). The best model structure for the last 7 y of study indicated survival varied between age-classes and varied among years for AHY birds (Figure 1). An intercept-only model was all that could be supported for the HY age-class for the last 7 y of study because too few HYs were banded during this time. Annual survival for the HY age-class over the last 7 y of study was 0.891 (SE = 0.116, 95% confidence interval [CI] = 0.439–0.989). The annual survival of HYs in some years seemed to be similar to AHYs (Figure 1), but the mean estimates of age-specific survival from the entire study period were SHY = 0.633 (SE = 0.031, 95% CI = 0.570–0.691), and SAHY = 0.719 (SE = 0.016, 95% CI = 0.687–0.748).

The top-ranked model suggested fidelity varied among years, but not between age-classes (Table 1). It seemed that in some years, fidelity was high (i.e., 100% of the band-tailed pigeons returned to Colorado), whereas in other years ∼30% of the birds did not return (Figure 2). Mean fidelity was 0.858 (SE = 0.029, 95% CI = 0.791–0.906) from 1969 to 1981.

Recovery rate

Sixty-two percent of banded birds that were harvested and reported were shot in Colorado. Of the remainder, 18% were reported shot in Mexico, 16% in New Mexico, 3% in Arizona, 1% in California, and <1% in both Utah and Washington. The top ranked model suggested that recovery rate varied among years and age (1969–1974, Figure 3). There were no models within 11 AIC units of the top model. The mean annual recovery rate for HYs was 0.015 (SE = 0.002, 95% CI = 0.012–0.020) and for AHYs was 0.011 (SE = 0.001, 95% CI = 0.010–0.012). There was no straightforward approach to include sex in the recovery rate modeling strategy because too many individuals were of unknown sex when banded. However, post hoc we compared the raw proportions of individuals recovered that did have sex identified upon capture, and we found no difference between sexes (female = 0.009, SE = 0.001, 95% CI = 0.008–0.011; male = 0.011, SE = 0.001, 95% CI = 0.009–0.013).

Our estimates of annual survival and recovery rate for the years 1969–1974 were similar to those reported by Kautz and Braun (1981). The point estimates for HY and AHY survival for the entire study were slightly lower than those reported by Kautz and Braun (1981), but well within the stated CIs for their estimates. In addition, our estimate of adult annual survival was similar to that of Coastal band-tailed pigeons reported by Casazza et al. (2015). Annual survival of AHYs seemed greater in the latter part of our study; however, the magnitude of uncertainty about these estimates precluded statistical significance.

We believe our study methods met the required assumptions needed for valid inference from banding-band return modeling (Brownie et al. 1985) and release–recapture modeling (Burnham et al. 1987), with a few minor exceptions. There was a small number of AHYs for which sex could not be assigned (n = 41 or 0.2% of the AHY sample). Ability to accurately identify the sex of HYs was much less certain (n = 2,271 or 75.4% of the HY sample). Thus, we only considered sex-specific models for birds banded in the AHY age-class. We created a separate “unknown” class for birds banded as HY (and the 41 AHYs of unknown sex) when modeling the effects of sex on survival and fidelity.

Some authors have suggested the nomadic nature of band-tailed pigeons was related to changing food supplies, whereas others have noted high site fidelity (Smith 1968; Braun 1972; Gutiérrez et al. 1975). On average it seemed that, conditional on surviving the year, ∼15% of band-tailed pigeons did not return to breed in Colorado. We estimated that in a few years, fidelity was exceptionally high (at or near 100% return), whereas in other years fidelity was near the average. Sample sizes were too small to accurately estimate annual fidelity from 1975 to 1981, but the point estimate for this time period was near the average. This relatively consistent level of fidelity may suggest nomadism related to food, but it may also suggest a simple interchange of individuals within a single population. It is also worth noting that ∼1% of band recoveries were from within the range of the Pacific Coast subspecies, indicating some interchange between the two subspecies.

Recovery rate estimates varied strongly among years, suggesting harvest rate may have varied among years. Similar to Kautz and Braun (1981), we also found recovery rate slightly greater for HYs than AHYs. How our estimates of recovery rate relate to actual harvest rates is unknown, but worth speculation. Braun (1972) reported that, based on hunter surveys, reporting rates from harvested birds were “high” due to publicity and hunter surveys in Colorado and New Mexico. Braun (1972) noted that many birds were also shot in Mexico, where there were no public surveys and that reporting rate was likely much lower than in the United States. However, because few band-tailed pigeons had been previously banded in the Arizona, Colorado, New Mexico, or Utah, banded pigeons were a novelty in Mexico, possibly enhancing their probability of being reported.

