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The ecology of Richardson’s Merlins on the Canadian prairies Hodson, Keith Alan 1976

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THE UNIVERSITY OF BRITISH COLUMBIA THE ECOLOGY OF RICHARDSON'S MERLINS ON THE CANADIAN PRAIRIES BY KEITH ALAN HODSON B . S c , U n i v e r s i t y of B r i t i s h Columbia, 1970 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE I n the Department of P l a n t Science We accept t h i s t h e s i s as conforming to the req u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA 1976. In p resent ing t h i s t he s i s in p a r t i a l f u l f i l m e n t o f the requirements f o r an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study. I f u r t h e r agree that permiss ion fo r ex ten s i ve copying o f t h i s t he s i s f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s r ep re sen ta t i ve s . It i s understood that copying or p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l ga in s h a l l not be a l lowed without my w r i t t e n permi s s ion . Department o The U n i v e r s i t y of B r i t i s h Columbia 20 75 Wesbrook P l a c e Vancouver, Canada V6T 1W5 Date To face t i t l e page NEWLY FLEDGED RICHARDSON'S MERLIN i i ABSTRACT I t i s the grass l a n d ecosystem which supports n e s t i n g M e r l i n s on the Canadian p r a i r i e s . This study considered the e f f e c t s of some f a c t o r s i n t e r -f e r i n g w i t h processes i n operation i n the n a t u r a l f u n c t i o n i n g of the grass-land ecosystem that i n f l u e n c e s M e r l i n p o p u l a t i o n s . An important p a r t of t h i s study was the comparison of s e l e c t e d h a b i t a t features near Hanna, A l b e r t a , where a segment of the p r a i r i e M e r l i n p o p u l a t i o n continues to nest, and those near K i n d e r s l e y , Saskatchewan, where M e r l i n s once nested but no longer are present. A d d i t i o n a l l y , data on ne s t i n g ecology of Richardson's M e r l i n s were gathered along the South Saskatchewan R i v e r , and near Hanna, i n southern A l b e r t a , during the summers of 1 9 6 8 - 1 9 7 4 . The absence of n e s t i n g merlins near K i n d e r s l e y appears to be r e l a t e d to changing human land use patterns i n that area. Since 1 9 5 1 , 6 2 7 „ of the land near K i n d e r s l e y has been c u l t i v a t e d , w h i l e the comparable f i g u r e near Hanna, where M e r l i n s continue to n e s t , i s 2 6 7 o . A i r photo study of t e r -r i t o r i e s around M e r l i n nest s i t e s showed that 5 2 . 3 7 » of the area presumed to have been used by M e r l i n s f o r hunting i n the K i n d e r s l e y area came under c u l t -i v a t i o n since the 1 9 4 0 ' s , as compared to 25 .57<, f o r the Hanna t e r r i t o r i e s over the same p e r i o d . I n t e r p r e t a t i o n of 4 0 M e r l i n t e r r i t o r i e s i n use sin c e 1 9 7 1 i n the Hanna area, has revealed that M e r l i n s were hunting i n areas which averaged 787o grassland. Increase i n c u l t i v a t i o n i n both areas s i n c e the e a r l y 1 9 6 0 ' s has been about 7 7 » , l e a v i n g K i n d e r s l e y M e r l i n t e r r i t o r i e s w i t h l e s s than 4 2 7 0 grassland by 1 9 7 1 . Assuming that at l e a s t 5 0 7 , g r a s s l a n d i s req u i r e d w i t h i n a M e r l i n hunting t e r r i t o r y i n order to provide s u f f i c i e n t small b i r d prey, i t was concluded that the K i n d e r s l e y area has not been a ' i i i prime M e r l i n n e s t i n g area since the 1 9 4 0 ' s , and that increases i n c u l t i v a t i o n since the e a r l y 1 9 6 0 ' s has probably reduced grassland below the th r e s h o l d necessary to support n e s t i n g M e r l i n s . I t i s f e l t t hat the heavy use of d i e l d r i n i n the e a r l y 1 9 6 0 ' s probably had the e f f e c t of adm i n i s t e r i n g the "coupe de grace" to the K i n d e r s l e y M e r l i n s , p o s s i b l y through the r e d u c t i o n of grassland b i r d s . A n a l y s i s of prey remains at nests i n d i c a t e d that the d i e t of these p r a i r i e M>erlins was composed of 507=, Horned Larks, 377=, C h e s t n u t - c o l l a r e d Longspurs, 6 7 » sparrows, 47« b l a c k b i r d s and 37, others (passerine b i r d s , shore-b i r d s , rodents, e t c . ) . D e s t r u c t i o n of the h a b i t a t s of these prey species must be viewed as d e s t r u c t i o n of the h a b i t a t f o r n e s t i n g M e r l i n s . The now common p r a c t i c e of seeding open c a t t l e range to c r e s t e d wheatgrass and other a l i e n monocultures i s l i k e l y to lead to f u r t h e r r e g r e s s i o n i n the p r a i r i e ecosystem and to reduce or e x t i n g u i s h populations of passerines and, consequently, M e r l i n s . I n A l b e r t a , from 1 9 7 1 - 1 9 7 4 , an average of 7 0 . 9 7 . of occupied nest s i t e s ( s i t e s w i t h a p a i r of M e r l i n s present p r i o r to egg l a y i n g ) were a c t i v e , at l e a s t to the egg l a y i n g stage; 54.97<> of nests w i t h eggs hatched young, and 8 5 . 3 7 o of these s u c c e s s f u l nests produced f l e d g l i n g s . Average c l u t c h s i z e was 4 . 6 eggs, of which an average of 3 . 5 hatched per nest with eggs hatching, producing 3 . 2 fledged young per nest with young reaching f l e d g i n g age. A net p r o d u c t i v i t y of 0 . 6 9 f l e d g l i n g s per occupied nest s i t e was determined. Hatching success f o r a l l eggs was 5 7 . 8 7 o , and 8 4 . 4 7 , of the young hatched survived to f l e d g i n g . These f i g u r e s i n c l u d e r e s u l t s from 1 9 7 3 when a s i n g l e storm accounted f o r a f a i l u r e of 4 1 . 4 7 , of the nests i n the Hanna area. Egg h a t c h a b i l i t y appears to be r e l a t e d ( p < ^ . 0 2 ) to DDE and D i e l d r i n r e s i d u e s , i v probably r e s u l t i n g from l o c a l a p p l i c a t i o n of these p e s t i c i d e s f o r grasshopper c o n t r o l ; e g g s h e l l d e n s i t y was i n v e r s l e y r e l a t e d to DDE l e v e l s . I n A l b e r t a , p e s t i c i d e l e v e l s do not appear to have been s u f f i c i e n t to cause a p o p u l a t i o n d e c l i n e . R e s u l t s from banding of n e s t l i n g s and trapping and banding of adults showed that males u s u a l l y r e t u r n to the general area where they have nested i n previous years whereas females may move as f a r as 100 miles from an e a r l i e r n e s t i n g s i t e . Apparent records of two b i r d s , one of each sex, breeding at an age of one year, are i n t e r e s t i n g i n that most r a p t o r s do not breed u n t i l t h e i r second or t h i r d year. Longevity of these b i r d s i n the w i l d i s unknown but one i n d i v i d u a l was known to have been 5 years o l d . V ACKNOWLEDGEMENTS Financing of f i e l d work f o r t h i s p r o j e c t during the 1973 and 1974 f i e l d seasons was provided by the Canadian W i l d l i f e S ervice (C.W.S.). Thanks are e s p e c i a l l y due to Mr. Richard Fyfe of the Edmonton, A l b e r t a , o f f i c e of tha t S e r v i c e , r e s p o n s i b l e f o r the support, who provided encou-ragement f o r t h i s p r o j e c t since i t s i n c e p t i o n i n 1970. My p r o j e c t on Me r l i n s i s a p a r t of a much wider program e s t a b l i s h e d by the Toxic Chemical Secti o n (C.W.S.) to monitor r a p t o r p o p u l a t i o n changes and p e s t i c i d e residue l e v e l s i n ra p t o r s on the Canadian p r a i r i e s . Acknowledgement i s accorded C.W.S. f o r the use, from t h e i r f i l e s , of popul a t i o n data, p e s t i c i d e residue data, and eg g s h e l l data, c o l l e c t e d before I commenced work on t h i s p r o j e c t . S p e c i a l thanks goes to Mrs. Lynne Kemper f o r her work during the 1972 f i e l d season, f o r only through her complete and d e t a i l e d observations while working f o r C.W.S. i s a continuous account of the p r a i r i e M e r l i n p o p u l a t i o n a v a i l a b l e since 1971. To Miss U r s u l a Banasch, who provided the l i n k between f i e l d operations and the Edmonton o f f i c e , I am most g r a t e f u l . To the people named below and many others I g r a t e f u l l y acknowledge a s s i s t a n c e and advice: Mr. Harry Armbruster, W i l d l i f e Technician, Mr. Bob Wroe, Range S p e c i a l i s t , Mr. Randy Semeniuk, Mr. Tom Donald, Mr. Tom R u s s e l l , Mr. Pat H a r r i s , Mr. Jim Windsor, Mr. Mike Bradley, Mr. John l e Jeune, and Mr. Glen Fox. P a r t i c u l a r thanks f o r the sometimes menial task of c o l l e c t i n g p l a n t specimens, and feathers and other prey remains, and f o r the a i d i n tr a p p i n g and banding M e r l i n s go to K i p , Ken and K e l l y Fyfe who spent periods of each f i e l d season v i w i t h me, to my brother Kim who spent much of the 1971 f i e l d season w i t h me, and to my b r o t h e r - i n - l a w Darren E t h i e r who spent most of h i s 1973 and 1974 school vacations w i t h me. I hope they got as much out of those f i e l d seasons as I d i d from t h e i r h elp. During the 1973-74 and F a l l 1974 sessions spent at the U n i v e r s i t y of B r i t i s h Columbia a number of people were e s p e c i a l l y h e l p f u l . These i n c l u d e Mr. Steve Borden who undertook the computer a n a l y s i s of much f i e l d data, Mrs. M. North who provided source m a t e r i a l f o r v e g e t a t i o n and h i s t o r i c a l aspects of the study area, and the f o l l o w i n g members of my committee: Dr. V.C. Runeckles, Chairman; Dr. V.C. B r i n k , Supervisor; Dr. R.J. Copeman, A s s i s t a n t P r o f e s s o r , Department of P l a n t Science; Dr. Ian McTaggart Cowan, Dean, F a c u l t y of Graduate Studies, and P r o f e s s o r , Department of Zoology; Dr. R.C. Fitzsimmons, A s s i s t a n t P r o f e s s o r , Department of P o u l t r y Science; and Mr. Richard Fyfe, Research B i o l o g i s t , Canadian W i l d l i f e S e r v i c e . Very s i n c e r e thanks go to Mr. and Mrs. R. Fyfe of F o r t Saskatchewan, Mr. and Mrs. J . Armstrong of Hanna, and Mrs. M. Seminiuk of Edmonton a l l of whom provided meals and accommodation f o r sometimes very d i r t y and weary workers, and to Mr. and Mrs. R. Gore of S c o t f i e l d who very k i n d l y provided a place f o r a f i e l d t r a i l e r out of which much of the study was c a r r i e d out during 1973 and 1974 f i e l d seasons. Thanks too must be accorded Mr. B r i a n Davies, Warden George C. R e i f e l , Waterfowl Sanctuary, and Mr. Frank Beebe of the B.C. P r o v i n c i a l Museum, r e t i r e d . Through t h e i r c o n s i d e r a t i o n and f r i e n d s h i p an i n t e r e s t and f a s c i n a t i o n f o r r a p t o r s was k i n d l e d i n the e a r l y 1960's, and sustained. Receipt of a i r phots from the Canadian W i l d l i f e S e r v i c e L i b r a r y , Edmonton, A l b e r t a , from the A l b e r t a Department of Lands and For e s t s Photo v i i L i b r a r y , from the A l b e r t a Department of Geography A i r Photo L i b r a r y , and from the N a t i o n a l A i r Photos L i b r a r y i s acknowledged. To my s i s t e r - i n - l a w , Mrs. H o l l y Linden, who undertook the f i n a l t y ping of t h i s t h e s i s , I am very g r a t e f u l . Above a l l , thanks go to my w i f e Heather who, f o r more than two years,put up w i t h my long absences i n the f i e l d . v i i i TABLE OF CONTENTS PAGE ABSTRACT i t ACKNOWLEDGEMENTS , v TABLE OF CONTENTS v i i LIST OF TABLES x LIST OF PLATES x i i LIST OF FIGURES x i 1. INTRODUCTION x 2. GENERAL BIOLOGY OF THE RICHARDSON' S MERLIN 2 3. MAJOR PHYSICAL AND BIOLOGICAL FEATURES OF THE STUDY AREA 4 3.1 Physiography 4 3.2 Climate 6 3.3 Vegetation 7 3.4 Fauna 9 3.5 A g r i c u l t u r e 10 4. METHODS . 17 4.1 M e r l i n Surveys 1971-74 17 4.2 Land Use C l a s s i f i c a t i o n 18 4.3 The Impact of P e s t i c i d e s 20 5. OBSERVATIONS AND RESULTS 22 5.1 Pop u l a t i o n Dynamics of Richardson's M e r l i n 22 5.1.1 M e r l i n Populations i n Southern A l b e r t a , 1971-1974: Some Aspects 22 5.1.2 M e r l i n P r o d u c t i v i t y 25 i x 5. OBSERVATIONS AND RESULTS (Continued) PAGE 5.2 F a c t o r s A f f e c t i n g M e r l i n Populations 25 5.2.1 Weather 25 5.2.2 P r e d a t i o n 30 5.2.3 Human Disturbance 31 5.2.4 P e s t i c i d e s 33 5.2.5 Disease 38 5.2.6 Land Use Change 38 6. DISCUSSION 46 7. SUMMARY AND CONCLUSIONS 58 8. LITERATURE CITED 6 0 9. APPENDICES 6 2 I P e s t i c i d e Residue Levels i n Eggs of Richardson's M e r l i n 1971-1973. 62 I I A n a l y s i s of Fact o r s of Nest S i t e s Used by Richardson's M e r l i n A l b e r t a . 63 I I I Prey Species U t i l i z e d by Richardson's M e r l i n 65 IV Common Pla n t s of the Study Area ' 72 V M e r l i n P opulation and Nest Data 78 VI M e r l i n P e s t i c i d e Data. 82 VI I Chemical Names of I n s e c t i c i d e s 83 X TABLE LIST OF TABLES PAGE 1 Change i n the Human Popu l a t i o n of the Census D i v i s i o n s of the Study Area. 15 2 Occupancy and Use of Nest S i t e s by Richardson's M e r l i n , 1971-1974. 23 3 P r o d u c t i v i t y of Richardson's M e r l i n , 1971-1974. 26 4 "Reproductive Success of Pigeon Hawks" 27 5 A Comparison of P e s t i c i d e Residue Levels and R a t c l i f f e I ndices f o r Richardson's M e r l i n Eggs. 34 6 A Comparison of P e s t i c i d e Residue Levels i n M e r l i n Eggs C o l l e c t e d During Incubation and Eggs C o l l e c t e d Dead A f t e r Incubation. 36 7 A Comparison of P e s t i c i d e Residue L e v e l s i n Eggs from " S u c c e s s f u l " Nests and Eggs from "Unsuccess-f u l " Nests. 37 8 The Areas Under C u l t i v a t i o n i n the Census D i v i s i o n s of the Study Area. 39 9 Percentage of Areas under C u l t i v a t i o n w i t h i n M e r l i n Hunting T e r r i t o r i e s Based on Studies of A i r Photos Taken between the 1940's and the 1970's. 40 10 The Acreage Under C u l t i v a t i o n (Permit and Lease) i n The S p e c i a l Areas of A l b e r t a . 41 11 A Comparison of C e r t a i n Environmental Factors Around Nest S i t e s near Hanna, A l b e r t a , and Kind-e r s l e y , Saskatchewan. 43 12 Sale of D i e l d r i n i n Some Rural M u n i c i p a l i t i e s of Saskatchewan, 1955-1965. 