INFLUENCES OF SELECTED WEATHER VARIABLES ON PREDATION OF WILD TURKEY FEMALES AND NEST SUCCESS D. Kevin Lowrey 1,2 Department of Wildlife and Fisheries, Box 9690, Mississippi State University, Mississippi State, MS 39762, USA Stan R. Priest 3 Department of Wildlife and Fisheries, Box 9690, Mississippi State University, Mississippi State, MS 39762, USA George A. Hurst Department of Wildlife and Fisheries, Box 9690, Mississippi State University, Mississippi State, MS 39762, USA Brandon S. Weemy Department of Wildlife and Fisheries, Box 9690, Mississippi State University, Mississippi State, MS 39762, USA Abstract: We explored relationships among weather, predation, and wild turkey ( Meleagris gallopavo sylvestris ) female nest success in Mississippi. Females ( n 82) were monitored via telemetry during the nesting period on Tallahala Wildlife Management Area from 1991–95. We compared annual nest success rates to cumulative rainfall and to number of rainfall events, April–May and March–June of each year. Cumulative rainfall was not significant ( P 0.188); however, number of rainfall events was negatively correlated ( r – 0.90, P 0.04) to annual nest success rates during both periods. Fewer rainfall events were associated with higher nest success rates. Weather variables were measured near nests ( 50 m) in sympatric habitats and compared between successful ( n 8) and unsuccessful females ( n 24). Maximum temperature ( P 0.01), number of rainfall events ( P 0.002), and total rainfall ( P 0.002) were significantly different between successful and unsuccessful females. Successful females had higher maximum temperatures at the nest site, fewer rainfall events, and less cumulative rainfall. Weather variables associated with known dates of predation of females ( n 19) were compared with weather variables associated with randomly selected dates ( n 19). Days since rain ( P 0.005) and amount of rainfall on the date of predation ( P 0.003) were significantly different from random dates. It rained on the date a female was preyed upon or 1– 8 days before predation, differing significantly from random dates. We suggest land managers be aware of influences of weather on predation, understand that we cannot manage weather conditions, and explore alternate methods of increasing nest success. Proceedings of the National Wild Turkey Symposium 8:173–178 Key words: Meleagris gallopavo sylvestris, Mississippi, nest success, predation, rainfall, weather, wild turkey. In many studies of wild turkey productivity, weather has been identified as a density-independent factor that may cause turkey populations to fluctuate (Stoddard 1963, Healy and Nenno 1985, Lint 1990). Porter et al. (1990) suggested that weather, not harvest, regulated turkey populations in New York. Several studies addressed effects of weather on poult survival rate (Williams et al. 1973, Healy and Nenno 1985) and nest success rate (Priest 1995, Lowrey 1999). Other studies concentrated on how weather events affected timing of reproductive events in the spring, resulting in poor female condition during the nesting season (Korschgen 1 E-mail address: kevin_lowrey@deq.state.ms.us 2 Present address: 468 Mockingbird Lane, Madison, MS 39110, USA. 3 Present address: Anderson–Tully Company, Vicksburg, MS 39181, USA. 173 1967, Vangilder et al. 1987, Hurst 1992). Many of these studies reported predation to be a more important factor in wild turkey production than weather (Williams et al. 1973, Healy 1992). Some studies suggested a relationship be- tween predation and weather (Siess 1989, Priest 1995, Rob- erts and Porter 1998). Our objectives were to determine if annual nest success is related to number of rainfall events and cumulative rainfall amounts during the nesting period, to compare weather variables associated with successful nests to unsuccessful nests, and to determine if rainfall amounts and patterns affect predation of nesting and non- nesting female wild turkeys throughout the year. STUDY AREA The study area was the 14,410-ha (35,592-ac) Talla- hala Wildlife Management Area (Tallahala WMA) located within the Bienville National Forest in parts of Scott, New- ton, Smith and Jasper counties, Mississippi. The area is in the Hilly Coastal Plain Province and the Blackland Soil Resource Area (Pettry 1977). Climate was mild with a mean annual temperature of 18 ° C (65 ° F) and mean annual pre- cipitation of 144 cm (56.16 