In a 1965–1966 study of the mourning dove, Zenaida macroura, from 15 widely separated areas in the United States, Tomlinson (1968) reported an estimated 32–66% of mourning dove bands retrieved were reported by hunters. The 66% reporting rate was associated with states with high publicity for their banding programs or states with conservation agencies that actively collected bands from hunters, whereas the 32% rate was what mourning dove hunters reported in general. Using reports of reward bands placed on birds in 1970–1971, Reeves (1979) estimated reporting probability for mourning doves in the Central Management Unit, which includes Colorado and New Mexico, as 0.45. Reeves (1979) considered this value as unaided by solicitation or publicity. As part of the Reeves (1979) study, Braun and Funk (1977) estimated the reporting rate for mourning dove bands from Colorado was 55.2%. In more contemporary studies (2003–2011) of mourning doves, Otis et al. (2008) and Sanders and Otis (2012) estimated U.S. band reporting probabilities (P) from “toll” (P = 0.495, SE = 0.015) and “web-address” (P = 0.535, SE = 0.021) bands (neither of which was used for our band-tailed pigeon study), and they concluded there was limited regional variation in reporting rates. Otis et al. (2008) used a reporting rate of 0.25 for Mexico, but they did not explain their rational. These band reporting studies suggest that during our study the reporting rate for band-tailed pigeons should have been high, as noted by Braun (1972), and possibly at or above the 66% number reported by Tomlinson (1968) for mourning doves shot in states that publicized banded mourning doves. If we use a reporting rate of 0.70 for the United States and 0.25 for Mexico (possibly too low based on novelty status), this would suggest that from 1969 to 1974 the mean annual harvest rate was ∼0.020 for HY and 0.014 for AHY band-tailed pigeons (overall mean = 0.015). If we use a reporting rate of 0.50 for the United States and 0.25 for Mexico, this would suggest that from 1969 to 1974 the mean annual harvest rate was ∼0.028 for HY and 0.020 for AHY band-tailed pigeons (overall mean = 0.021).

Hunters were required to obtain a special permit to hunt band-tailed pigeons from 1970 to 1974, thereby allowing managers to collect data on harvest (Braun et al. 1975). The estimated annual harvest of band-tailed pigeons in Colorado during the early years of our study was as follows: 0 in 1969; 541 in 1970; 1,723 in 1971; 820 in 1972; 363 in 1973; and 792 in 1974 (Pacific Flyway Study Committee and Central Flyway Webless Migratory Game Bird Technical Committee 2001). Because Colorado is at the northern end of the range for the Interior band-tailed pigeon subspecies, it is likely that all band-tailed pigeons in the harvest were from birds that spent the breeding season in the state. We can use these numbers, in conjunction with the harvest rate of birds in Colorado, to estimate abundance by using a Lincoln estimator (Lincoln 1930; Otis 2006; Alisauskas et al. 2011): Ni = Hi/hi, where Hi is total harvest in year i and hi is harvest rate in year i. Using reporting rates of 0.50 and 0.70, we estimated the annual harvest rates for band-tailed pigeons in Colorado from 1970 to 1974. These harvest rate estimates do not include birds harvested from outside of Colorado, nor do they include data from 1969 (when the season was closed). Thus, we estimate a range of annual abundance, with the range based on the two reporting rates. Annual abundance estimates on 1 September were (in the thousands): 62–88 for 1970, 39–54 for 1971, 57–80 for 1972, 58–98 for 1973, and 83–120 for 1974. The mean annual abundance was 59,911–88,290 for 1970–1974.

There are concerns about the status of band-tailed pigeons, primarily because standard methods for assessing trends in populations over time are lacking. Population level studies are also uncommon. Presently, annual hunting regulations are based on models informed with limited data. The data set we examined and the demographic estimates provide a glimpse of population demography for a now distant period (1969–1981) that may not relate well to the present. Advances in analytical methodology now need to be followed by advances in techniques for data collection, especially with regard to informing estimates of band-tailed pigeon abundance. Conservation of secretive migratory birds involves more than regulatory mechanisms as it is highly likely that habitats supporting secure nesting, brood rearing, and foraging through all seasons of the life cycle are key to maintaining and increasing populations of band-tailed pigeons.

It seems unlikely that the large-scale banding program in Colorado in 1969–1981 will be replicated, mainly for logistical reasons. In addition, we suspect that hunter interest and harvest rates have decreased, making sufficient recoveries of banded birds more difficult to obtain. To determine how fewer recoveries or less sampling effort might affect precision of estimates, we used the simulation routine in program MARK to estimate precision of survival under various scenarios (White and Burnham 1999). For simulations, we used a simple live encounter-dead recovery model (Sa2 p. ra2 F.), our parameter estimates (SHY = 0.633, SAHY = 0.719, p = 0.110, rHY = 0.037, rAHY = 0.034, F = 0.858), and a 6-y study. When annual sample sizes were 3,000 AHY and 600 HY (approximately what occurred during our study the first 6 y) coefficient of variation [CV](SHY) = 0.06 and CV(SAHY) = 0.04. When reporting probabilities were halved, CV(SHY) = 0.07 and CV(SAHY) = 0.05. When both reporting probability and sampling effort were halved, CV(SHY) = 0.10 and CV(SAHY) = 0.07. The simulation results suggest that studies with reduced effort may still be worthwhile, depending on desired precision of estimates. However, other methods to estimate demographic parameters or abundance (e.g., point counts) need to be weighed against the required effort.