54 13 Species U t i l i z e d as Prey by M e r l i n s . 66 14. P a i r Density and Species Abundance of Grassland B i r d s i n Grassland and A g r i c u l t u r a l Regions. 67 15 "Average Number of B i r d s Recorded at Roadside Stops" 69 x i LIST OF FIGURES FIGURE PAGE 1. The Vegetation of the Canadian P r a i r i e s . 5 2. The P a l l i s e r T r i a n g l e . 12 3. Temperature and P r e c i p i t a t i o n f o r Hanna and Medicine Hat, A l b e r t a , June 1973. 29 4. Simple Regression G r a p h - - DDE as the Dependent V a r i a b l e and the R a t c l i f f e Index as the Independent V a r i a b l e . 35 5. Feeding Behaviour of Grassland B i r d s . 70 x i i LIST OF PLATES PAGE FRONTISPIECE: Newly Fledged Richardson's M e r l i n . P l a t e 1. M e r l i n Nest S i t e Along the South Saskat-chewan R i v e r . 3 P l a t e 2. M e r l i n Nest S i t e i n an Upland Aspen Grove. 3 P l a t e 3. M e r l i n Nest S i t e i n an Abandoned Wind-break. 45 P l a t e 4. M e r l i n Nest S i t e i n Springtime Showing Magpie Nests Used by M e r l i n s . 45 P l a t e 5. M e r l i n Nest S i t e i n an Abandoned Wind-break Showing E f f e c t of C a t t l e on Wind- 49 breaks. P l a t e 6. " H i g h l i n i n g " and Breakage by C a t t l e Using an Abandoned Windbreak as S h e l t e r . 49 1 1. INTRODUCTION Richardson's M e r l i n , a d i s t i n c t i v e r e s i d e n t of the gras s l a n d s , meadows and aspen parklands of the Great P l a i n s of North America, continues to e x i s t d e s p i t e the d e s t r u c t i o n of most of the o r i g i n a l p r a i r i e sod f o r a g r i c u l t u r e . This study of what has happened and i s happening to t h i s b e a u t i f u l r a p t o r , an organism at the apex of a p r a i r i e t r o p h i c pyramid, r e f l e c t s the extent and nature of man's m o d i f i c a t i o n of grass l a n d systems. I n Western Canada, where the M e r l i n i s r e s i d e n t from A p r i l u n t i l September, the a g r i c u l t u r a l system has almost overwhelmed the n a t u r a l system, but i t i s not too l a t e to i d e n t i f y the prime f a c t o r s i n f l u e n c i n g M e r l i n d i s t r i b u t i o n and abundance. The i d e n t i f i c a t i o n of these f a c t o r s , a purpose of the studies reported here, may be an a i d i n determining the ways i n which the p r a i r i e environment may accomodate man as w e l l asccomponents of the n a t u r a l p r a i r i e system, such as the M e r l i n . 2 2. GENERAL BIOLOGY OF THE RICHARDSON'S MERLIN The M e r l i n i s a member of the Genus Falco and i s one of s i x species n a t i v e to North America, v i z . the M e r l i n , F a l c o columbarius; American K e s t r e l , F a l c o s p a r v i u s ; Aplomado Falcon, Falco f e m o r a l i s , ; P r a i r i e Falcon, F a l c o  mexicanus; Peregrine Falcon, Falco peregrinus; and Gyrf a l c o n , Falco r u s t i c o l u s . But f o r the k e s t r e l , the M e r l i n i s the s m a l l e s t . Female M e r l i n s weigh i n the range of 185 g- to 255 g. The males, as i n most other r a p t o r s p e c i e s , are about one t h i r d smaller than the females and weigh about150g to 220 g (Amadon and Brown, 1968). The female M e r l i n i s thus about the s i z e of a small pigeon. Temple (1970) recognizes three subspecies of M e r l i n s i n North America, F a l c o  columbarius columbarius (formerly F.c_. columbarius and F.c_. b e n d i r e i ) summering i n the t a i g a zone, F.£. s u c k l e y i of the P a c i f i c northwest, and F.c_. r i c h a r d s o n i i n e s t i n g i n the p r a i r i e - p a r k l a n d , the b i r d of t h i s study. Richardson's M e r l i n s are c h a r a c t e r i s t i c a l l y b i r d s of open country; other M e r l i n s are found breeding i n h e a v i l y f o r e s t e d areas but i n such areas they hunt over n a t u r a l openings and r i v e r v a l l e y s . M e r l i n s are extremely ener-g e t i c , and capture most prey on the wing i n the f a s t , long-winged p u r s u i t s c h a r a c t e r i s t i c of f a l c o n s . T h e i r c h i e f prey are small b i r d s , but i n s e c t s , at times, form a s i g n i f i c a n t p a r t of t h e i r d i e t . As i n the case with other f a l c o n s , M e r l i n s do not b u i l d t h e i r own nest, but use nests b u i l t by other r a p t o r s , crows, or magpies, or simply nest on the ground i n an elevated p o s i t i o n on a c l i f f or h i l l t o p . Four or f i v e , or occasio-n a l l y six,eggs are l a i d , and i n c u b a t i o n , which begins some time a f t e r the f i r s t egg i s l a i d , l a s t s about a month. The young M e r l i n s spend about a month i n the nest before they begin to f l y . 3 P l a t e 1. M e r l i n Nest S i t e Along the South Saskatchewan R i v e r P l a t e 2. M e r l i n Nest S i t e i n an Upland Aspen Grove To face page 5 4 3. ' MAJOR PHYSICAL AND BIOLOGICAL FEATURES OF THE STUDY AREA 3.1 Physiography The Great P l a i n s of North America, c e n t r a l l y l o c a t e d on the c o n t i n e n t , extend from the G u l f of Mexico to the A r c t i c Ocean. From south c e n t r a l Canada southward, the p l a i n s are p r a i r i e . By d e f i n i t i o n , p r a i r i e s are s t r e t c h e s of medium to t a l l grasses i n temperate c l i m a t e s w i t h an annual p r e c i p i t a t i o n of between 20 and 40 inches per year; however the popular connotation of p r a i r i e s i s g rassland and i n c l u d e s the short-grassed p l a i n s or steppes, the t a l l grassed p l a i n s and the s p a r s e l y t r e e d g r a s s i a n d - f o r e s t t r a n s i t i o n (savannah). I n the Canadian p l a i n s provinces ( p r a i r i e provinces) t h i s " - p r a i r i e " extends roughly i n a t r i a n g l e bounded on the south by the I n t e r n a t i o n a l Boundary, on the west by the Rocky Mountains, and on the north and east by a l i n e drawn from north of Edmonton, A l b e r t a to the Red R i v e r V a l l e y of southern Manitoba (Figure 1). I t i ncludes the short grass and mixed grass p r a i r i e s of A l b e r t a and Sask-atchewan, the t a l l grass p r a i r i e of the Red R i v e r V a l l e y of Manitoba, and the aspen parkland. The landscape of the Canadian p r a i r i e s i s not simply the f l a t and open p l a i n p i c t u r e d by those u n f a m i l i a r w i t h the r e g i o n , but i s a broken and d i v e r s e composition of p l a i n s and h i l l s and i n c i s e d r i v e r v a l l e y s . A basement of Precambrian rock whose o r i g i n s date back at l e a s t two b i l l i o n years under-l i e much of the p r a i r i e from depths of h a l f m i le to over two m i l e s . Periods of inundation by sea waters, sedimentation, r e t r e a t of seas,and e r o s i o n by warm water and i c e are presented i n the h i s t o r y of the p r a i r i e bedrock and landscape. The impact of P l e i s t o c e n e i c e over the past m i l l i o n years i s recorded i n n e a r l y a l l p a r t s of t h i s study area -- i n i n t e r u p t e d drainage, t i l l sheets, l a c u s t r i n e s o i l s , s a n d h i l l s and other t e r r a i n f e a t u r e s . The F i g u r e 1. The Vegetation of the Canadian P r a i r i e s ( a f t e r Smith, 1972) 6 last retreat of ice began over 12,000 years ago; the varied s u r f i c i a l materials of the wake became the parent materials of the present prairie soils. Differential melting of the ice resulted in the formation of today's small pothole lakes and sloughs; meltwater cut channels, many of which today, being largely devoid of water, are referred to as coulees. A few upland areas, such as the Cypress H i l l s and the Porcupine H i l l s , escaped the work of advancing and retreating ice. The comparatively level macrotopography over which the present prairie vegetation and s o i l is established slopes from an elevation of about 1200 metres in the southwestern Alberta immediately east of the Rocky Mountains to below 220 metres in northern Manitoba, where i t merges with the Precambrian rocks of the Canadian Shield. Numberous upland areas occur on the plains rising 300 to 800 metres above the general surface of the plain. Ex-amples are the Cypress H i l l s of Alberta and Saskatchewan and Riding Mountain in Manitoba. Many proglacial lakes, now largely drained, l e f t sediments which are level and which are the parent materials of some of the best agricultural soils of the plains. Windsorting of soils, occurring as deglaciation pro-ceeded (and is s t i l l occurring in a minor way), has been responsible for ex-tensive loessial loams, blow-out lands, and sandhills. 3.2 Climate The northerly latitude and the mid-continent location of the prairie provinces accounts for wide fluctuations in temperature and the great variations in precipitation from year to year. The Rocky Mountains to the west impede the flow of mild, moist, Pacific air and place this study area in a rain shadow with limited total precipitation. The Rockies however do not impede but in a sense contribute to the flow of cold air from the north, particularly in winter, and to the flow of warm air from the south in summer. The climate of the study 7 area i s one of extremes both i n time and l o c a t i o n w i t h p r e c i p i t a t i o n that can range from 15.24 cm (6 in) i n a dry year to 70.12 cm (28 in) i n a wet year i n the same l o c a t i o n and temperatures which can range from a January low of -45.5° C (-50°F) to a 42.2°C (108°F) J u l y h i g h . Mean annual p r e c i p i t a t i o n (30 y r normals) over the general area i n which the study area i s loc a t e d i s 30 cm (11.79 i n) and i s 50 cm (19.65 i n ) f o r the wetter s e c t i o n s -- the A l b e r t a F o o t h i l l s , the aspen parkland, the Red Ri v e r V a l l e y , and eminences above the p l a i n such as the Cypress H i l l s . The r e g i o n of lowest p r e c i p i t a t i o n i s found i n the so u t h - c e n t r a l i n t e r i o r of the r e g i o n on the boundary of south-eastern A l b e r t a and south-western Saskatchewan. Mean temperatures f o r the p r a i r i e r e g ion range between -10°C (14°F) and -2°C (-1°F) i n January to between 18°C (64.4°F) and 20°C (65.3°F) i n J u l y . A concise account of f a c t o r s r e s p o n s i b l e f o r c l i m a t i c c o n d i t i o n s on the Canadian p r a i r i e s i s given by Laycock ( i n Smith, 1972). 3.3 Vegetation The o r i g i n a l grasslands of the p r a i r i e provinces included the t a l l grass p r a i r i e s of the Red R i v e r V a l l e y , the mixed grass p r a i r i e s covering the major p o r t i o n of the r e g i o n and extending i n t o the aspen parkland to the n o r t h , and the short grass p r a i r i e of southern A l b e r t a and Saskatchewan. Today l i t t l e of the t a l l grass p r a i r i e v e g e t a t i o n remains and much of the mixed grass p r a i r i e has disappeared as w e l l f o r t h i s v e g e t a t i o n has gone under the plow p r o v i d i n g some of the most productive c e r e a l lands of the world. The areas of mixed grass p r a i r i e and short grass p r a i r i e , not s u i t a b l e f o r arable a g r i c u l t u r e , support a la r g e c a t t l e i n d u s t r y . General d e s c r i p t i o n s of p r a i r i e v e g e t a t i o n are given by Laycock ( i n Smith, 1972) and by Webb, Johnston, and Soper ( i n Hardy, 1967); a d e t a i l e d d e s c r i p t i o n f o r A l b e r t a i s given by Moss (1955). 8 Probably the most complete d e s c r i p t i o n of the p r i s t i n e n a t u r a l v e g e t a t i o n of the Canadian p r a i r i e s i s that by Watts (1960). Today the lands of the P a l l i s t e r t r i a n g l e i n which the study i s undertaken are, f o r the most p a r t , rangelands i n the A l b e r t a p o r t i o n and lands c u l t i v a t e d f o r c e r e a l production i n the Saskatchewan p o r t i o n . The non-c u l t i v a t e d grasslands of both of these areas are a cover of species main-tained by low r a i n f a l l , i n keeping w i t h the average of 14.64 i n (37.19 cm) p r e c i p i t a t i o n per year (based on a 1921-1927 average, Hanna). These grass-lands f a l l l a r g e l y i n t o the mixed p r a i r i e c l a s s i f i c a t i o n of Johnson et a l (1970) and are dominated by species of the spear grass-wheatgrass ( S t i p a -Agropyron) a s s o c i a t i o n . On the A l b e r t a side of the study area some lands are e s t a b l i s h e d to the shortgrass (Bouteloua-Stipa) a s s o c i a t i o n but composition i s dependent on the nature of grazing pressure; species such as groundsage (Artemesia f r i g i d a) and low sedge (Car ex eleo.charis) are more common on the more h e a v i l y grazed lands. Extensive patches of w i l l o w ( S a l i x .§_£.), wolf w i l l o w (Eleagnus commutata), and snowberry (Symphoricarpos o c c i d e n t a l i s ) , as w e l l as other shrubs occur on these grasslands throughout the area. Trees of the grasslands occur i n n a t u r a l groves of poplar (Populus tremuloides) and w i l l o w ( S a l i x sp.) around slough margins, and as stands of p o p l a r , w i l l o w , and Manitoba maple (Acer negundo) which are the remnants of windbreaks planted about s e t t l e r s ' homesites at some e a r l i e r time. Many of the grassland areas of t h i s r e g ion were at one time (see 3.5) under c u l t i v a t i o n , abandoned during the depression of the 1920's and 1930's, and through a slow process of n a t u r a l succession reached t h e i r present s t a t e of " n a t u r a l " g rassland. Today much of the g r a s s l a n d i s again being ploughed under, and reseeded to domesticated a l i e n species such as c r e s t e d wheatgrass 9 (Agropyron cristatum) i n an e f f o r t to increase the p r o d u c t i v i t y of the grass-land f o r l i v e s t o c k husbandry. 