Mapping Nutritional Resources Across Landscapes: Why, How, Results Cow Summer Body Fat Dynamics 18 2.9 Kcal DE /g (64% DDM) Percent Body Fat 16 14 Began Nutrition Treatments Began nutrition treatments 12 2.65 Kcal DE/g (58% DDM) 10 8 6 2.40 Kcal DE/g (53% DDM) High Medium Low 4 2 March June October Probability of Breeding 1.0 0.8 100 80 300 60 20 0 (1/x) 290 Low Nutr. 280 0 20 Sep 4 Oct 18 Oct 1 Nov 27 Sep 11 Oct 25 Oct 8 Nov 1996 0.2 0.0 y = 266.7 x 1.5543 Medium Nutr. 40 0.6 1997 0.4 High Nutr. Breeding Date (Julian day) Cumulative Breeding (%) Autumn Breeding Dynamics 5 10 15 20 Total Body Fat (%) 25 270 260 5 10 15 20 Total Body Fat (%) 25 Calf Growth and Winter Survival 1997 140 100 High Nutr. 80 Days of Winter Survived (no.) Body Mass (kg) 120 100 80 60 40 60 40 20 y = -109.9 + 3.1656x - 0.0120088x Medium Nutr. 2 r 2 = 0.63; P < 0.001 0 60 80 100 120 140 120 140 1998 100 80 Low Nutr. y = 4.1 + 0.479x r 2 = 0.38; P < 0.001 60 40 20 Weaned 20 1 Jul 0 60 27 Aug 80 100 26 Oct 26Body Dec Mass (kg) Beginning 1995-1999 Summer Nutrition (DE) – Animal Performance Relationships Percent of maximum 120 Poor Poor Marginal Marginal Good Good Excellent Excellent <2.4 (53% DDM) 2.4-2.7 2.7-2.9 >2.9 (64% DDM) 100 80 60 40 20 0 Digestible energy of diets (kcal/gram of food) Calf growth Yearling growth Yearling preg Adult fat Adult preg Breeding times Winter survival Total body fat (%) 25 r 2 = 0.90; P < 0.001 20 15 10 5 0 1 2 3 4 5 6 LIVINDEX 7 8 9 Documenting Patterns of Nutritional Limitations in Wild Herds 22 elk herds in 5 states 12 years (1998 to 2010) Twice/year captures ~ 3,000 elk sampled Wild Elk Capture Coastal Data Collected: Body fat Washington Inland NW Montana Age Pregnancy status Lactation status Body mass Blood, urine, fecal samples Oregon Cascades Idaho Rocky Mtns Wyoming Colorado Autumn Body Fat of Lactating Wild Elk Cows Coastal Plains and Foothills Autumn body fat (%) 15 Cascade Foothills and Mountains Inland Northwest Central Rocky Mountains Adequate nutrition 10 5 Highly deficient nutrition 0 Rectangle denotes 1 herd; Each dot represents 1 year of data; Vertical lines are + 1 S.E. Pregnancy Rates of Lactating Cow Elk 66% 62% 95% 89% 64% 100% 86% 94% 78% 86% 89% 70% 80% 45% 76% 86% 82% 70% 89% 88% 72% 78% 83% Average Pregnancy Rate Pregnancy Rates by Lactation Status 1 0.9 0.8 All Cows (Age 3-13) 0.7 0.6 0.5 0.4 0.3 Lact Cows (Age 313) Reject the hypothesis that nutritional influences are of little or no practical relevance and the hypothesis that summer nutrition is ubiquitously satisfactory and thus is of little or no practical relevance Instead, The relevant questions are how severe, where, why, and what can be done about it? Digestible Energy Content of Consumed Forage: Our Nutrition Variable for Resource Mapping DE plays a key, usually dominant role in virtually all life processes . There is considerable precedence…National Research Council presents