in). The area averaged 200 – 230 frost-free days per year. Tallahala WMA was comprised of 37% mature pine ( Pinus spp.) stands, 17% pine – hardwood stands (30 – 70% pine), and 11% pine regeneration areas. Loblolly pine ( Pinus taeda ) predominated, but shortleaf pine ( P. echinata ) and longleaf pine ( P. palustris ) also occurred in upland sites, and spruce pine ( P. glabra ) was found in bottomland sites. Home sites and forest openings made up 5% of the area and the remaining 30% of the area was occupied by mature ( 80 yr old) bottomland hardwood stands. METHODS Turkeys were captured with cannon nets from early January to early March (winter) and from early July to late August (summer), 1991 – 95 (Bailey et al. 1980). Turkeys were processed and released at their capture site. Each turkey was classi fi ed as an adult or subadult based on molt pattern (Williams 1981). Females were indi- vidually marked with patagial wing tags (Knowlton et al. 1964) and numbered leg bands. A transmitter weighing 107 g (3.7 oz.) and operating in the 150 – 152 MHz range (Wildlife Materials, Carbondale, Illinois, USA) was fi tted “ backpack ” style to each female (Williams et al. 1968, Everett et al. 1979). Females ( n 82) were monitored by telemetry during 1991 – 95. Weather stations were set up approximately 50 m (55 yd) from incubating females ( n 32) in sympatric habitat. Weather variables were measured using a soil ther- mometer, rain gauge, maximum and minimum thermome- ter, and hydrometer. Soil thermometers were placed in the ground approximately 10 cm (4.0 in) deep and left for the duration of incubation. Soil temperature was recorded at 1200 hr each day. Thermometers and hydrometers were placed at 30 cm (11.8 in) above ground to simulate average height of incubating females and some potential predators. Stations were checked at approximately 6-hr intervals (i.e., 0600 hr, 1200 hr, 1800 hr, 2400 hr). Weather data were also obtained from the Mississippi Agriculture Experiment Sta- tion (MAES) near Newton, Mississippi, and were used in some analyses. The MAES station was located about 20 km (12.42 mi) northeast of the study area. Data from the MAES were consistent with weather data experienced on Tallahala WMA (Miller 1997). Annual nest success rates were compared to number of rainfall events and amount of rainfall during the incubation period (Apr – May). For the purpose of this study, nest suc- cess equaled the number of successful nests divided by the number of nests attempted. While some studies utilized a pattern of localized movements to determine a nest attempt, our study conservatively required a female to begin incu- bation before it was considered a nest attempt. A nest was determined to be successful if it hatched at least 1 egg. Nest site weather data and nest success data on wild turkey females from 1991 – 92 were obtained from Priest (1995). We repeated the same analyses for the entire nesting season (Mar – Jun). Spearman rank correlation analyses (SAS Insti- tute 1992) were performed to determine if number of rain- fall events and amount of rainfall were correlated to nest success. We also compared weather variables associated with successful females to unsuccessful females. Nest site weather station data were averaged for each female ’ s indi- vidual incubation period and pooled across years. A Kruskal – Wallis test was used to compare each variable Higher temperatures, fewer rainfall events, and lower rainfall amounts were associated with successful turkey nests. ( W. Porter ) 174 Habitat, Weather, and Population Dynamics between successful and unsuccessful nests (SAS Institute 1992) at the 0.05 signi fi cance level. Finally, we used the MAES data to compare rainfall amounts and patterns between days on which predation of females occurred ( n 19) and random days ( n 19) from 1991 – 95. Known predation events throughout the year, not just the nesting season, were used in the analyses. Variables associated with the date of predation and vari- ables associated with randomly selected dates were com- pared using a Kruskal – Wallis test (SAS Institute 1992). RESULTS Nest success averaged 20.8% and ranged from 13% in 1991 to 27% in 1992 over the 5 study years (Tables 1, 2). We found no correlation between total rainfall during April – May or March – June and annual nest success. We found a negative correlation between number of rainfall events during