3.4 Fauna H i s t o r i c a l l y , the North American bison (Bison bison) was the most important l a r g e mammal i n the p r a i r i e ecosystem and i t ranged over our study area. This magnificant animal s u f f e r e d a l o s s of h a b i t a t through ranching and farm homesteading and was slaughtered u n c o n t r o l l a b l y u n t i l by 1888 ex-ceedingly few were l e f t , a decimation unprecedented i n n a t u r a l h i s t o r y annals. Today, few bison l i v e outside E l k I s l a n d N a t i o n a l Park i n c e n t r a l A l b e r t a and Wood B u f f a l o N a t i o n a l Park i n northern A l b e r t a . These parks were set up, i n p a r t , as conservation areas f o r these animals. To some degree the niche i n the p r a i r i e ecosystem once occupied by the b i s o n , i s , i n our study area, occupied by domestic range c a t t l e . When the b u f f a l o disappeared from the p r a i r i e as a source of l i v e l i h o o d f o r n a t i v e people and s e t t l e r s a l i k e , the hunting pressure turned on the pronghorn antelope ( A n t i l o c a p r a  americana) and the mule deer (Odocoileus v i r g i n i a n u s ) ; these l i k e w i s e were con s i d e r a b l y reduced i n numbers by the turn of t h i s century. Today, thanks to conservation measures and enforced hunting r e g u l a t i o n s , pronghorn antelope and mule deer, as w e l l as the w h i t e t a i l e d deer, occur i n the general study area and are now common on many pa r t s of the p r a i r i e s . K i t foxes (Vulpes  velox) have disappeared from the p r a i r i e s , but red foxes (Vulpes f u l v a ) and coyotes (Canis l a t r a n s ) are common; wolves (Canis l u p u s ) , although no longer r e s i d e n t , are seen o c c a s i o n a l l y when they move south from the northern f o r e s t s i n w i n t e r . The p r a i r i e g r i z z l y (Ursus arctos h o r r i b i l i s ) i s today found only i n the Swan H i l l s n orth of the aspen parkland; mountain l i o n s ( F e l i s concolor) are o c c a s i o n a l l y seen on the p r a i r i e s adjacent to the F o o t h i l l s of the Rocky Mountains. S i m i l a r l y , moose (Alces alces) normally r e s i d e n t i n the mountains have been reported on the 10 grasslands of western A l b e r t a ; one such animal was seen by the author i n the study area i n the summer of 1971. Small f u r - b e a r i n g animals such as beaver (Castor canadensis), muskrat (Ondatra z i b e t h i c a ) , and mink (Mustela vison) are numerous i n many marsh areas of the p r a i r i e s and provide an important source of revenue f o r many trappers. Other small mammals in c l u d e j a c k r a b b i t s (Lepus sp.) badgers (Taxidea t a x u s ) , porcupines ( E r e t h i z o n dorsatum), gr o u n d s q u i r r e l s ( C i t e l l u s s p . ) , pocket gophers (Thomomys talpoide's), bats ( C h i r o p t e r a ) , and mice and moles ( M i c r o t i n a e ) . For complete l i s t s and n a t u r a l h i s t o r i e s of p r a i r i e animals the reader i s r e f e r r e d to such works as those of Soper (1964) f o r mammals, and S a l t and Wilk, (1958) f o r b i r d s . B i r d l i f e i s v a r i e d on the study area, and on the p r a i r i e s g e n e r a l l y , from s p r i n g through f a l l . The numbers of b i r d s are not many i n wi n t e r but increase greatly,each s p r i n g w i t h migrants. Of the many migrants of the p r a i r i e s the best known are the multitudes of waterfowl that cloud the s k i e s each s p r i n g and f a l l . P r a i r i e r a p t o r s i n c l u d e many species that breed there, as w e l l as many others which winter there, or pass "through" i n mi g r a t i o n . Among the more notable summer r e s i d e n t s are Golden Eagles ( A q u i l a c h r y s a e t o s ) , Feruginous Hawks (Buteo r e g a l i s ) , P r a i r i e Falcons, and Richardson's M e r l i n s . Gallinaceous b i r d s i n c l u d e S h a r p t a i l e d Grouse (Ped rioectes, p h a s i a n e l l u s ) of the p l a i n s , Ruffed Grouse (Bonasa umbellus) of r i v e r i n e t h i c k e t s , and the introduced Pheasant (Phasianus c o l c h i c u s ) common i n and near c u l t i v a t e d land. 3.5 A g r i c u l t u r e I t i s p o s s i b l e to review the impact of white men on the study area, and on the p r a i r i e s g e n e r a l l y , i n many contexts. Inasmuch as a g r i c u l t u r a l impact on M e r l i n h a b i t a t i s so apparent and f a r reaching, and since the 11 recorded h i s t o r y of the study area i s l a r g e l y a g r i c u l t u r a l , the account which f o l l o w s emphasizes a g r i c u l t u r a l development as summarized by Gray (1967). The study area near Hanna, i n southeastern A l b e r t a , and K i n d e r s l e y , i n southwestern Saskatchewan, l i e i n what has been c a l l e d the P a l l i s e r T r i a n g l e (Figure 2). I t was named a f t e r the f i r s t e x p l o r a t i o n s . o f t h i s area by Capt. John P a l l i s e r i n the p e r i o d 1867-1870. I n h i s surveys, P a l l i s e r concluded that these lands were a northern extension of the Great American Desert and would never support a v i a b l e a g r i c u l t u r e ; h i s f o r e c a s t , made without a surmise of 20th century technology, was i n p a r t wrong. P a l l i s e r ' s journeys were mainly on the periphery of the study area; h i s surveys made i n a s e r i e s of dry years, and acc o r d i n g l y h i s judgement of the q u a l i t y of the land to produce crops was based on a s u p e r f i c i a l acquaintance. While P a l l i s e r ' s T r a i n g l e d i d i n c l u d e m i l l i o n s of acres of land which should not have been broken by the plow, i t included m i l l i o n s of acres s u i t a b l e f o r dry-land c e r e a l production. P a l l i s e r ' s judgement of the a g r i c u l t u r a l p o t e n t i a l was on the whole c o r r e c t c o n s i d e r i n g the s t a t e of a g r i c u l t u r a l technology of the day, f o r c e r e a l v a r i e t i e s and dryland farming techniques, f o r a northern p r a i r i e a g r i c u l t u r e , had not evolved i n North America. I t was the i n t r o d u c t i o n of summer f a l l o w i n g by the Fede r a l Indian Head A g r i c u l t u r a l S t a t i o n , Sask-atchewan, i n 1885, however, that turned the P a l l i s e r T r i a n g l e i n t o a produc-t i v e breadbasket. I n the system of summer f a l l o w i n g , crops are grown every second or t h i r d year; between the crop years the land i s ploughed and c u l t i v a t e d to prevent weed growth and to produce a summer dust mulch, measures which conserve s o i l moisture. The moisture that accumulates during the f a l l o w years can be used during the ensuing crop year. Summer f a l l o w i n g made the growing of 12 F igure 2. The P a l l i s e r T r i a n g l e -- denoted by coarse v e r t i c a l l i n e s . ( a f t e r Dawson and Younge, 1940) 13 s a t i s f a c t o r y crops p o s s i b l e where without i t none would be p o s s i b l e . However, i n a succession of extremely dry years, i n the study area, even summer f a l l o w i n g can not conserve enough moisture to produce a crop, and the f i n e l y t i l l e d mulch becomes i n c r e a s i n g l y subject to wind e r o s i o n . I n the droughts of the 1920's and 1930's, i t was the bone-dry land i n the summer f a l l o w which blew away. P r i o r to the t u r n of the century there was a growing fear that;the P a l l i s e r T r i a n g l e area would be taken i n t o the U n i t e d States of America. As the rush of American homesteaders i n t o the American west c l o s e d o l d b u f f a l o pastures to the cattlemen, American ranchers were forced westward and n o r t h -ward. The wide open and u n s e t t l e d spaces of the P a l l i s e r T r i a n g l e , on both sides of the border, l o o k e d - i n v i t i n g to the d i s p l a c e d beef producers. As the American ranching i n d u s t r y d r i f t e d north of the border, an aggressive Canadian s e t t l e m e n t ^ p o l i c y developed by S i r C l i f f o r d S i f t o n , then the M i n i s t e r of the I n t e r i o r of the Canadian government, urged that lands of the P a l l i s e r T r i a n g l e be farmed by Canadians and non-Americans. Between 1896 and 1914 the Dominion Government gave away m i l l i o n s of acres of homestead lands and the r a i l w a y s and land companies s o l d m i l l i o n s more. Immediately upon the d e c l a r a t i o n of war on August 14, 1914, a great d r i v e was launched by the Dominion Government to b r i n g more land under c u l t i -v a t i o n ; over 4,800,000 a d d i t i o n a l p r a i r i e acres were brought i n t o c u l t i v a t i o n i n 1915. The pressure f o r more g r a i n production kept up f o r the next four years and 12,000,000 c u l t i v a t e d acres were added to the c u l t i v a t e d t o t a l on the p r a i r i e s ; of t h i s more than 5,500,000 acres were broken i n the P a l l i s e r T r i a n g l e . The season of 1915 was notable f o r the high per acre g r a i n y i e l d s and the fabulous crops d i d much to s t i m u l a t e settlement; y i e l d s .of 30 to 40 14 bushels of wheat to the acre were common i n the P a l l i s e r T r i a n g l e and the average y i e l d of 25 bushels was destined to be a record f o r 40 years. The b i g crop of 1915 s p e l l e d tragedy f o r the shortgrass p r a i r i e s of the P a l l i s e r T r i a n g l e though, because i t r e s u l t e d i n the breaking of thousands of acres of submarginal land. Crop f a i l u r e followed crop f a i l u r e on marginal lands i n 1917, 1918, 1919 and 1920. In southeastern A l b e r t a movement of s e t t l e r s out of the country bordering Saskatchewan became almost a stampede between 1921 and 1926. The 1926 census recorded more than 10,,000 abandoned farms, h a l f of them were north of Medicine Hat and i n ,the area of t h i s study. The A g r i c u l t u r a l Extension Department i n A l b e r t a reversed i t s previous p o l i c y of emphasis on g r a i n and l i t t l e emphasis on l i v e s t o c k to one of l i v e s t o c k f i r s t and g r a i n second and began an a c t i v e program of de-population i n 1926. F a m i l i e s d e s i r i n g to emigrate from the s t r i c k e n land were given f r e e passage, and t r a i n load a f t e r t r a i n load of d e s t i t u t e farmers were transported to the north and to the west. G r a d u a l l y a large part of eastern A l b e r t a was taken out of g r a i n growing and returned to grasslands by l a r g e l y "autogenic" processes. I n 1927 a S p e c i a l Areas Board was set up i n A l b e r t a to manage those lands i n the south-east where much land had been deserted and r e v e r t e d to the crown. The p a t t e r n s of settlement and land abandonment are r e f l e c t e d i n the changes i n human pop-u l a t i o n s , i n the F e d e r a l census, 1906-1971, f o r my study area (Table 1). In the K i n d e r s l e y , Saskatchewan, area the brown s o i l s are,of a l l the s o i l s of the P a l l i s e r T r i a n g l e , most s u i t e d to the growing of g r a i n ; i n ad-d i t i o n , de-population was not encouraged i n t h i s area by the Saskatchewan government. Consequently, many people remained on the land on the Saskatchewan side of the border and the s t r u g g l e to produce crops continued i n t o the d r e a d f u l TABLE 1. CHANGES IN THE HUMAN POPULATION OF THE CENSUS DIVISIONS OF THE STUDY AREA 1906 1911 1921 1931 1941 1951 1961 1971 Haima, A l t a . D i v i s i o n 479 16,984 30,678 25,261 15,920 13,182 15,020 12,991 K i n d e r s l e y , Sask. D i v i s i o n 1,111 12,480 35,483 42,632 36,346 30,721 32,994 30,947 (From: Canada Census) !6 years of the 1930's. The depression of the 1930's brought an economic d i s a s t e r of equal magnitude to the c o n t i n u i n g p h y s i c a l d i s a s t e r of drought. The calamatous c o l l a p s e of farm p r i c e s between 1930 and 1933 reduced farm purchasing power to near zero, and the p r i c e s f o r crops that.were produced b a r e l y p a i d f o r t h e i r t r a n s p o r t a t i o n ; t o market. On top of weather- and economic a d v e r s i t i e s , there were plagues of i n s e c t s (grasshopper, sawfly etc.) and b l i g h t s of p l a n t disease (wheat r u s t , c e r e a l smuts etc.) to face. U n t i l the grasshoppers descended i n clouds i n 1933 a good crop was i n the making i n p a r t s of the P a l l i s e r T r i a n g l e . A go,od crop was i n prospect too i n 1935 u n t i l the r u s t attacked... By the l a t e 1930's a glimmer of hope appeared w i t h the r e t u r n of the r a i n s and the b e t t e r i n g of the world economy; g r a d u a l l y the b a t t l e against d r i f t i n g t o p s o i l was won w i t h new farming techniques and extensive reseeding; the f i g h t against i n s e c t s gained the upper hand thanks to new and i n n o v a t i v e c o n t r o l programs, and- the problem of wheat r u s t was l a r g e l y overcome by the development of r u s t r e s i s t a n t s t r a i n s of wheat. By the 1940's the " b a t t l e had been won" and the p a r t of Saskatchewan i n our study area was r e t u r n i n g to an a g r i c u l t u r a l economy based,again l a r g e l y on the growing of wheat and other ce r e a l s . 17 4. METHODS 4.1 M e r l i n Surveys, 1971-74 F o l l o w i n g the 1968 observation of a p a i r of M e r l i n s n e s t i n g i n the ranching country of southern A l b e r t a near Hanna, a reconnaissance survey f o r n e s t i n g M e r l i n s wa« undertaken i n 1969 by the Canadian W i l d l i f e S e r v i c e ; i n t h i s survey 14 p a i r s were found n e s t i n g . In 1970 a more i n t e n s i v e survey lo c a t e d 23 p a i r s n e s t i n g . The area along the South Saskatchewan R i v e r system i n A l b e r t a a l s o was surveyed i n t h i s same season and 30 p a i r s were found. 