livestock nutritional requirements guidelines several ways, including DE content of food. Integrating Plant Succession, Disturbance, and Elk Nutrition Habitat predictor variables Stand/site characteristics Stand age Animal response variables Understory vegetation characteristics Canopy cover Animal foraging outcomes Bite mass Composition Bite rate BA, QMD, TPH, HT Repro, Surv Biomass Season DDP, DDE DP, DE Climate Foraging time Palatability Soils Performance outcomes Integrating Plant Succession, Disturbance, and Elk Nutrition Macroplots: n = 349 total n = 89 w/elk Nooksack S.A. Willipa Hills S.A. Springfield S.A. Examples of Accepted/Avoided Forage Accepted Forage Selected Forage Neutral Forage Avoided Forage Deciduous shrubs: Big leaf maple Hazelnut Cascara Forbs: Queen’s cup beadlily Northern bedstraw False Solomon’s seal Oxalis Most grasses Deciduous shrubs: Alder Elderberry Salmonberry Many forbs Ferns: Lady fern Evergreen shrubs: Salal Oregon grape Rhododendron Ferns: Deerfern Swordfern Most conifers Digestible Energy of Elk Diets Digestible energy (kcal/g) 3.4 3.2 3.0 2.8 2.6 Maintenance = 2.7 kcal/g 2.4 2.2 MHZ & PSFZ WHZ 2.0 1.8 0 300 600 900 1200 Neutral + selected biomass (kg/ha) 1500 TSME & ABAM Habitats All Seasons All Study Areas TSHE Habitats Autumn All Study Areas TSHE Habitats Summer All Study Areas 3.2 3.2 3.2 3.2 2.8 2.8 2.8 2.8 2.4 2.4 2.4 2.4 A 2.0 0 400 800 1200 2.0 1600 0 Accepted biomass (kg/ha) B 400 2.0 1200 0 800 Accepted biomass (kg/ha) TSHE Habitats All Seasons Nooksack Digestible energy (kcal/g) Digestible energy (kcal/g) TSHE Habitats All Seasons All Study Areas 400 800 1200 1600 Accepted biomass (kg/ha) TSHE Habitats All Seasons Willipa Hills 1 3.2 C D 2.0 0 400 TSHE Habitats All Seasons Springfield 3.2 3.2 1 2 3 2 2.8 2.8 3 2.8 4 2.4 2.4 E 2.0 0 400 800 1200 Accepted biomass (kg/ha) 2.4 4 F 2.0 0 400 800 1200 1600 Accepted biomass (kg/ha) G 2.0 0 300 800 1200 Accepted biomass (kg/ha) 600 900 Accepted biomass (kg/ha) Equations to Predict Dietary Digestible Energy (DDE) n r2adj P MHZ & PSZ Habitats, All Study Areas DDE = a + b1(NeuB) + b2(SelB) – b3(NeuB*SelB) 14 0.43 0.036 WHZ Habitats, Nooksack DDE = a + b1(NeuB) + b2(SelB) – b3(NeuB*SelB) 19 0.39 0.015 WHZ Habitats, Willipa Hills DDE = a + b1(NeuB) + b2(SelB) – b3(NeuB*SelB) 26 0.46 0.001 WHZ Habitats, Springfield DDE = a + b1(NeuB) + b2(SelB) – b3(NeuB*SelB) 28 0.57 0.0001 Equationsa,b a NeuB = Neutral spp biomass; SelB = Selected spp biomass b P < 0.075 individually for each predictor variable. Overstory canopy cover (%) Overstory canopy cover (%) Overstory Canopy Cover Development 100 80 60 Western hemlock zone 40 20 0 0 100 200 300 400 500 600 700 100 80 Pacific fir zone 60 40 Mountain hemlock zone 20 0 0 100 200 300 Stand age (years) 400 500 Understory Production (kg/ha) Understory Production (kg/ha) 3,500 Springfield, OR 3,000 Conifer E. Shrub For. Ferns PTAQ Sedge Grass Forb D.Shrub 