Please note: The Journal of Fish and Wildlife Management is not responsible for the content or functionality of any supplemental material. Queries should be directed to the corresponding author for the article.

Table S1. Band-tailed pigeon Patagioenas fasciata live(L)-dead(D) recovery data used to estimate annual survival and fidelity, 1969–1981. Each row describes a unique capture history and the number of individuals by age-class and sex that share this capture history. The first column describes a capture history as LDLDLD, etc., for 13 yearly capture–recovery occasions. For example, row 1 represents individuals captured during occasion 1 that were recovered dead during occasion 1. Row 2 represents individuals that were initially captured during occasion 1, recaptured alive in both occasions 2 and 3, and never encountered again. The number of individuals that share a capture history by age-class (after-hatch-year [AHY], or hatch-year [HY]) and sex (female [F], male [M], or unknown sex [U]) is identified in columns 2–7. For example, two HY-U share the first capture history, one HY-U the second capture history, and one AHY-M the third capture history.

Found at DOI: http://dx.doi.org/10.3996/112015-JFWM-110.S1 (10 KB Text File).

Reference S1. Braun CE. 1976. Methods for locating, trapping and banding band-tailed pigeons in Colorado. Colorado Division of Wildlife, Special Report 39.

Found at DOI: http://dx.doi.org/10.3996/112015-JFWM-110.S2 (14.7 MB PDF).

Reference S2. Braun CE, Funk HD. 1977. Mourning dove studies in Colorado, 1964–1974. Colorado Division of Parks and Wildlife, Special Report XXX (Accepted).

Found at DOI: http://dx.doi.org/10.3996/112015-JFWM-110.S3 (637 KB PDF).

Reference S3. Braun CE, Brown DE, Pederson JC, Zapatka TP. 1975. Results of the Four Corners cooperative band-tailed pigeon investigation. Washington, D.C.: U.S. Department of the Interior, U.S. Fish and Wildlife Service Resource Publication 126.

Found at DOI: http://dx.doi.org/10.3996/112015-JFWM-110.S4 (8320 KB PDF).

Reference S4. Otis DL, Schulz JH, Scott DP. 2008. Mourning dove (Zenaida macroura) harvest and population parameters derived from a national banding study. Washington, D.C.: U.S. Department of Interior, U.S. Fish and Wildlife Service, Biological Technical Publication FWS/BTP-R3010-2008.

Found at DOI: http://dx.doi.org/10.3996/112015-JFWM-110.S5; also available at http://nctc.fws.gov/resources/knowledge-resources/pdf/mourningdove08.pdf (June 2016) (1291 KB PDF).

Reference S5. Pacific Flyway Study Committee and Central Flyway Webless Migratory Game Bird Technical Committee. 2001. Pacific and Central Flyways management plan for the Four Corners population of band-tailed pigeons. Portland, Oregon: Pacific Flyway Council, U.S. Fish and Wildlife Service.

Found at DOI: http://dx.doi.org/10.3996/112015-JFWM-110.S6; also available at http://pacificflyway.gov/documents/ibp_plan.pdf (June 2016) (381 KB PDF).

Reference S6. Sanders TA. 2014. Band-tailed pigeon population status, 2014. Washington, D.C.: U.S. Department of the Interior, U.S. Fish and Wildlife Service, Division of Migratory Bird Management.

Found at DOI: http://dx.doi.org/10.3996/112015-JFWM-110.S7; also available at: https://www.fws.gov/migratorybirds/pdf/surveys-and-data/Population-status/Band-tailedPigeon/Band-tailedPigeonPopulationStatus14.pdf (June 2016) (2979 KB PDF).

We thank all who reported locations of band-tailed pigeons in Colorado during 1969–1981, and especially private landowners for permission to access their lands for capture of band-tailed pigeons. Numerous field personnel of the Colorado Division of Wildlife facilitated the fieldwork. The banding efforts were aided by summer field assistants and multiple graduate students, including P.D. Curtis, J.E. Kautz, and J.A. White. The Accelerated Research Program for Migratory Webless Game Birds of the U.S. Fish and Wildlife Service provided the initial stimulus and financial support. We thank J.E. Kautz, R.E. Tomlinson, D.A. Haukos, M.L. Casazza, and B.A. Collier for reviewing earlier drafts of this manuscript. We thank K.H. Knudsen for help locating past reports. This study was supported throughout by the Colorado Division of Wildlife through Federal Aid to Wildlife Restoration Project W-88-R. H.D. Funk provided administrative support throughout the period of study.

Any use of trade, product, or firm names is for descriptive purposes only and does not imply endorsement by the U.S. Government.

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Author notes

Citation: Seamans ME, Braun CE. 2016. Estimation of band-tailed pigeon band recovery and population vital rates in Colorado, 1969–1981. Journal of Fish and Wildlife Management 7(2):369–376; e1944-687X. doi: 10.3996/112015-JFWM-110

The findings and conclusions in this article are those of the author(s) and do not necessarily represent the views of the U.S. Fish and Wildlife Service.

Supplemental Material