'Following the l o c a t i o n of these two s i z e a b l e populations a study by Canadian W i l d l i f e Service of the "dynamics" of Richardson's M e r l i n was proposed; i n 1971 t h i s study was begun. During the author's absence i n A f r i c a i n 1972 the study was undertaken by Mrs. Lynne Kemper under c o n t r a c t to the Canadian W i l d l i f e S e r v i c e . The data presented i n t h i s r e p o r t i s based on M e r l i n surveys by v e h i c l e and by r i v e r boat undertaken i n 1971, 1972, 1973 and 1974. The data are derived mainly from two areas i n southern A l b e r t a v i z . a p o r t i o n of the ranching country about Hanna, A l b e r t a and the area along the South Saskatchewan R i v e r as i t flows through A l b e r t a . R i v e r survey i n c l u d e d a short s t r e t c h of the Bow R i v e r upstream from i t s confluence w i t h the Oldman R i v e r ; the con-fluence of the Bow and Oldman marks the beginning of the South Saskatchewan R i v e r . In each year these areas were surveyed thoroughly, a l l occupied s i t e s were mapped, and a l l a c t i v e nests were "climbed" to determine reproductive success and to gather other i n f o r m a t i o n . Banding of young M e r l i n s and trapping and banding of a d u l t M e r l i n s at nest s i t e s provided some l i m i t e d i n f o r m a t i o n on f i d e l i t y of p a i r s and r e -use of n e s t i n g s i t e s by M e r l i n s from year to year. 18 Although M e r l i n s now do not i n h a b i t the K i n d e r s l e y area of Sask-atchewan, a l l s i t e s i n the K i n d e r s l e y area of Saskatchewan, where M e r l i n s had been n e s t i n g i n the 1960's and 1970's,were checked i n 1972 and 1974. No n e s t i n g M e r l i n s were found. A p a i r was found i n 1972 near the A l b e r t a border which began two cl u t c h e s but which hatched no eggs, and another p a i r was found n e s t i n g i n Saskatchewan near the South Saskatchewan R i v e r . Fox (1971) reported that f o r the K i n d e r s l e y area the l a s t reported n e s t i n g was i n 1962 and M e r l i n s have not been found since then by R. Fyfe (personal communication) i n p e r i o d i c checks. Throughout the M e r l i n surveys records were kept of features of each n e s t i n g s i t e which were deemed to a f f e c t n e s t i n g M e r l i n . The type of rec o r d i n g i s given i n Appendix 9.2. From an a n a l y s i s of these data, at l e a s t some of the h a b i t a t requirements of M e r l i n s may be perceived and comparisons of s i t e s i n use w i t h s i t e s no longer i n use can be made. During these surveys prey remains were c o l l e c t e d from M e r l i n nests and associated p l u c k i n g perches f o r l a t e r i d e n t i f i c a t i o n . A c o l l e c t i o n of b i r d s k i n s of prey species made on the study area aided i n the i d e n t i f i c a t i o n of prey remains. 4.2 Land Use C l a s s i f i c a t i o n Land use patter n s since 1945-1950 were determined through i n t e r -p r e t a t i o n of a e r i a l photographs. Three s e r i e s of photographs were used both f o r A l b e r t a and f o r Saskatchewan. For A l b e r t a , photographs used were a f i r s t set made between 1949 and 1952, a second set made between 1962 and 1965, and a t h i r d set made i n 1971. The corresponding s e r i e s of photographs from Sask-atchewan were a f i r s t set made i n 1946, a second set made between 1956 and 1961, and a t h i r d set made i n 1971. F o r t y M e r l i n s i t e s occupied near Hanna, i. A l b e r t a during t h i s study p e r i o d and 15 s i t e s of M e r l i n s known from the area of 19 K i n d e r s l e y , Saskatchewan were stud i e d i n t h e i r r e s p e c t i v e periods of photo a v a i l a b i l i t y . T h i r t y - n i n e s i t e s along the South Saskatchewan R i v e r were also s t u d i e d i n a set of photographs from 1971. The area w i t h i n a one mile r a d i u s around each M e r l i n s i t e was mapped i n t o areas of rangeland and c u l t i v a t e d land. A f u r t h e r c l a s s i f i c a t i o n of rangeland was not p o s s i b l e with the sc a l e of photography a v a i l a b l e . .From photo i n t e r p r e t a t i o n a p i c t u r e of land use change, over a p e r i o d of about 25 years, was obtained and a comparison between land use changes at M e r l i n s i t e s i n use i n A l b e r t a and land use changes at M e r l i n s i t e s no longer i n use i n Saskatchewan was p o s s i b l e . An overview of land use changes i n both A l b e r t a and Saskatchewan can be derived from s t a t i s t i c s i n Canada Year Books, based on the ten year census. S t a t i s t i c s on land use f o r Census D i v i s i o n s i n which study areas i n each province f e l l were organized f o r comparative s c r u t i n y . Present land use patterns were determined by the simple mapping about M e r l i n s i t e s i n use i n the Hanna area. From a p r i o r i M e r l i n behaviour observations i t was b e l i e v e d that M e r l i n were hunting up to one mile away from t h e i r n e s t i n g s i t e s , and f o r t h i s reason an area of one mile r a d i u s about nest s i t e s was mapped. A r e p r e s e n t a t i v e c o l l e c t i o n of common and dominant p l a n t species was made at M e r l i n s i t e s under c o n s i d e r a t i o n . When a d e t a i l e d c l a s s i f i c a t i o n of rangeland i n the Hanna area of A l b e r t a was contemplated i t was q u i c k l y evident that the rangeland i n the Hanna area of A l b e r t a f e l l i n t o the mixed p r a i r i e category (Johnson et a l . 1970), w i t h d i f f e r e n c e s observed i n the rangeland due to d i f f e r e n t i a l g r azing by l i v e s t o c k . The South Saskatchewan R i v e r area study was d i f f e r e n t from that of 20 Hanna i n that very l i t t l e land was under c u l t i v a t i o n . This area too f a l l s i n t o the mixed p r a i r i e and shortgrass p r a i r i e c l a s s i f i c a t i o n of rangeland and i s very l a r g e l y subjected to l i v e s t o c k g r a z i n g ; however, the species composition of t h i s area d i f f e r e d somewhat from that of Hanna as a rep-r e s e n t a t i v e c o l l e c t i o n of common and dominant p l a n t species made along the South Saskatchewan R i v e r w i l l show. The S u f f i e l d M i l i t a r y Reserve along the South Saskatchewan R i v e r , except f o r some concessions, has not e x p e r i -enced l i v e s t o c k g r a z i n g f o r many years. 4.3 The Impact of P e s t i c i d e s Work c a r r i e d out s i n c e 1969 f o r the Canadian W i l d l i f e S e r v i c e has taken a s p e c i a l i n t e r e s t i n the e f f e c t s of p e s t i c i d e s on M e r l i n s . As afore-mentioned, M e r l i n s , l i k e other r a p t o r s are at the "top" of the t r o p h i c pyramid and tend t h e r e f o r e t o consume the accumulation of c e r t a i n c h l o r i n a t e d hydrocarbons, which degrade very l i t t l e i n the t r o p h i c chain. A sample of i eggs was c o l l e c t e d each year and a n a l y s i s of residue of DDE, d i e l d r i n , heptachlor epoxide, and mercury i n them was made. DDE i s a breakdown metabolite of DDT which occurs very, s h o r t l y a f t e r i t s entry i n t o b i o l o g i c a l t i s s u e . An e g g s h e l l index, termed the R a t c l i f f e Index ( R a t c l i f f e , 1967) was determined f o r each egg. The outer dimensions of eggs c o l l e c t e d were measure and then the contents were sent to L.M. Reynolds of the Ontario Research Foundation to be analyzed f o r p e s t i c i d e residues by gas chromatography. This a n a l y s i s was done f o r the Canadian W i l d l i f e S e r v i c e as a p a r t of the C.W.S. p e s t i c i d e residue monitoring program. For a complete d e s c r i p t i o n of a n a l y s i s procedure r e f e r to F y f e , e_t _ a l . , 1969. Eggshells from samples c o l l e c t e d had membranes removed, were washed and were then allowed to dry at room temperature. Once dry, the s h e l l s were weighed to 0.01 g. From e g g s h e l l measurements and weights, the R a t c l i f f e Index was determined f o r a comparative measure of e g g s h e l l d e n s i t y . This index, defined by the formula R l = weight (mg) length X width (mm) has been used widely i n recent comparative studies of egg s h e l l c h a r a c t e r i s t i c s i n r a p t o r s , (Cooke (1973), review). P e s t i c i d e and residue data were obtained f o r the years 1968-1973 (unpublished C.W.S. residue data). Using residue l e v e l s and egg s h e l l i n d i c e s , s t a t i s t i c a l analyses were run to t e s t the c o r r e l a t i o n between i n c r e a s i n g p e s t i c i d e residue l e v e l s and decreasing e g g s h e l l d e n s i t y or th i c k n e s s . In order to t e s t the a s s o c i a t i o n between p e s t i c i d e residue l e v e l s and egg hatch-a b i l i t y , residue l e v e l s i n eggs c o l l e c t e d from nests i n which young had sub-sequently hatched were compared with l e v e l s i n eggs from nests which f a i l e d to hatch young. A l s o , eggs c o l l e c t e d at random were compared w i t h "dead e§gs" (eggs which f a i l e d to hatch). As a pa r t of the C.W.S. residue monitoring program, average residue l e v e l s f o r each year i n which eggs were c o l l e c t e d gave: an i n d i c a t i o n of the changes i n l e v e l o f p e s t i c i d e contamination i n the p r a i r i e environment (Appendix 1). The e f f e c t s of the use of h e r b i c i d e s was not considered i n t h i s study H e r b i c i d e s have not been widely used i n the p r a i r i e environment, although as more areas of grassland are put i n t o c e r e a l production the use of h e r b i c i d e s w i l l increase. Use of 2-4-D and r e l a t e d chemicals may have an e f f e c t of reducing seed-bearing "weed" species that support some grassland b i r d s . 2 2 5 . OBSERVATIONS AND RESULTS 5 . 1 P o p u l a t i o n Dynamics of Richardson's M e r l i n 5 . 1 . 1 M e r l i n populations i n Southern A l b e r t a , 1 9 7 1 - 1 9 7 4 ; Some aspects. In the surveys undertaken i n the 1 9 7 1 , 1 9 7 2 , 1 9 7 3 and 1 9 7 4 seasons (see 4 . 1 ) i n f o r m a t i o n was obtained on the use of n e s t i n g s i t e s . Occupancy of a M e r l i n s i t e was determined by the presence of adults and t h e i r v o c a l and aggressive behaviour. I n many cases the occupancy of a s i t e was evident p r i o r to egg l a y i n g but the b a s i c t e s t f o r s i t e use was the l a y i n g o f eggs. In Table 2 , a r e s u l t of four years observations, i t i s recorded that 7 0 . 9 7 . of s i t e s occupied were used f o r egg l a y i n g and 5 4 . 9 7 o s i t e s used saw young reared to fl e d g i n g age. A l l immature M e r l i n s 'studied since work began i n 1 9 6 8 were banded i f and where p o s s i b l e ; by 1 9 7 4 over 5 0 0 had been banded. I n a d d i t i o n , from 1 9 7 0 -1 9 7 4 1 5 9 adult M e r l i n s , a l l that could be trapped, were banded; of these, 2 5 adults c a r r y i n g bands have been retrapped at a c t i v e M e r l i n s i t e s and from these recaptures some comments can be made. From the n e s t i n g and banding observations i t seems h i g h l y probable that p a i r s p a r t each year and only by chance mate again i n ensuing years. I t i s also h i g h l y probable that males r e t u r n to the same s i t e i n successive years while females do not. Of 1 2 adult male trap-recaptures, 9 were on the same s i t e , 2 were l e s s than 2% miles away from t h e i r e a r l i e r occupancies and one was 8 miles away. The one. capture of a male, banded as a n e s t l i n g , was als o at a l o c a t i o n 8 miles away from where i t was banded. By comparison, of 1 0 adult female trap-rrecaptures, o n l y 2 were captured on the same s i t e , another 3 were captured over 1 0 m i l e s away, 3 were captured between 1 0 and 2 0 miles away, and 2 were captured more than 7 5 miles away from the handing s i t e . 23 TABLE 2. OCCUPANCY AND USE OF NEST SITES BY RICHARDSON * S MERLIN 1971-1974 Year Area # Nesting Success- Fledging Success S i t e s i A c t i v e A c t i v e S i t e s % of % of • - O c c u p i e d d S i t e s % Hatching % Fledging Successful Occupied S i t e s S i t e s 1971 Hanna 33 69.7(23) 78.3(18) 73.9(17) 94.4(17) 54.5(18) 1972 Hanna 38 78.9(30) 53.3(16) 50.0(15) 93.8(15) 42.1(16) 1973 Hanna 31 60.6(25) 48.0(12) 44.0(11) 91.6(11) 41.9(13) 1974 Hanna 34 79.4(27) 81.4(22) 77.7(22) 94.4(21) 61.7(21) Mean Hanna 77.2 65.3 61.4 93.3 50.1 1971 SSR* 34 67.6(23) 69.5(16) 47.8(11) 68.8(11) 32.3(11) 1972 SSR* 37 56.6(21) 47.6(10) 42.9( 9) 80.8( 8) 24.3( 9) 1973 SSR* 35 62.9(22) 59.1(13) 45.6(10) 76.9(10) 28.6(10) 1974 SSR* 49 71.4(35) 68.5(24) 57.1(20) 83.3(20) 40.8(20) Mean SSR* 64.6 61.2 48.3 . 77 A 31.5 Mean Hanna & SSR* 70.9 63.3 54.9 85.3 40.8 * South Saskatchewan R i v e r a S i t e s w i t h p a i r s present p r i o r to egg l a y i n g b S i t e s w i t h p a i r s producing eggs c S i t e s w i t h p a i r s hatching out young. 24 Two female M e r l i n s banded as n e s t l i n g s were retrapped as n e s t i n g a d u l t s , one 14 miles away and the other 96 miles away from the banding s i t e s . There were a t o t a l of 22 merlins trapped as adults that were r e -trapped at a l a t e r date, of these, 16 were recaptured the year a f t e r banding, 4 were recaptured the second year a f t e r banding, and 5 were recaptured the t h i r d year a f t e r banding. Of the three b i r d s banded as n e s t l i n g s and captured as breeding a d u l t s , one was captured the year f o l l o w i n g banding, one was captured as a breeding female at two years of age; and the t h i r d , a male, was captured as a breeding b i r d at three years of age. The n e s t i n g data f o r the Hanna area and the South Saskatchewan R i v e r area are given separately i n Table 2 since the n e s t i n g s i t e s grouped f a i r l y w e l l i n t o two. However, movement of b i r d s from area to area does e x i s t as i n d i c a t e d by the recapture of two breeding females. One b i r d was trapped as a breeding b i r d along the South Saskatchewan R i v e r and was retrapped one year l a t e r near Hanna. The second b i r d was banded as a n e s t l i n g on the South Saskatchewan R i v e r and retrapped the f o l l o w i n g year as a breeding b i r d near Hanna. This l a s t b i r d i s notable i n that i t was e v i d e n t l y breeding at one year of age while most r a p t o r s , i t i s b e l i e v e d , come of breeding age i n t h e i r second or t h i r d years. I t may be of i n t e r e s t to mention that one male trapped as a breeding b i r d was a l s o e v i d e n t l y breeding at one year of age, as judged by i t s plumage. The words " e v i d e n t l y breeding" are used since there has been some suggesting that.these b i r d s may only be replacements f o r the b i r d o r i g i n a l l y n e s t i n g at that s i t e a f t e r some misfortune has. 25 b e f a l l e n a member of a p a i r . This view however has not be s u b s t a n t i a t e d through any observation. 