April – May and March – June and annual nest success (Table 3). Maximum temperature, number of rainfall events, and amount of rainfall were different between successful ( n 8) and unsuccessful ( n 24) females (Table 4). Remaining variables were not related ( P 0.05). Higher tempera- tures, fewer rainfall events, and lower rainfall amounts were associated with successful nests (Table 4). Predation of females was associated with lower num- ber of days since rain and higher amounts of rainfall on the day of predation. Days since rainfall and amount of rainfall on date of predation were signi fi cantly different between days on which females were depredated and random days (Table 5). All remaining variables were not signi fi cant ( P 0.05). DISCUSSION The role of rainfall in nest success is not completely understood (Healy and Nenno 1985). Many land managers have speculated that certain weather events may affect wild turkey productivity. We found that annual nest success was not signi fi cantly correlated with total amount of rainfall. This result contrasted with other studies emphasizing total rainfall amounts as the primary environmental in fl uence on nest success (Pattee and Beasom 1980, Roberts and Porter 1998). However, Priest (1995) found no correlation between nest success and total rainfall amount on Tallahala WMA. Although total rainfall was not correlated to annual nest success, there was a signi fi cant difference in the amount of rainfall between successful and unsuccessful nests. Less rainfall was observed at successful nests than unsuccessful nests. Perhaps successful females selected habitats that pro- vided more overhead cover accounting for lower rainfall totals. Nest sites of successful females on Tallahala had less canopy closure than those of unsuccessful females, but had more vegetative cover in the 0 – 0.9 m (0 – 30 in) range (Lowrey 1999). There could have been some spatial varia- tion between the location of the nest and the weather station; however, every effort was made to reduce this possibility by locating the weather station in similar habitat. Stand type could affect the amount of rainfall reaching ground level; however, there was no signi fi cant site effect in relation to stand type. Pine and hardwood basal areas were similar for successful and unsuccessful nest sites (Lowrey 1999). Priest (1995) suggested that a given amount of rainfall on 1 day may be less important than the same amount spread across several days. Rolley et al. (1998) reported the decline in turkey recruitment in a Wisconsin study was due to cold and wet spring weather. In New York, Roberts and Porter (1998) found that nest survival was negatively asso- ciated with both daily rainfall and cumulative departure from normal seasonal rainfall. In our study, the number of rainfall events was signi fi cantly correlated to nest success and was signi fi cantly different between successful and un- successful nests. This result suggests the importance of rainfall patterns on turkey productivity. The number of times it rains during the incubation period may be more critical to nest success than the total amount of rainfall. These data deal with a relatively small sample of successful females; however, further research may yield a predictive model based on number of spring rainfall events. Average relative humidity was slightly higher for suc- cessful nests; however, relative humidity was not signi fi - cantly different between successful and unsuccessful nests. Priest (1995) also found no signi fi cant difference in relative humidity between successful and unsuccessful nests on Tal- Table 2. Total rainfall amounts, total number of rainfall events, and nest success rates on Tallahala WMA, Mississippi, for March – June 1991 – 95. Year Nest success rate (%) Rainfall (cm) Number of rainfall events 1991 13 88.01 61 1992 27 38.37 35 1993 17 57.59 38 1994 22 43.15 39 1995 25 58.77 36 Table 1. Total rainfall amounts, total number of rainfall events, and nest success rates on Tallahala WMA, Mississippi, for April and May 1991 – 95. Year Nest success rate (%) Rainfall (cm) Number of rainfall events 1991 13 61.29 41 1992 27 10.48 15 1993 17 24.31 20 1994 22 20.10 15 1995 25 9.17 15 Weather Influences on Predation Lowrey et al. 175 lahala WMA. High relative humidities were associated with warm nights and increased predator movement on Tallahala WMA (Wilson 1997). Maximum temperature