2,500 2,000 1,500 1,000 500 0 1 6 9 3,500 14 18 21 24 30 34 37 40 44 55 58 63 73 92 153 307 479 642 Springfield, OR 3,000 Age > 50 years, Mean “accepted” biomass = 50 kg/ha 2,500 2,000 Avoided Neutral Selected 1,500 1,000 500 0 1 6 9 14 18 21 24 30 34 37 40 44 55 58 63 73 92 153 307 479 642 Stand Age (years) Digestible Energy of Elk Diets Digestible energy (kcal/g) 3.4 Age > 50 years, Mean “accepted” biomass = 50 kg/ha 3.2 3.0 2.8 Maintenance 2.6 2.4 2.2 MHZ & PSFZ WHZ 2.0 1.8 0 300 600 900 1200 Neutral + selected biomass (kg/ha) 1500 Equations to Predict Forage Biomass Groups Equationsa,b n r2adj P MHZ & PSZ Habitats, All Study Areas AccB = a - b1(CC) + b2(CC2) + b3(HWd) 70 0.70 <0.001 NeuB = a - b1(CC) + b3(HWd) 70 0.72 <0.001 SelB = 1/(a + b1*CC) 70 0.30 <0.001 a AccB = accepted spp biomass; NeuB = neutral spp biomass; SelB = selected spp biomass; CC = canopy cover; HWd = hardwood stems. b P < 0.05 individually for each predictor variable. Equations to Predict Forage Biomass Groups n r2adj AccB = a - b1(CC) + b2(CC2) + b3(HWd) 276 0.68 <0.001 NeuB = a - b1(CC) + b2(CC2) + b3(HWd) 276 0.65 <0.001 SelB = a - b1(CC) + b3(HWd) 276 0.46 <0.001 Equationsa,b P WHZ Habitats, By Study Area a AccB = accepted spp biomass; NeuB = neutral spp biomass; SelB = selected spp biomass; CC = canopy cover; HWd = hardwood stems. b P < 0.05 individually for each predictor variable. Nutrition Succession Trajectories Western Hemlock-Salal Habitats Dietary DE (kcal/g) 3.4 Nooksack 3.2 3.0 Willipa Hills 2.8 Springfield Maintenance 2.6 2.4 2.2 0 50 100 150 200 250 300 350 400 450 500 550 600 650 Stand Age (years) Nutrition Succession Trajectories Western Hemlock-swordfern Habitats Dietary DE (kcal/g) 3.6 Nooksack 3.4 3.2 Hardwood-riparian 3.0 2.8 Maintenance Springfield Willipa Hills 2.6 2.4 2.2 0 50 100 150 200 250 300 350 Stand Age (years) 400 450 500 550 600 Nutrition Succession Trajectories Mountain Hemlock & Pacific Fir Habitats Dietary DE (kcal/g) 3.4 Nooksack 3.2 3.0 2.8 Willipa Hills Maintenance Springfield 2.6 2.4 0 50 100 150 200 250 300 Stand Age (years) 350 400 450 500 The Maps Pacific Fir & Mtn Hemlock Zones Location: Preg; Autumn BF Nooksack: 95%; 13% Forks: 62%; 6% Wynoochee: 53%; 6% Green River: 89%; 10% White River: 94%; 11% Willapa Hills: 78%; 6% Mount St. Helens: 68%; ~5.5% Suislaw, Untr.: 18%; ~2.9% Impr.: 41%; ~3.9% Springfield: 80%; 10% Steamboat: 88%; 7% Toketee: 72%; 9% Location: Preg%; Autumn BF% Nooksack: 95%; 13% Forks: 62%; 6% Wynoochee: 53%; 6% Green River: 89%; 10% White River: 94%; 11% Willapa Hills: 78%; 6% Mount St. Helens: 68%; ~5.5% Suislaw, Untr.: 43%; ~2.9% Impr.: 49%; ~3.9% Springfield: 80%; 10% N. Umpqua R.: 80%; 8% Habitat Attributes-11 Herds I. DDE Nutrition Categories Category kcal/g of forage Poor Low Marginal High Marginal <2.45 2.45 to 2.59 2.60 to 2.74 Low Good High Good 2.75 to 2.89 2.90 to 3.04 Excellent >3.05 ______________________________________________ II. Forest Overstory Canopy Cover: <35%; 35-70%; >70% III. Forest zones: High elev. including Alpine, MHZ, PSZ Low elev. including WHZ, DFZ, SSZ Herd-level Nutritional Resources 100 Percent of area 80 <2.45 2.45-2.60 2.60-2.75 2.75-2.90 2.90-3.05 >3.05 60 40 20 0 A A A A A A A R R R R O O O O W W W W W W W , , , , . ., ., d r. r. s, e, k, s, s, l l R t R R k e c l p n e i n r fi le o sa ch ua Im en it e g e F ,U k , o q aH e h n H o r i o p w p W n G t. aw la la m pr l y l No S s i s S U i i . W W N. Su Mt Su 100 100 90 90 90 80 80 80 70 60 Pregnant (%) 100 Pregnant (%) Pregnant (%) Herd-level Pregnancy & DDE 70 60 70 60 50 50 50 40 40 40 30 30 0 1 02 33 30 4 065 r 2 = 0.67 5 69 r 2 = 0.73 r 2 = 0.64 10 7 12 15 15 20 25 of >2.90 areaPercent with DDEwith >2.60 DDE Percent ofPercent area with DDE>2.75 of area > High good > Low good > High marginal 30 Herd-level Pregnancy & DDE 100 x1 Pregnant (%) 90 15% >2.75 DDE 12% >2.75 DDE 80 9% >2.75 DDE 70 6% >2.75 DDE 60 3% >2.75 DDE 50 0% >2.75 DDE 40 0 Coastal or Cascade foothill areas lacking high elevation habitats 5 10 15 20 25 30 Percent of area with 2.6 to 2.75 DDE (x 2) 35 Preg (%) = 39.3 + 3.03(x1) + 1.48(x2), where x1 = % of area >2.75 DDE (P = 0.0004) 2 R adj x 2 = % of area 2.6 to 2.75 DDE (P = 0.0109) = 0.83, P = 0.0009 Herd-level Pregnancy & DDE 100 Pregnant (%) 90 x1 15% >2.75 DDE 12% >2.75 DDE 80 9% >2.75 DDE 70 6% >2.75 DDE 60 3% >2.75 DDE 50 0% >2.75 DDE 40 0 Coastal or Cascade foothill areas lacking high elevation habitats 5 10 15 20 25 30 Percent of area with <35% Can Cov (x 2) 35 Preg (%) = 39.3 + 3.06(x1) + 1.50(x2), where x1 = % of area >2.75 DDE (P = 0.0001) 2 R adj x 2 = % of area <35% Can Cov (P = 0.0023) = 0.87, P = 0.0002 Herd-level Body Fat & DDE Ingesta-free body fat (%) 14 12 10 8 6 4 r 2 = 0.65 2 0 3 6 9 12 15 Percent of area with >2.75 DDE 14 12 Ingesta-free body fat (%) Ingesta-free body fat (%) 14 10 8 6 4 12 10 8 6 4 2 r 2 = 0.44 r = 0.71 2 0 1 2 3 4 5 6 Percent of area with >2.90 DDE 7 2 0 5 10 15 20 25 Percent of area with >2.60 DDE 30 Herd-level Body Fat & DDE Ingesta-free body fat (%) 15 x1 12 15% >2.75 DDE 12% >2.75 DDE 9 9% >2.75 DDE 6% >2.75 DDE 6 3% >2.75 DDE 0% >2.75 DDE 3 0 Coastal or Cascade foothill areas lacking high elevation habitats 5 10 15 20 25 30 Percent of area with 2.6 to 2.75 DDE (x 2) 35 IFBF(%) = 3.54 + 0.512(x1) + 0.130(x2), where x1 = % of area >2.75 DDE (P = 0.0047) 2 R adj x2 = % of area 2.6 to 2.75 DDE (P = 0.28) = 0.66, P = 0.014 Herd-level Body Fat & DDE Ingesta-free body fat (%) 16 14 x1 12 15% >2.75 DDE 10 12% >2.75 DDE 8 9% >2.75 DDE 6% >2.75 DDE 6 4 2 3% >2.75 DDE 0% >2.75 DDE 0 Coastal or Cascade foothill areas lacking high elevation habitats 5 10 15 20 25 30 Percent of area with <35% Can Cov ( x 2) 35 IFBF(%) = 3.28 + 0.518(x1) + 0.152(x2), where x1 = % of area >2.75 DDE (P = 0.0031) 2 R adj x2 = % of area <35% can cov (P = 0.15) = 0.70, P = 0.0087