5.1.2 M e r l i n P r o d u c t i v i t y Table 3 contains " r e p r o d u c t i v e " data by year and by re g i o n . In compiling t h i s t a b l e , two assumptions were made: 1) nests i n which 5 eggs hatched or 5 young fledged had clu t c h e s of 5 eggs, except i n cases where 6 eggs were a c t u a l l y seen, and 2) b i r d s reaching banding age were considered to have reached f l e d g i n g age, unless i t was noted otherwise. P r o d u c t i v i t y was 4.58 eggs per nest w i t h eggs and 3.23 young fledged per nest w i t h young f l e d g i n g (averages over the four y e a r s ) , Fox (1971) during the pe r i o d of the K i n d e r s l e y m e r l i n d e c l i n e gives a comparable f i g u r e of 2.7 young per nest (Table 4) f o r p r a i r i e n e s t i n g m e r l i n s . Net p r o d u c t i v i t y i s the number I. of young produced per t o t a l nest s i t e s occupied, whether i n f a c t each nest s i t e produces young or not. This would be a s e n s i t i v e i n d i c a t o r of the s t a t e of h e a l t h of the p o p u l a t i o n , but has not been used because most st u d i e s of rap t o r s have not followed through on the n e s t i n g c y c l e from i t s beginning, and only gives f i g u r e s f o r the number of s i t e s s u c c e s s f u l in^producing eggs or young. I n t h i s study, a net production of 0.69 f l e d g i n g young per nest s i t e occupied was determined. 5.2 Factors A f f e c t i n g M e r l i n Populations 5.2.1 Weather The lowest p r o d u c t i v i t y i n the four years of t h i s study occurred (Table 3) i n 1973 i n the Hanna area. This can be a t t r i b u t e d to a storm o c c u r r i n g between June 13 and June 15. During the two weeks p r i o r to t h i s storm a l l nests i n the Hanna area had been checked f o r hatching, as many TABLE 3 PRODUCTIVITY OF RICHARDSON'S MERLIN, 1971-1974 Year Region 1 Eggs per Nest 2 Hatch per Nest 3 4 F l e d g l i n g s Eggs per f o r Nest A n a l y s i s Hatching Success 7o Fledging SuccessT. 7 Net P r o d u c t i v i t y 1971 Hanna SSR* 4.85(19) 5.1K 9) 3.77(17) 3.89( 9) 3.58(17) 3.57( 7) 15 7 67.7 72.5 95.1 81.0 0.89 0.77 1972 1973 Hanna SSR* Hanna SSR* 4.24(29) 4.81(16) 4.00(18) 4.30(10) 2.88(17) 2.68( 6) 3.08(12) 4.68( 6) 2.94(16) 3.22( 9) 2.73(11) 3.56( 9) 26 13 9 4 26.8 73.7 73.0 86.4 0.61 0.90 as 1974 Hanna SSR* Mean 1971-1974 4.76(25) 4.57(30) 3.52(21) 3.64(19) 3.30(20) 2.90(19) 10 14 4.58(156) 3.51(107) 3.23(108) 98 1. of nests w i t h eggs 2. of nests w i t h hatching 3. of nests w i t h f l e d g i n g 4. removed p r i o r to hatching, i . e . apparently v i a b l e 5. of eggs l a i d 6. of young hatched 7. # f l e d g l i n g s / n e s t s i t e occupied * South Saskatchewan R i v e r ** No comparable data. 62.2 44.1 57.8 89.2 81.4 84.4 0.80 0.63 0.69 27 TABLE 4. REPRODUCTIVE SUCCESS OF PIGEON HAWKS Eggs Per Nest Young 3 Hatching Hatching 1 1 Per Success F a i l u r e Nest % % Great P l a i n s pre-1950 Forested P r a i r i e 1950-1969 1950-1969 4.7(10) c 4.1(9) 4.5(10) 4.3(3) 92 4.0(16) 98 2.7(17) 49 0(3) 13(16) 41(17) a. young of any age; thus the minimum # eggs hatched b. percent of nests wit h advanced young which contained one or more unhatched eggs c. sample s i z e (adopted from Fox, 1971) 28 nests were j u s t at the hatching stage. The storm l a s t e d f o r two days, during the p e r i o d the winds gusted to more than 50 mph, over 4 inches of r a i n f e l l i n the study r e g i o n , w i t h some nearby areas r e p o r t i n g up to 8 inches, and the d a i l y maximum temperature f e l l to 52° (Figure 3), the lowest of the month. I F o l l o w i n g t h i s storm, over a 5 day p e r i o d , a l l ' n e s t s were again checked, w i t h the f o l l o w i n g f i n d i n g s : A c t i v e nests p r i o r to storm 26 A c t i v e nests a f t e r storm 15 I. Two ne s t s , s t i l l a c t i v e , were found with both l i v e young and dead young; one of these nests had one dead and one had two dead young. Of the eleven nests no longer a c t i v e , one was found with two dead i. young, s i x were found w i t h c o l d and dead eggs, two were found w i t h e g g s h e l l remnants ( i n d i c a t i n g that something, probably crows, had already found the deserted n e s t ) , and two nests were empty. s Other species l i v i n g at the same n e s t i n g s i t e s as the M e r l i n s a l s o s u f f e r e d losses from the storm. F i v e magpie n e s t s , two w i t h three dead young and three w i t h one dead young were found: two crow n e s t s , one w i t h f i v e c o l d i dead eggs and one with 4 dead young were found and two feruginous hawk n e s t s , both with one dead young were found. Beneath one feruginous hawk nest a dead adult feruginous hawk was found, apparently have succumbed during the storm. One M e r l i n nest had been blown f i f t e e n f e e t out of the nest t r e e and the M e r l i n eggshells were found twenty f e e t beyond. A r e p o r t r e f e r r r i n g to the a f f e c t s of adverse weather on r a p t o r s i s that by Rauchenbauch (1969) i n which he discusses r a p t o r populations i n West 29 Figure 3. Temperature and P r e c i p i t a t i o n f o r Hanna and Medicine Hat, A l b e r t a , June, 1973. (Data from Environment Canada Atmospheric Envi r o n -ment Service) To face page 29 TEMPERATURE - LINE GRAPH MEDICINE HAT HANNA JUNE 30 Germany. He w r i t e s ( t r a n s l a t i o n ) : "Of a t o t a l of 43 checked buzzard c l u t c h e s , 26 were deserted, and i n the r e s t o f the " s u c c e s s f u l " ones - - 9 young d i e d i n the nest. Only 18 young were fledged. That gives the average of 0.4 v. . fledged young/clutch s t a r t e d . Most of the cl u t c h e s f a i l e d during (due to) a f i v e day r a i n f a l l , even though the four to f i v e week o l d young were already almost a l l feathered". I t must be noted t h a t , d e s p i t e having experienced much the same co n d i t i o n s during the storm under c o n s i d e r a t i o n , the M e r l i n s along the South Saskatchewan R i v e r seem not to have s u f f e r e d from the bad weather c o n d i t i o n s . Indeed, the hatching success and f l e d g i n g success of M e r l i n s i n 1973 along the South Saskatchewan R i v e r were the hi g h e s t of the three years f o r which data were a v a i l a b l e . Three reasons are advanced to e x p l a i n why t h i s might have been so: f i r s t , M e r l i n s n e s t i n g along the South Saskatchewan R i v e r c o n s i s t e n t l y nested up to a week e a r l i e r than those around Hanna, judging from the development of young M e r l i n s : eggs ther e f o r e i n nests along the r i v e r would have hatched p r i o r to the storm; secondly, most of the trees i n which M e r l i n s nested along the r i v e r were along the shore i n the r i v e r bottom and th e r e f o r e out of the main b l a s t of high winds; and t h i r d l y , M e r l i n s n e s t i n g along the South Saskatchewan R i v e r were n e s t i n g i n b i g trees w i t h much l e a f cover, and i n dense stands which would serve>,as • p r o t e c t i o n against the i elements; nest trees around Hanna by comparison were much sm a l l e r , o f t e n sparsely l e a f e d , and o f f e r e d l i t t l e p r o t e c t i o n against the elements. 5.2.2 Predation P r e d a t i o n of nests by crows i s f e l t to be an important f a c t o r a f f e c t i n g n e s t i n g success i n M e r l i n p o p u l a t i o n s . Throughout the s p r i n g of 31 1971 53 % hours were spent i n a b l i n d observing a p a i r of M e r l i n s ; instances of the M e r l i n s f i g h t i n g w i t h crows which came too near t h e i r nest were r e -corded 56 times. These M e r l i n s were attempting to nest i n an open feruginous hawk's nest, an attempt which f a i l e d some time during the p e r i o d of egg l a y i n g . I t i s suggested that the crows f i n a l l y succeeded i n breaking through the M e r l i n ' s defences during the egg l a y i n g p e r i o d and had destroyed the c l u t c h . At another nest a c l u t c h l a i d by May 1 had disappeared by May 19 and a second c l u t c h was begun i n a nearby nest by May 27. Numerous f i g h t s w i t h crows were observed with t h i s M e r l i n p a i r . By J u l y 4, four of the eggs had hatched and a f i f t h was pi p p i n g . On that day the nest was being observed from a d i s t a n c e , and a p a i r of crows and t h e i r o f f s p r i n g were seen approaching the M e r l i n s i t e w i t h obvious i n t e n t . One p a r t i c u l a r l y determined crow was about to r a i d the nest when i t was beset by the M e r l i n s who drew feathers from the crow i n the process of d r i v i n g i t away. Nearby most s i t e s i n which M e r l i n s were ne s t i n g crows were a l s o n e s t i n g . Never was there any i n d i c a t i o n of nest predation -by magpies or of = q u a r r e l i n g between these b i r d s and M e r l i n s . Great horned owls are probable predators of M e r l i n s . One nest which M e r l i n s were using i n the sp r i n g of 1971 had only a great horned owl regur-g i t a t e d p e l l e t i n i t the f i r s t week of J u l y . At another s i t e the in c u b a t i o n of eggs ceased about h a l f way and only the female was to be found. On c l o s e i n s p e c t i o n of the s i t e , feathers and a wing of the male M e r l i n was found. The marked antagonism of M e r l i n s f o r great horned owls was very evident i n any encounter between the two sp e c i e s ; i t i s t h i s antagonism which provided the means f o r drawing ad u l t M e r l i n s i n t o a trap f o r banding. 5.2.3 Human Disturbance The f i r s t form of human disturbance considered i n t h i s study was that 32 caused by personal v i s i t s to n e s t i n g s i t e s f o r the purpose of s c i e n t i f i c study. I t must be s t r e s s e d that i n a l l v i s i t s to nests i n t h i s study, care was taken to keep d u r a t i o n of v i s i t s to a minimum, and these were not made i f weather 1 L c o n d i t i o n s were adverse. For a l l nestings recorded over the p e r i o d of t h i s study no r e l a t i o n s h i p between the number of v i s i t s to a near s i t e and the nest-in g success of that p a i r could be e s t a b l i s h e d . I t i s f e l t that i f c o n s i d e r a t i o n i s given to the c r i t i c a l periods of egg l a y i n g and egg hatching and v i s i t s are of minimum, du r a t i o n during e a r l y development, v i s i t s to nests by researchers have no s i g n i f i c a n t e f f e c t on n e s t i n g success. A more d i r e c t e f f e c t on M e r l i n s that cannot be ignored i s that of shooting. At one nest s i t e four young people were caught i n the act of shooting a hen M e r l i n that had been s u c c e s s f u l i n r a i s i n g 5 young. The t r a n s f e r a l of these young to other occupied M e r l i n nests nearby was .'effected. Subsequent i n q u i r y i n d i c a t e d that t r a v e l l i n g from i s o l a t e d grove to i s o l a t e d grove to shoot crows, magpies, and other "vermin" i n c l u d i n g hawks, was a w i d e l y p r a c t i s e d d i v e r s i o n f o r many people on Sunday afternoons i n the Hanna area. Further checks of M e r l i n s i t e s on the same day the shooting r e f e r r e d to above occurred revealed three s i t e s w i t h dead crows that had been shot r e c e n t l y . At one of these s i t e s where a M e r l i n nes r^f had had young a few days e a r l i e r , a dult or young M e r l i n s were not to be found. With M e r l i n s so v o c a l and obvious at n e s t i n g s i t e s the e f f e c t s of a few such shooters on M e r l i n numbers i s very s e r i o u s . I t was only a few years ago that annual "vermin" shoots were h e l d i n the Hanna area; the p a r t i c u l a r t a r g e t s were ground s q u i r r e l s and hawks but doubtless other species were shot. Discussions w i t h members of the Rod and Gun Club, and s p o r t i n g goods s t o r e operators were h e l d and now 33 most show very favourable attitudes towards raptors; many local ranchers, particularly, voice their criticism of weekend shooting by people from local towns on their rangelands. It is by the towns' people mainly that shooting probably w i l l continue to take a t o l l of nesting Merlins. 5.2.4 Pesticides The correlation analysis of residue levels and Ratcliffe Indices favours a significant relationship between DDE residues and Ratcliffe Indices (r = .369); no relationships between Ratcliffe Indices and dieldrin, hepta-chlor epoxide, or mercury residues were indicated (Table 5). A cause and effect relationship is strongly indicated by the highly significant regres-sion of log. DDE against Ratcliffe Indices (p<\01) (Figure 4). (For justification of the use of log DDE, see Blus et a l . t 1972). Eggs collected from 1971 until 1974 for pesticide analysis, were put into two categories, (a) those collected at"random" some time during incu-bation, and (b) those collected after hatching which had failed to hatch (Table 6). A significant (negative) relationship between DDE levels' and nest success was indicated, but none with other pesticide residues. Eggs collected from nests where eggs were l e f t for incubation were also put into two categories, (a) those from nests which succeeded in hatching the remaining egg(s), and (b) those from nests which did not suc-ceed in hatching any egg (Table 7). A highly significant relationship between hatching failure and levels of DDE and dieldrin is indicated. 34 TABLE 5. A COMPARISON OF PESTICIDE RESIDUE LEVELS AND RATCLIFFE INDICES FOR RICHARDSON'S MERLIN EGGS. P e s t i c i d e Standard C o r r e l a t i o n Residue - 1" Average D e v i a t i o n C o e f f i c i e n t DDE 14.73 D i e l d r i n 0.67 Heptachlor Epoxide 0.61 Mercury 0.19 18.14 -0.3693 0.74 -0.06519 0.75 -0.01641 0.20 -0.01513 1 i n pa r t s per m i l l i o n (ppm) wet weight. 