was signi fi cantly higher for successful nests than unsuccessful nests. Priest (1995) found no signi fi cant difference between successful and un- successful nests but reported successful nests were associ- ated with higher maximum temperatures. Minimum temper- atures did not differ. In contrast, Roberts and Porter (1998) found daily nest survival was positively associated with heating degree days, in a southcentral New York study. Wildlife managers could consider the importance of aspect (e.g., south facing) and basal area in their forest manage- ment plans. More sunlight would increase temperatures of potential nest sites and may increase nest success. Predation of nests and nesting females has been well documented on Tallahala WMA and throughout the United States. Some of the lowest nest success rates ever reported were from Tallahala WMA (Miller 1997). Perhaps the conversion of hardwood stands to pine plantations is creat- ing less quality nesting habitat while providing ideal habitat for predators. Palmer et al. (1993 b ) stated that low repro- duction on Tallahala WMA was due to predation of nests and that this predation was related to the last rainfall event. Miller et al. (1998 a ) summarized 13 years of reproductive data on Tallahala WMA and found that 51.1% of nest failures were due to predation. Miller et al. (1998 b ) also found that predation was the main mortality factor for female wild turkeys on Tallahala WMA. In another Missis- sippi study, Palmer et al. (1993 a ) found that 95% of female wild turkey mortality was due to predation. Roberts et al. (1995) found that predation accounted for more annual mortality than poaching, hunting, and wounding combined. In western Massachusetts, predation accounted for 92% of nest losses (Vander Haegen et al. 1988). Predation is thought to be a more important factor in nest success than weather conditions (Speake 1980, Palmer et al. 1993 b ). We suggest that relationships between the 2 factors are impor- tant to female productivity and survival. In our study, 19 females were known to be killed by predators. It rained on the days 10 of these birds were killed. The remaining birds had a rain event 1 – 8 days before their death. This result was signi fi cantly different from random days selected during the same periods. Rainfall amounts on the date of predation were higher than rainfall amounts on random dates. When turkeys are wet, they may be more detectible by predators (Palmer 1990, Roberts and Porter 1998). Past research has shown that predation exerts a great in fl uence on turkey productivity. Our results suggest a strong relationship between predation and certain weather variables. Not only do we know that predation lowers nest success rates, our results indicate that certain weather events Table 3. Results of Spearman rank correlation analyses for amount of rainfall and number of rainfall events versus annual nest success, on Tallahala WMA, Mississippi, 1991 – 95. Variables Mar – Jun Apr – May r S P – value r S P – value % success vs. rainfall – 0.70 0.188 – 0.70 0.188 % success vs. rainfall events – 0.90 0.037 – 0.89 0.040 Table 4. Summary of Kruskal – Wallis test results for differences between weather variables associated with successful and unsuccessful nests on Tallahala WMA, Mississippi, 1991 – 95. Variable a Fate b Mean SE Test statistic P – value RELHUM U 56.8 3.07 1.88 0.17 S 61.2 2.77 MAXTEMP U 78.5 1.06 6.15 0.01 c S 82.8 0.99 MINTEMP U 54.4 1.12 0.21 0.65 S 52.9 2.19 SOILTEMP U 65.7 0.18 2.07 0.15 S 65.9 0.95 RAINFALL U 3.96 0.58 13.8 0.002 c S 1.00 0.25 EVENTS U 5.08 0.47 13.8 0.002 c S 1.63 0.18 aRELHUM relative humidity, MAXTEMP maximum daily temperature, MINTEMP minimum daily temperature, SOILTEMP daily soil temperature, RAINFALL total amount of rain, EVENTS number of days with a measurable amount of rainfall. bU unsuccessful nest, S successful nest. cSigni fi cant difference at 0.05. Table 5. Summary of Kruskal – Wallis test results for differences between weather variables associated with days when females were preyed upon and weather variables associated with random days on Tallahala WMA, Mississippi, 1991 – 95. Variable Class e Mean SE Test statistic P – value DAYS a R 4.15 0.879 8.06 0.004 f P 1.42 0.515 AMOUNT b R 0.05 0.010 8.92 0.003 f P 1.22 0.216 DAYS4 c R 0.94 0.222 1.41 0.234 P 1.42 0.279 AMOUNT4 d R 2.23 0.251 0.01 0.905 P 2.49 0.376 DAYS8 c R 2.05 0.370 1.03 0.308 P 2.57 0.361 AMOUNT8 d R 4.57 0.529 0.29 0.588 P 3.38 0.442 DAYS20 c R 6.00 0.616 0.27 0.605 P 6.26 0.687 AMOUNT20 d R 10.4 0.778 0.90 0.343 P 8.40 0.783 aDAYS number of days since rain. bAMOUNT Amount of rain (cm) on date. cDAYS4, DAYS8, DAYS20 number of rainfall events at 4, 8, and 20 days prior to date. dAMOUNT4, AMOUNT8, AMOUNT20 total amount of rain (cm) at 4, 8, and 20 days prior to date. eClass depredated (P) and random (R). fSigni fi cant difference at 0.05. 176 Habitat, Weather, and Population Dynamics may be enhancing predators ’ ability to destroy nests and kill turkeys. This is important when considering ways to in- crease nest success: habitat management alone may not be enough. MANAGEMENT IMPLICATIONS Successful females were associated with less rainfall, fewer rainfall events, and warmer temperatures; however, managers cannot control any of these weather conditions. Predation of nests (eggs) and females was enhanced by certain weather conditions. Perhaps predator management via trapping and hunting could be used to reduce predation rates and increase nest success. Since we cannot manage weather conditions and predator management may not be a practical option, we must focus our attention to habitat management. Managers may use silvicultural practices (e.g., prescribed burning) to enhance habitat conditions for nesting and lessen the impact of predation. ACKNOWLEDGMENTS We thank the following for their support of the project: Mississippi Department of Wildlife, Fisheries and Parks through Federal Aid In Wildlife Restoration, W – 30; U.S. Forest Service; National Wild Turkey Federation (NWTF); and Mississippi Chap- ter NWTF. This study was a part of the Mississippi Cooperative Wild Turkey Research Project. We also appreciate the editorial comments provided by fellow researchers. We operated under Mississippi State University Animal Care and Use Committee (IACUC) 93 – 030. LITERATURE CITED Bailey, W., D. Dennet, H. Gore, J. Pack, R. Simpson, and G. Wright. 1980. Basic considerations and general recom- mendations for trapping the wild turkey. Proceedings of the National Wild Turkey Symposium 4:10 – 23. Everett, D. D., D. W. Speake, and W. K. Maddox. 1979. Wild turkey ranges in Alabama mountain habitat. Pro- ceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies 33:233 – 238. Healy, W. M. 1992. Population in fl uences: environment. Pages 129 – 143 in J. G. Dickson, editor. The wild turkey: biology and management. Stackpole Books, Harrisburg, Pennsylvania, USA. ——— , and E. S. Nenno. 1985. Effect of weather on wild turkey poult survival. 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Pro- ceedings of the Annual Conference of the Southeastern Association of Game and Fish Commissioners 22:16 – 30. Wilson, T. S. 1997. Raccoon and opossum home ranges, movements, and habitat use in a managed forest of central Mississippi. Thesis, Mississippi State Univer- sity, Mississippi State, Mississippi, USA. Don Kevin Lowrey (pictured) gradu- ated from Clemson University in 1992 with a B.S. in Aquaculture, Fisheries, and Wildlife Biology. He received a M.S. in Wildlife Ecology from Mississippi State University (MSU) in 1999. This research repre- sents part of his thesis work at MSU. Along the way he has held several seasonal positions, the most recent with Ducks Unlimited. Currently he lives in Brandon, MS, with his wife and 3-year old son. His interests in- clude upland game management, wa- terfowl management, and animal damage control. He is also an avid hunter and fi sherman. George Hurst is a graduate of North Carolina State University (M.S.) and Mississippi State University (Ph.D.). He recently retired from teaching and research at Mississippi State University. His research emphasis has included bobwhite quail, white-tailed deer, and wild turkeys. Much of his career was spent researching wild turkeys. He coordinated the Mississippi Cooperative Wild Turkey Research Project which was initiated in 1983 and ended in 1998. Stan R. Priest received a B.S. in forestry and M.S. in wildlife ecology from Mississippi State University. He has been working as a wildlife biologist for Anderson-Tully Co. in Vicksburg, Mississippi, since 1994. Brandon S. Weemy is a graduate of South Dakota State University (B.S.). He is currently working on his M.S. degree in wildlife sciences at Mississippi State University. 178 Habitat, Weather, and Population Dynamics