35 F i g u r e 4. Simple Regression Graph — DDE as the Dependent V a r i a b l e and R a t c l i f f e I n d i c e s as the Independent V a r i a b l e . ' DDE - parts per m i l l i o n wet weight ( l o g a r i t h m i c scale) t = -5.089 . P C .01 Data from unpublished f i l e s of Canadian W i l d l i f e S e r v i c e , Edmonton. To face page 35 NON-ZERO DATA Y = 1-E57 + -0-2E42 * LOG X N = UBB X Ld Ld Lu U • < or 0.7. DDE 36 TABLE 6. A COMPARISON OF PESTICIDE RESIDUE LEVELS IN MERLIN EGGS COLLECTED DURING INCUBATION AND EGGS COLLECTED DEAD AFTER INCUBATION Pesticide''" Residue Mean Resudue L e v e l i n Eggs C o l l e c t e d During Incubation (ppm) N=112 Dead a f t e r Incubation (ppm) N=40 2 P r o b a b i l i t y DDE 12.83 24.66 0.03 D i e l d r i n 0.59 0.70 0.41 Hept. Epox. 0.56 0.46 0.27 Hg. 0.19 0.20 0.93 1 residue data from unpublished f i l e s of Canadian W i l d l i f e S e r v i c e , Edmonton. 2 p r o b a b i l i t y : p r o b a b i l i t y that d i f f e r e n c e s between equal means would be as great as the observed through chance alone. 37 TABLE 7. A COMPARISON OF PESTICIDE RESIDUE LEVELS IN EGGS FROM "SUCCESSFUL" NESTS AND EGGS FROM "UNSUCCESSFUL" NESTS. P e s t i c i d e Mean P r o b a b i l i t y -Residue l e v e l i n Eggs from  Successful Unsuccessful Nests Nests^ (ppm) (ppm) N=78 N=53 DDE 9.97 26.81 0.001 D i e l d r i n 0.54 0.83 0.02 Hept. Epox 0.54 0.60 0.66 Hg. 0.19 0.19 0.93 1. Su c c e s s f u l Nest: a nest that hatched any eggs 2. Unsuccessful Nest: a nest which f a i l e d to hatch any eggs 3. P r o b a b i l i t y : p r o b a b i l i t y that d i f f e r e n c e s between equal means would be as great as the observed through chance alone * Residue data from unpublished f i l e s of Canadian W i l d l i f e S e r v i c e , Edmonton f 38 5 . 2 . 5 Disease I n a nest i n a t r e e near the South Saskatchewan R i v e r two h e a l t h y young M e r l i n s about three weeks o l d were found along w i t h a t h i r d i n a very weak c o n d i t i o n . The s i c k b i r d was taken f o r care and a canker s i m i l a r to that caused by the protozoan p a r a s i t e Trichomonas g a l l i n a e i n other r a p t o r s was found adhering to the tongue and mouth l i n i n g ; t h i s c o n d i t i o n i s known as "frounce" and i s q u i t e w e l l known to f a l c o n e r s . I t i s f a i r l y common i n pigeons and domestic f o w l . Recently, i t s treatment w i t h the drug "Emtryl" (Dimetridazol) has proved very s u c c e s s f u l i n r a p t o r s , and so the M e r l i n \ V was t r e a t e d w i t h t h i s drug. W i t h i n four days of the beginning of treatment the canker had disappeared and the M e r l i n had regained enough strength to take food without being force fed. 5 . 2 . 6 Land use change One prime f a c t o r a f f e c t i n g the p r a i r i e M e r l i n p o p u l a t i o n i s man-induced land use change. Sources of information used to p o r t r a y land use change i n the study area come l a r g e l y from the Canada Census r e p o r t s , 1 9 4 1 to 1 9 7 1 (Table 8 ) , a e r i a l photographs (Table 9 ) , and records of the S p e c i a l Areas of A l b e r t a Board's minutes (Table 1 0 ) . A s u b s t a n t i a l expansion of a g r i c u l t u r e between 1 9 4 1 and 1 9 7 1 i n the K i n d e r s l e y area from which M e r l i n s have vanished, as compared to that of the Hanna area where they s t i l l e x i s t , can be seen i n Canada Census s t a t i s t i c s . By the end of t h i s p e r i o d the K i n d e r s l e y area had at l e a s t 677o of land under c u l t i v a t i o n as compared to 2 6 7 ° f o r the Hanna area. A. 127o increase at l e a s t i n c u l t i v a t i o n during t h i s p e r i o d i s i n d i c a t e d f o r the K i n d e r s l e y area as compared to only 47, f o r the Hanna area. A i r photo study of s p e c i f i c areas now or i n the past known to be M e r l i n s i t e s show a s i m i l a r d i f f e r e n c e i n land TABLE 8. The Areas Under C u l t i v a t i o n i n the Census D i v i s i o n s of the Study Area 1941 Hanna, A l t a . (acres) 1,133,506 % 23.1 K i n d e r s l e y , Sask.(acres) 2,431,338 % 55.5 1951 1,338,255 27.2 2,480,875 56.7 1961 1,331,82? 25.02 2,830,066 64.7 1966 1,397,513 25.8 2,883,959 65.8 1971 1,408,079 26.4 2,886,171 67.5 1 crop and f a l l o w 2 r e d u c t i o n due to enlargement of Census D i v i s i o n from Canada Census S t a t i s t i c s to VO TABLE 9. Percentage of Areas under C u l t i v a t i o n W i t h i n M e r l i n Hunting T e r r i t o r i e s Based on Studies of A i r Photos taken between the 1940 's and the 1970 's. Hanna, A l b e r t a K i n d e r s l e y , Saskatchewan 1949 1946 15 51 1962-I65 21 1971 22 1956-61 48 1971 58 1 cropland and hayland ( c u l t i v a t e d ) TABLE 10. The Acreage Under C u l t i v a t i o n (Permit and Lease) i n the S p e c i a l Areas of A l b e r t a 1963 1964 1966 1967 1968 1969 1970 1971 1972 Acres 275,122 273,579 270,470 273,587 273,741 274,885 275,356 391,688 391,155 % of T o t a l * 3.5 3.5 3.5 3.5 3.5 3.5 3.5 5.0 5.0 * S p e c i a l Areas T o t a l Acreage 7,838,826 acres (Stewart and P o r t e r , 1942) from S p e c i a l Areas of A l b e r t a Board's Minutes 4 2 use between the two areas (Table 9 ) ; v i z . 5 8 7 , of the t e r r i t o r y was under c u l t i v a t i o n i n the K i n d e r s l e y area, as compared to 2 2 7 , f o r the Hanna area. In the K i n d e r s l e y area a 37, decrease i n c u l t i v a t e d land from the 1 9 4 0 ' s to the 1 9 5 0 ' s i s apparent i n a i r photos (Table 9 ) . This i s because 3 of the 1 4 M e r l i n s i t e s s t u d i e d i n 1 9 4 6 a i r photos appeared as t o t a l l y c u l t i v a t e d ateas. During the 1 9 4 0 ' s these s i t e s were reseeded to c r e s t e d wheatgrass (Agropyron cristatum) i n the Teo Lake Community pasture to I c o n t r o l e r o s i o n of s o i l by wind. This reseeded pasture appeared as c u l t -i v a t e d crops i n the e a r l y photographs. By the 1 9 5 0 ' s t h i s pasture had taken on the nature of grassland and was considered to be such i n the data from t L v the a i r photos. A i r photo study of 3 9 M e r l i n s i t e s along the South Saskatchewan R i v e r show th a t 9 6 7 . of the area w i t h i n a. one m i l e r a d i u s of n e s t i n g s i t e s i s g r a ssland. The hunting t e r r i t o r i e s s p e c i f i c a l l y s t u d i e d range between 847> minimum and 1 0 0 7 , maximum of land i n g r a s s l a n d . Since 1 9 7 1 an increase i n the amount of c u l t i v a t i o n i n the Hanna area i s to be noted; t h i s i s c l e a r l y recorded i n the S p e c i a l Areas of A l b e r t a Board's minutes (Table 1 0 ) . These show an increase of 1 1 6 , 4 3 7 acres of land i under the j u r i s d i c t i o n of the S p e c i a l Areas Board ( i . e . not deeded land) f o r which leases and permits f o r c u l t i v a t i o n were issued between 1 9 7 0 and 1 9 7 1 . This i s about a 1 . 5 7 o increase i n c u l t i v a t i o n f o r a l l of the S p e c i a l Areas excluding deeded land. Deeded land accounts f o r about 1 9 . 5 7 , of the lands i n the S p e c i a l Areas (Stewart and P o r t e r , 1 9 4 2 ) . A s t a t i s t i c a l comparison between Hanna and K i n d e r s l e y of a number of f a c t o r s which/might c h a r a c t e r i z e M e r l i n nest s i t e s i s given i n Table 1 1 ' . I n g e n e r a l , i t can be s a i d that s i t e "stands" i n the Hanna area were l a r g e r , 43 TABLE 11. A Comparison of C e r t a i n Environmental Factors of Nest S i t e near Hanna, A l b e r t a , and K i n d e r s l e y , Saskatchewan  Feature Category Hanna K i n d e r s l e y X 2 P a =29) (N=14) # % # 7o Stand Acer-Populus 25 86.2 9 64.3 .30 Tree A c e r - S a l i x 2 6.9 1 7.1 2.58 .30 Type Populus-Caraaana 2 6.9 4 28.6 S i t e 10»-19' 2 6.9 4 28.6 Tree 20'-29 f 18 62.1 7 50.0 3.54 .20 Height 30»-39' 9 31.0 3 21.4 S i t e Upland Grove 6 20.7 4 28.6 O r i g i n Windbreak 23 79.3 10 71.4 .68 .50 S i t e Impenetrable 8 27.6 6 42.9 Density Open 21 72.4 8 57.1 .98 .50 S i t e 100m- 5 17.2 7 50.0 S i z e 100m+ 24 82.8 7 50.0 5.06 .05 (1 or dia) Presence Yes 23 79.3 8 57.1 of Bare No 6 20.7 6 42.9 3.05 .10 Branches Presence Yes 26 89.7 6 42.9 of Bare No 3 10.3 8 57.1 10.75 .01 Trunk Presence Yes 20 69.0 13 92.9 of Under No 99 31 1 7.1 3.02 .10 Cover Presence Yes 29 100.0 7 50.0 of u No 0 0 7 50.0 17.1 .001 N e s t ( s ) D a P r o b a b i l i t y that u t i l i z e d S i t e s (Hanna) were the same as n o n - u t i l i z e d s i t e s ( K i n d e r s l e y ) b Nests of crows, hawks, or magpies which could be taken over as nests by M e r l i n . 44 had more bare branches extending above the leafy portions of the windbreak or grove, had more trees whose trunks were bare above the level of two feet, and had more often bare ground beneath the "stand". There were a greater number of upland groves (i.e. natural clumps of Populus around low areas and sloughs) in the Kindersley area. Merlins made greater use of windbreaks and groves which were also used by cattle for shelter in the Hanna area. Nest t "stands" in the Kindersley area tended to differ from those of the Hanna area in thatconly 507=, of the Kindersley "stands" had nests of crows, hawks or mag-pies which would be potential nests for Merlins. It may be surmised that changes probably attributable to agriculture made the Kindersley area no longer a suitable area for these other species to nest. 45 P l a t e 3. M e r l i n Nest S i t e i n Abandoned Windbreak P l a t e 4. M e r l i n Nest S i t e i n Springtime Showing Magpie Nests Used by M e r l i n s To face page 45 46 6. D i s c u s s i o n M e r l i n s n e s t i n g along p r a i r i e r i v e r s are recorded i n the w r i t i n g s of Houseman (1894) and Dippie (1895),,both of whom c o l l e c t e d M e r l i n s along the Bow R i v e r and observed M e r l i n s hunting over the grasslands. Brooks (1896) c i t e s a personal communication r e p o r t i n g that: "Falco r i c h a r d s o n i i i s a common enough b i r d throughout most of the Rocky Mountain r e g i o n . I t breeds i n the Saskatchewan country i n such numbers that one of our c o l l e c t o r s took four sets of eggs i n a s i n g l e season." I n the f i r s t two decades of t h i s century the r a p i d breaking of the p r a i r i e sod, d r a i n i n g of sloughs, the c l e a r i n g of groves of t r e e s , and other a c t i v i t i e s attendent on homesteading, would have r e s u l t e d i n a major l o s s of v.. h a b i t a t and, probably, a major pop u l a t i o n d e c l i n e . During the 1920's and the 1930's the Hanna country was almost vacated by farmers and as the p l a i n s beg^an the slow process of change to grasslands again, the abandoned windbreaks became a v a i l a b l e f o r m e r l i n n e s t i n g s i t e s . Bent , (1931) w r i t e s of Richardson's pigeon hawk ( M e r l i n ) : "This b e a u t i f u l l i t t l e f a l c o n , the p a l e s t of the North American m e r l i n s , i s a b i r d of the Great P l a i n s , breed-i n g mainly i n southern A l b e r t a and Saskatchewan, i n Montana, and i n northwestern North Dakota. I t s summer home i s on the wide r o l l i n g p l a i n s and p r a i r i e s , where they are dotted w i t h small groves of p o p l a r s , aspens, cottonwoods, and other deciduous t r e e s . " Dependence of M e r l i n s on numbers of passerine prey species to nest s u c c e s s f u l l y i s documented i n t h i s study. Conversion of much of the n a t i v e grassland which supports passerine populations to a g r i c u l t u r e has destroyed much of the M e r l i n s ' t r a d i t i o n a l h a b i t a t and much more so i n the K i n d e r s l e y area than i n the Hanna area. In the e a r l y homesteading days from 1895 to 1920, extensive d e s t r u c t i o n of n a t i v e grassland probably produced a marked 47 decline in Merlins in both areas where larger percentages of land with better so i l and moisture occur. In the years following the 1914-18 War, relatively much more of the land in the Hanna area was abandoned and has returned to I . quasi-native grassland. Thirty-six percent more land in the Kindersley area has been brought under cultivation since the 1940's than that in the Hanna area. Moreover, census statistics indicate a 12% increase in cultivated land in the Kindersley area.) during this period, as compared to a 4% increase in the Hanna area. Air photo study of Merlin hunting ranges in both the Kindersley area and the Hanna area show the amount of land under cultivation since the 1940's increased 7%. Differences between land use changes apparent from census statistics and those apparent from air photo study can be explained by the more restricted areas (i.e. hunting territories) considered in air photos. Most Merlin terrritories were at one time occupied farmsites, and these were probably located^on the best farming locations. In the Kindersley area a greater number of these farms would have remained under cultivation since the original sod breaking. In the Hanna area virt u a l l y a l l broken land went back to grass-land, and the better farming sites around old homesteads would be the f i r s t areas to be subjected to cultivation again. From the air photo study of 40 Merlin sites in use successfully since 1971, 95%, had more than 50% of their area grassland; of the remaining sites one was 40%, grassland and one 45%,. In the Kindersley area, air photo study revealed that between 1956 and 1961, the period in which Merlin sites were recorded in the area, 66%, of the 15 sites were more than 50%, grassland; by 1971 only 27%, of these sites were more than 50%, grassland. 48 I f , as the above r e s u l t s suggest, a hunting t e r r i t o r y must have a minimum of about 50% grassland to support a p a i r of M e r l i n s , then the Kind-e r s l e y area cannot be considered as having been a prime M e r l i n n e s t i n g area, at l e a s t s i n c e the 1940's. I t i s suggested that the r e d u c t i o n of grasslands i n the K i n d e r s l e y area since 1961 could have had the e f f e c t of lowering the amount of grassland below the th r e s h o l d necessary to support M e r l i n s f o r many Ki n d e r s l e y s i t e s . Therefore, r e d u c t i o n of grass l a n d emerges as a probable f a c t o r i n the d e c l i n e of M e r l i n s i n the K i n d e r s l e y area. Another very obvious change i n Merl i n s i t e s i n the K i n d e r s l e y area since the time of t h e i r u t i l i z a t i o n by M e r l i n s i s the disappearance of nests of hawks, magpies, and crows which M e r l i n s use. Seven of the 14 s i t e s once occupied by M e r l i n s had no nests of any k i n d which might have been taken over by M e r l i n s i n 1974. I t i s assumed that " c l e a n " farming p r a c t i c e s and extensive c u l t i v a t i o n s have probably c o n t r i b u t e d to the disappearance of hawks, crows and magpies as n e s t i n g b i r d s i n many areas. I n any event, where there are no vacant nests to u t i l i z e , M e r l i n s do not nest i n the p r a i r i e h a b i t a t . D e s t r u c t i o n of n e s t i n g t r e e s by c a t t l e has been suggested as a p o s s i b l e reason f o r the disappearance of M e r l i n s from the K i n d e r s l e y area. (Fox, 1964). When c a t t l e are kept near o l d windbreaks or other trees they w i l l use them f o r s h e l t e r and shade and i n doing so are o f t e n very d e s t r u c t i v e to the tr e e s . However, M e r l i n s were found n e s t i n g even i n a s i n g l e tree on a number of occasions on the p r a i r i e s , and i n none of the K i n d e r s l e y s i t e s u t i l i z e d by Me r l i n s i n the 1950's and 1960's was there any s u b s t a n t i a l r e d u c t i o n of nest-ing trees by the 1970's. M e r l i n nest trees c h a r a c t e r i s t i c a l l y have l i t t l e underbrush and the trunks are well-rubbed up to a l e v e l of about 4' to 5'. 49 P l a t e 5. M e r l i n Nest S i t e i n Abandoned Windbreak Showing E f f e c t of C a t t l e on Windbreaks P l a t e 6. " H i g h l i n i n g " and Breakage by C a t t l e Using an Abandoned Windbreak as s h e l t e r . 50 A notable feature of most of the s i t e s formerly occupied i n the K i n d e r s l e y area i s the l a c k of t h e i r use by c a t t l e . I t must be noted, however, that at many l o c a t i o n s on the p r a i r i e s are remnants of one-time windbreaks and groves that have been u t t e r l y destroyed by c a t t l e and i t i s p o s s i b l e that i n some cases c a t t l e may be instrumental i n the e l i m i n a t i o n of M e r l i n n e s t i n g s i t e s . Much of the increase i n c u l t i v a t i o n i n the Hanna area i s a r e s u l t of a program i n the S p e c i a l Areas by which s e r a i g r a s s l a n d and some climax grassland i s being ploughed and c u l t i v a t e d f o r crops f o r a number of years, a f t e r which they are reseeded:-in e x o t i c grasses such as c r e s t e d wheatgrass. This program i s an attempt to reach a higher l e v e l of primary p r o d u c t i v i t y of grasslands to support more c a t t l e ; the process i s being encouraged throughout the p r a i r i e s (Johnson, 1969). These c u l t i v a t e d grasslands l a c k d i v e r s i t y i n cover species and seem to support low populations of passerine b i r d s (personal o b s e r v a t i o n ) ; i n time, however, they may take on some of the species d i v e r s i t y of n a t i v e grasslands ( n a t i v e grasslands here includes o l d f i e l d as w e l l as climax g r a s s l a n d ) . The Teo Lake pasture i n the Kind-e r s l e y area was seeded to c r e s t e d wheatgrass i n the 1940's f o r wind er o s i o n c o n t r o l . I n e a r l y a e r i a l photographs i t appeared as c u l t i v a t e d land but now has taken on the appearance of a n a t i v e grassland. This pasture land-had enough passerine b i r d s on i t i n the 1950's and 1960's to support n e s t i n g M e r l i n s . Under present management programs i t i s proposed that these c u l t i v a t e d grassland be ploughed and reseeded every few years i n order to keep primary p r o d u c t i v i t y h i g h . Since the widespread i n t r o d u c t i o n of c h l o r i n a t e d hydrocarbon p e s t i c i d e s , e s p e c i a l l y DDT, i n t o many world ecosystems a f t e r 1947, p o p u l a t i o n declines i n a number of birds of prey have been noted. In p a r t i c u l a r , peregrine falcons (Falco peregrinus) i n the northern hemispere, and ospreys (Pandion  haliaetus) and bald eagles (Haliaeetus leucocephalus) i n eastern North America have been affected.. Chlorinated hydrocarbon residues were generally considered the p r i n c i p a l cause of these declines but i t was not u n t i l R a t c l i f f e (1967) noted a s i g n i f i c a n t decrease i n the density of raptor egg s h e l l a f t e r 1947, and showed a c o r r e l a t i o n between decrease i n egg s h e l l weight and an increase i n egg s h e l l breakage that a p h y s i o l o g i c a l basis for these declines was delineated. Hickey and Anderson (1968) have shown that for at l e a s t three species of birds of prey, i n populations where decreases i n egg s h e l l weights have, -amounted to 197« or more, populations were found to be e i t h e r i n a state of decline or had been eixtirpated. These eggshell reductions are based on a comparison of eggs c o l l e c t e d before 1947 (the preorganochlorine i n s e c t i c i d e use date) with eggs c o l l e c t e d after 1947. In our study, egg s h e l l density was measured i n terms of R a t c l i f f e Indices; the regression of R a t c l i f f e Index against DDE i s h i g h l y s i g n i f i c a n t (p <C-01) (Figure 4). The hypothesis of thin eggshells explains well a pathway by which high l e v e l s of DDE can cause a reduction i n raptor pop-u l a t i o n s . This hypothesis states that higher residue l e v e l s r e s u l t i n thinner eggshells which, i n turn, r e s u l t i n lower h a t c h a b i l i t y of eggs (probably through breakage of eggs). I f h a t c h a b i l i t y i s too low, i n s u f f i c i e n t young are produced to replace adult mortality and a population decline r e s u l t s . High residue l e v e l s could also r e s u l t i n abberrant behaviour a f f e c t i n g hunting or care of the o f f s p r i n g so that posthatching mortality occurs and a lowered p r o d u c t i v i t y r e s u l t s . Much study of raptors today i s directed 52 towards i n v e s t i g a t i o n of t h i s phenomena (R. F y f e , personal communication). M e r l i n s would seem to be l e s s s u s c e p t i b l e to the e f f e c t s of DDE i n view of an apparently "normal" p r o d u c t i v i t y c o i n c i d i n g w i t h g e n e r a l l y high p e s t i -c i d e residue l e v e l s . Reduction i n the use of DDT (from which DDE i s de-r i v e d ) and other c h l o r i n a t e d hydrocarbon p e s t i c i d e s since 1971 w i l l , hope-f u l l y , r e s u l t i n some recovery of M e r l i n populations i n any areas a f f e c t e d . No p e s t i c i d e residue or egg s h e l l data f o r M e r l i n s are a v a i l a b l e from Saskatchewan during the p e r i o d of d e c l i n e , i . e . i n the 1960's. However, two eggs from a M e r l i n nest i n Saskatchewan j u s t west of the K i n d e r s l e y area i n 1972 had 2.78 and 3.24 ppm d i e l d r i n . Those l e v e l s of d i e l d r i n from the Saskatchewan M e r l i n s were the h i g h e s t , except forgone, of 166 samples c o l l e c t e d elsewhere, between 1968 and 1973, and f a r above the mean d i e l d r i n l e v e l of 0.70 ppm. Because of the very slow degradation of d i e l d r i n , i t s heavy use i n the 1960's could s t i l l account f o r i t s presence i n the K i n d e r s l e y environ-ment i n the 1970's. Data from our study c l e a r l y i n d i c a t e that both DDE and d i e l d r i n residue are s i g n i f i c a n t l y r e l a t e d to lowered h a t c h a b i l i t y of eggs, however, i t would appear that t h e i r e f f e c t s on A l b e r t a M e r l i n s were not s u f f i c i e n t to cause a d e c l i n e . One aspect of the heavy use of d i e l d r i n i n the K i n d e r s l e y area which was not s t u d i e d i s the e f f e c t i t had on the M e r l i n prey, i . e . on the small b i r d populations of that area. Any d e s t r u c t i o n of small b i r d s could only have had the e f f e c t of i n c r e a s i n g the amount of grasslands M e r l i n s needed to hunt over to get enough prey to s u s t a i n themselves and so reduce f u r t h e r the c a r r y i n g c a p a c i t y of the K i n d e r s l e y area f o r n e s t i n g M e r l i n s . Although the data are by no means s a t i s f a c t o r y f o r comparative purposes the f a c t that s u b s t a n t i a l q u a n t i t i e s of d i e l d r i n were s o l d i n the K i n d e r s l e y area i n the years 1958-1964 appears i n the records of the r u r a l m u n i c i p a l i t i e s (see Table 12). A f t e r 1964, d i e l d r i n use f o r i n s e c t c o n t r o l i n the area was discontinued and other compounds such as chlordane, a l d r i n , and endrin were used. The heavy and widespread use of d i e l d r i n i s p o s t u l a t e d as the f a c t o r which rendered the "coupe de grace" to the M e r l i n s of the Ki n d e r s l e y area.' Records of d i e l d r i n use i n the Hanna area were not a v a i l a b l e . Twelve m o r t a l i t i e s of n e s t l i n g M e r l i n s along the South Saskatchewan R i v e r which cannot be accounted f o r occurred on a p a r t i c u l a r s t r e t c h of the r i v e r which has been an area of low p r o d u c t i v i t y and nest success f o r both M e r l i n s and P r a i r i e Falcons since they were f i r s t looked at i n 1968. This area i s l o c a t e d w i t h i n the S u f f i e l d M i l i t a r y Reserve about h a l f way between Medicine Hat and the Saskatchewan border. At one nest i n t h i s area i n 1969, where 3 well-developed young were found dead, an egg had been c o l l e c t e d e a r l i e r f o r p e s t i c i d e a n a l y s i s . In i t both DDE and d i e l d r i n l e v e l s were f a r above the sample pop u l a t i o n mean of 20.47 and 0.70, being 65.8 and 2.10 ppm, r e s p e c t i v e l y . Of the eggs c o l l e c t e d from the same v i c i n i t y (SSR 23-SSR26) (n=18) i n t h i s and other years, 41% had DDE residue and 50%, had d i e l d r i n residues above the mean of the popu l a t i o n sample. (Appendix V I ) . This would perhaps suggest that these f a l c o n s are feeding on a popu l a t i o n of small b i r d s from a l o c a l contaminated area. Lockie e_t a l , 1969, i n a w e l l documented study presents .data, i n d i c a t i n g that d i e l d r i n used as a sheep dip was r e s p o n s i b l e f o r the very low prod-u c t i v i t y of Golden Eagles ( A q u i l a chrysaetos) i n Scotland i n the e a r l y 1960's. Fo l l o w i n g the 1966 ban of d i e l d r i n from use i n sheep dips p r o d u c t i v i t y returned to a l e v e l considered normal. The data suggested "an inverse r e l a t i o n s h i p between d i e l d r i n l e v e l and success i n breeding". A l e v e l of 54 TABLE 12. SALE OF DIELDRIN IN SOME RURAL MUNICIPALITIES OF SASKATCHEWAN 1955-1965 Year K i n d e r s l e y RM 289 Smiley Marengo Glidden 1955 - - - - -1956 - - - -1957 - - - -1958 $2,380.00 - - - 9 g a l 1959 1,479.00 - 34 g a l - 202 g a l 1960 3,960.00 - • 1627.00 360.00 177 g a l 1961 14,742.70" 2790.00 7915.00 360.00 1408 g a l 1962 12,788.00 3546.00 3624.00 360.00 519- g a l 1963 8,838.00 1636.00 4281.00 - 453 g a l 1964 - - 120.00 -1965 55 lppm seemed to be the level beyond which upsets to normal breeding occurred in Golden Eagles. The discovery of a Merlin in a nest which was apparently suffering from "Frounce" opens speculation into a possible cause of Merlin mortality, that of disease. Kocan and Herman (1971) state: "Naturally occurring infections of T. gallinae have been . reported in raptors, and i t is believed that these birds acquire their infections by eating infected pigeons (Stabler, 1941b; Stone and James, 1969). The large num-ber of feral pigeons in many areas of the country make an excellent source of T. gallinae for birds of prey. There has been some speculation that the decline of certain raptorial species may be directly related to their shift in diet from other wild birds to feral pigeons. Although there is no definite proof of this, the presence of naturally occurring trichomoniasis i s worthy of consideration when studying the population dynamics of birds of prey". Merlins (despite the old name of pigeon hawk) are not large enough to prey on pigeons, and i t is only speculation that perhaps i t is S tarlings (Sturnus vulgaris) which are providing an intermediate host for T. gallinae. Starlings are very common along c l i f f s of the South Saskatchewan River. Reproductive data for Merlins investigated during this study have been summarized (Table 2); the wide ranges of means between data from different years and different areas indicates considerable variation in re-productive success from year to year. Factors which can be considered "natural" (i.e. not influenced by man) that affect reproductive success and could be responsible for this variation have been outlined. In 1973 a cause of the low net production of young Merlins can be attributed to the severe storm during the c r i t i c a l hatching and post-hatching period. Failure of over 4G7o of the nests active prior to the storm can probably be attributed 56 d i r e c t l y to i t . Of over 500 young M e r l i n s produced during the study p e r i o d , twenty-four young M e r l i n s were found dead i n the nest. Of these deaths, 7 were i n the Hanna area and a l l but one were a t t r i b u t e d to the storm. The cause of the remaining m o r t a l i t y i s unknown. Along the South Saskatchewan R i v e r 17 m o r t a l i t i e s were discovered, and only 2 could p o s s i b l y have been a t t r i b u t e d to the storm discussed. The remaining 15, f o r which causes were unknown, were i n a l l years between 1969 and 1974, and at a l l stages of development. The interchange of i n d i v i d u a l M e r l i n s between the Hanna area and the South Saskatchewan R i v e r area serves to e s t a b l i s h the c o n t i n u i t y of the M e r l i n p o p u l a t i o n i n A l b e r t a . There i s no reason to suppose that a s i m i l a r movement between M e r l i n s of the Hanna area and M e r l i n s of the K i n d e r s l e y area d i d not e x i s t . The f a c t that a l l male r e t r a p s were a t , or nearbyj the p o i n t of o r i g i n a l capture lends support to the idea that i t i s the a d u l t male, or a male o f f s p r i n g r a i s e d i n the area, which keeps a s i t e under occupancy year a f t e r year. I f a l l M e r l i n s are removed over a wide area f o r a number of years so that there are no male M e r l i n s w i t h a past h i s t o r y of nest s i t e use f o r that area, reoccupancy of that area would become u n l i k e l y . I n an area l i k e K i n d e r s l e y where M e r l i n s have been absent f o r over a decade and where the few pasture,>areas which might be good M e r l i n n e s t i n g h a b i t a t are so i s o l a t e d from one another and from areas where M e r l i n s do p r e s e n t l y nest, i t seems very u n l i k e l y that they would be reoccupied again by n a t u r a l means. How-ever, i t i s p o s s i b l e that r e - i n t r o d u c t i o n of M e r l i n s , e s p e c i a l l y males, to the i s o l a t e d pockets of M e r l i n h a b i t a t would be s u c c e s s f u l i n r e - e s t a b l i s h i n g M e r l i n s to a formerly vacated area. This might be p o s s i b l e even i n the wheat country of Saskatchewan i f problems encountered by M e r l i n s i n attempting to navigate to a n e s t i n g t e r r i t o r y over oceans of wheat do not prove i n s u r r -mountable. At present, considerable time and research i s being devoted t o the breeding of r a p t o r s , i n c l u d i n g M e r l i n s , under a r t i f i c i a l c o n d i t i o n s , and the r e i n t r o d u c t i o n of these b i r d s to the w i l d (R. Fyfe, personal com-munication),. 58 SUMMARY AND CONCLUSIONS Changes on the Canadian p r a i r i e s w i t h the advent of white men have been many; m o d i f i c a t i o n of the p r a i r i e ecosystem has been as great as, or i g r e a t e r than, i n any of the major ecosystems of the continent and many p r a i r i e species of p l a n t s and animals have been brought to e x t i n c t i o n or very near to i t . The thundering of the f e e t of m i l l i o n s of bison has been repl a c e d by the bawling of c a t t l e and the humming of t i g h t wire on fence posts. The most dramatic changes and, from a n a t u r a l i s t s p o i n t of view, the most devastating changes have been e f f e c t e d by the plough. S o i l , unturned s i n c e the age of i c e , have been again exposed to the p r a i r i e sunshine, wind, and water, and an a g r i c u l t u r a l system l a r g e l y based on annual crops has been e s t a b l i s h e d over l a r g e t r a c t s of land. I n s p i t e of the great changes i n t h e ^ p r a i r i e ecosystem much n a t i v e l i f e remains or i s adapting to the new face of the p r a i r i e s . One such com-ponent of t h i s l i f e i s the M e r l i n and so long as there are trees w i t h s u i t -able nest s i t e s , and smaller b i r d s to hunt, some M e r l i n s "make a l i v i n g " . F o l l o w i n g the disappearance of M e r l i n s from the K i n d e r s l e y area of Saskatchewan i n very recent years (the l a t e 1960's) as recorded by f i e l d surveys, my study was i n s t i t u t e d w i t h the Canadian W i l d l i f e S e r v i c e support to i n t e n s i f y documentation of the major features of the p r a i r i e M e r l i n pop-u l a t i o n and h a b i t a t , and to probe the cause of the disappearance of M e r l i n s from the K i n d e r s l e y area of Saskatchewan. This study, focusing l a r g e l y on the two above mentioned areas, c o n s i s t e d of: a) Y e a r l y surveying of ne s t s , r e c o r d i n g of reproductive success, banding of n e s t l i n g M e r l i n s and banding and trapping a d u l t M e r l i n s ; t h i s was 59 done to o b t a i n data regarding nest s i t e reoccupancy, p r o d u c t i v i t y of M e r l i n s , and M e r l i n p o p u l a t i o n dynamics. b) I n v e s t i g a t i o n of past and curr e n t a g r i c u l t u r a l systems and t h e i r e f f e c t s on the M e r l i n population. c) Observation o f prey composition and in c i d e n c e , weather, disease, and human i n t e r f e r e n c e and t h e i r e f f e c t s on n e s t i n g M e r l i n s , and d) S t a t i s t i c a l a n a l y s i s of p e s t i c i d e residue l e v e l s i n egg t i s s u e s , and t h e i r r e l a t i o n s h i p to egg h a t c h a b i l i t y . I n c o n c l u s i o n , the f o l l o w i n g statements can be made: 1) P r a i r i e grassland areas have been i n a continuous s t a t e of change since the a r r i v a l of settlement, and p r a i r i e w i l d l i f e populations have been a f f e c t e d . 2) The most l o n g - l a s t i n g e f f e c t on M e r l i n s (and other p r a i r i e w i l d l i f e ) has been as a r e s u l t of the plough and the r e s u l t i n g r e d u c t i o n i n the d i v e r s i t y of p r a i r i e h a b i t a t . 3) Abandonment of p r a i r i e farmlands has r e - e s t a b l i s h e d grasslands i n dryer p a r t s of the p r a i r i e s and abandoned windbreaks have r e s u l t e d i n p o t e n t i a l n e s t i n g h a b i t a t f o r M e r l i n s and other p r a i r i e b i r d s . 4) The process of ploughing p r a i r i e grasslands s o i l s i s a cont-i n u i n g process, though, and of l a t e i s expanding; present a g r i c u l t u r a l p o l i c y i n A l b e r t a i a advocating the ploughing and reseeding of grasslands to e x o t i c species such as c r e s t e d wheatgrass; with the improved farming technology today these lands w i l l never be reclaimed by grasslands again. I t may there f o r e be concluded from my comparative study t h a t , i f the M e r l i n i s to be maintained on the Canadian p r a i r i e s , we must maintain a g r i -c u l t u r a l systems which w i l l leave large segments of the p r a i r i e landscape i n a near n a t u r a l s t a t e , such as the not - s o - h i g h l y modified r a n g e - l i v e s t o c k systems. The r e d u c t i o n of f a c t o r s which reduce the d i v e r s i t y of p r a i r i e l i f e , such as the use of p e s t i c i d e s and h e r b i c i d e s , must be considered a par t of maintaining the n a t u r a l p r a i r i e system. v 60 L i t e r a t u r e C i t e d Amadon, D., and L. Brown, 1968. Eagles, Hawks, and Falcons of the World. McGraw-Hill Co., N.Y. Bent, A.C. 1938. L i f e H i s t o r i e s of North American B i r d s of Prey. P a r t 2. U.S. N a t l . Museum B u l l . 170. 482 pp. B l u s , L . J . , C D . G i s h , A.A. B e l i s l e , & R.M. Prouty. 1972. Logarithmic R e l a t i o n s h i p of DDE Residues to E g g s h e l l Thinning. Nature, Lond., 235, 376.-7. Brooks, W.E. 1896. Remarks on Richardson's M e r l i n . I b i s 1896, 222-28. Cody, M.L., 1968. On the Methods of Resource D i v i s i o n i n Grassland B i r d Communities. Amer. N a t u r a l i s t 102(924): 107-147. Cooke, A.S. 1973. S h e l l Thinning i n Aviam Eggs by Environmental P o l l u t a n t s . E n v i r o n . P o l l u t . (4) pp 85-152. Dawson, C.A. & E.R. Younge. 1940. P i o n e e r i n g i n the P r a i r i e Provinces. The MacMillan Co., Toronto. D i p p i e , G.F. 1895. Nesting of Richatdson's M e r l i n . O o l o g i s t 12: 135-36. F i s h e r , A.K. 1863. The Hawks and Owls of the United S t a t e s . Wash., D.C. Fox, G.A. 1964. Notes on the Western Race of the Pigeon Hawk. Blue Jay 12 (4): 140-147. Fox, G.A. 1971. Recent Changes i n the Reproductive Success of the Pigeon Hawk. J . W i l d l . Mgmt. 35 (1): 122-128. Fyf e , R.W., J . Campbell, B. Hayson, and K. Hodson. Regional P o p u l a t i o n Declines and Organochlorine I n s e c t i c i d e s i n Canadian P r a i r i e Falcons. The Canadian F i e l d - N a t u r a l i s t 83 ( 3 ) ; 191-200. Gray, J.H. 1967. Men Against the Desert. Modern Press, Saskatoon. Hardy, W.G., ed. 1967. A l b e r t a , A N a t u r a l H i s t o r y . Evergreen Press L t d . , Vancouver. Hickey, J . J . , and D.W. Anderson. 1968. C h l o r i n a t e d Hydrocarbons and E g g s h e l l Changes i n R a p t o r i a l and F i s h - e a t i n g B i r d s . Science, N.Y. 162: 271-273. Houseman, J.E. 1894. Nesting Habits of Richardson's M e r l i n . O o l o g i s t 11:236-237. 61 L i t e r a t u r e C i t e d , cont'd. Johnston, A. 1970. C l a s s i f i c a t i o n of Rangelands i n A l b e r t a . A l b e r t a Department of A g r i c u l t u r e . Kocan, R.M., and CM. Herman, i n I n f e c t i o u s and P a r a s i t i c Diseases of Wild B i r d s . J.E. Davies, R.C. Anderson, L. Narstead, and D.O. T r a i n e r , e d i t o r s . Iowa State Univ. Press Iowa, 1971. L o c k i e , J.D., D.A. R a t c l i f f e , and R. B a l h a r r y . 1969. Breeding Success and Organochlorine Residues i n Golden Eagles i n west Scotland. J . A p p l i e d E c o l . 6(3): 381-389. Moss, E.H. 1955. The V e g etation of A l b e r t a . B o t a n i c a l Review 21(9): 493-567. Owens, R.A. 1971. MSc. Thesis (unpubl) The U n i v e r s i t y of Calgary. Calgary, A l t a . R a t c l i f f e , D.A. 1967. Decrease i n E g g s h e l l Weight i n C e r t a i n B i r d s of Prey. Nature 215: 208-210. Rochenbauch, D. 1969. "The Good and the Bad Years" (Tr.ansl. from German) Wi l d Und Hund. 72(15): 357-358. S a l t , W.R., and A.L. W i l d . 1958. The B i r d s of A l b e r t a . Gov. A l t a . Queen's P r i n t e r . Edmonton. Smith, P.J., ed. 1972. The P r a i r i e Provinces. U n i v e r s i t y of Toronto Press. Sopher, J.D. 1964. The Mammals of A l b e r t a . Gov. A l t a . Queen's P r i n t e r . Edmonton. Stewart, A., and W.D. P o r t e r . 1942. Land Use C l a s s i f i c a t i o n i n the S p e c i a l Areas of A l b e r t a . Ottawa Dom. of Can. Dept. of A g r i c . P u b l . #731 Tech. B u l l . #39. 73pp. Temple, S. 1970. MSc. Thesis. The e v o l u t i o n And Systematics of the North American M e r l i n s . C o r n e l l U n i v e r s i t y . I t h a c a , N.Y. Watts, F.B. 1960. The N a t u r a l Vegetation of the Southern Great P l a i n s of Canada. Geographical B u l l e t i n 14: 25-43. Weins, J.A. 1973. P a t t e r n and Process i n Grassland B i r d Communities. E c o l o g i c a l Monographs 43 (2): 248-62 APPENDIX 1 AGRICULTURAL CHEMICALS: MEAN RESIDUE LEVELS IN EGGS OF RICHARDSON'S MERLIN 1969-1973 Residue (ppm wet) DDE D i e l d r i n Heptachlor Mercury 1969 1 (N = 12) 23.2 0.99 1.05 0.36 2 (N = 2) 7.82 0.22 0.94 0.22 1970 1 (N = 38) 9.53 .33 0.70 0.22 2 (N = 5) 29.11 0.46 0.37 0.37 1971 1 (N = 13) 25.57 0.57 0.48 0.29 2 (N 4) 47.17 0.95 0.35 0.15 1972 1 (N = 44) 9.60 0.59 0.31 0.14 2 (N = 15) 20.60 0.78 0.33 0.22 1973 1 (N = 13) 10.76 1.40 0.88 0.07 2 (N = 18) -. 21.36 0.74 0.58 0.13 MEAN: 20.47 .70 .60 .22 1 Egg c o l l e c t e d during incubation p r i o r to hatching 2 Eggs c o l l e c t e d dead af t e r incubation. * Unpublished data from C.W.S. F i l e s , Edmonton 63 APPENDIX 11 ANALYSIS OF FACTORS OF NEST SITES USED IN ALBERTA BY RICHARDSON'S MERLIN Factor Category Nestings # 1 Site Tree Type Si t e Tree Height S i t e O r i g i n Undercover S i t e Density S i t e Size Presence of Bare Branches Bare Trunk Nest Type Nest Height Acer-Populus Acer-Salix Acer-Caragana 10' - 19' 20' - 29' 30' - 39' Over 39' Upland Grove River Grove Windbreak Bare Ground' Less Than 6" 6" - 2' Over 2' Impenetrable See Through Bare Less Than 100 yd 100 yd Over 300 yd Yes No Less Than 2' 2' - 6' Over 6* Open Enclosed 6' - 10' 11'- 15' 16'- 20' 21'- 26' Over 26' 84 10 57 15 56 35 45 .24 43 80 30 12 53 51 37 117 2 21 71 136 15 10 105 36 50 71 44 47 9 10 12 55.2 7.2 37.6 10.0 37.1 23.2 29.7 15.9 31.1 53.0 20.6 2.2 36. 34. ,3 .9 24.5 74.2 1.3 .14.0 47.2 90.0 10.0 6.6 69.6 23.8 41. 58. 37.6 35.9 7.7 8.5 10.3 64 APPENDIX 11 (Continued) Factor Category Nestings # % Nest Tree Type Acer Populus  S a l i x  Caragana 50 62 4 0 43.1 53.5 3.4 0 Nest Tree Height 0' - 9' 10' - 19' 20' - 29' 30* - 39' Over 39' 1 23 44 23 23 0.8 20.2 38.6 20.2 20.2 Canopy over Nest 0' - 5' 6' - 10' Over 10' 35 36 35 33.0 34.0 33.0 L i v e Branches Below Nest 0' - 5' 6' - 10' Over 10' 47 27 25 47.5 27.3 25.2 P o s i t i o n of Nest Distance To Roads Distance To Occupied B u i l d i n g s Against Trunk In Crotch On Limb Less than % mi \ - % mi % - 1 mi Over 1 mi Less Than \ mi \ - \ mi % - 1 mi Over 1 mi 48 42 16 31 3 33 33 0 2 13 85 45.2 39.6 15.2 31.0 3.0 33.0 33.0 0 2.0 13.0 85.0 Distance To Water Less Than \ mi - % mi % - 1 mi Over 1 mi 29 4 58 9 29.0 4.0 58.0 9.0 6 5 APPENDIX I I I PREY SPECIES USED BY RICHARDSON'S MERLINS The occurrences of feather and s k e l e t a l remains found and i d e n t i f i e d from 2 0 7 0 feather and other prey remains found i n nests and at p l u c k i n g perches i s given i n Table 1 3 . C h e s t n u t - c o l l a r e d longspurs ( C a l c a r i u s ornatus) and Horned Larks (Eremophila a l p e s t r i s ) together formed 9 7 7 , of prey remains. Of the stomach (crop) contents of Richardson's M e r l i n s examined by F i s h e r ( 1 8 6 3 ) , 2 contained b i r d remains, 1 i n s e c t s , and 1 was empty. Amadon and Brown ( 1 9 6 8 ) give the r e l a t i v e p roportions of food items of M e r l i n s as being about 8 0 % b i r d , 57, mammal and 1 5 7 , i n s e c t remains. Although n e s t i n g M e r l i n s are sustained almost e n t i r e l y by small b i r d s , rodent remains were found twice, and M e r l i n s are known to feed on in s e c t s i n the study area. A f a m i l y of new fledged M e r l i n s was seen pursuing and e a t i n g grasshoppers during a heavy hatch of these i n s e c t s (D. O'Dell, personal communication). I n s e c t remains d i d not show up i n prey remains c o l l e c t e d . I f M e r l i n s are hunting an area of one mile r a d i u s as behaviour observations would lead me to b e l i e v e , then, based on i n t e r p r e t a t i o n of 1 9 7 1 a i r photos f o r 4 0 M e r l i n s i t e s , t y p i c a l l y M e r l i n s are using an area composed of about 757o g r a s s l a n d and 257o c u l t i v a t e d land. I n a study immediately adjacent to my study area, Owens ( 1 9 7 1 ) has shown that breeding populations of p r a i r i e b i r d s have d e n s i t i e s o f ^ 2 7 . a n d 5 4 . p a i r s / 1 0 0 acres on grazed and undisturbed grasslands r e s p e c t i v e l y , 4 1 . 5 p a i r s / 1 0 0 acres on mowed hayland * ( n a t i v e ) , and only 6 . 3 and 7 . 5 p a i r s / 1 0 0 acres on seeded and f a l l o w c u l t i -vated land r e s p e c t i v e l y . I t i s obvious that M e r l i n s are hunting pre-dominantly over grasslands where small b i r d populations are g r e a t e s t . TABLE 13. 66 SPECIES UTILIZED AS PREY BY MERLINS Horned Lark (Eremophila a l p e s t r i s ) Chestnut - c o l l a r e d Longspur ( C a l c a r i u s ornatus) Sparrows: Vesper Sparrow (Poecetes gramineus) Savannah Sparrow (Passerculus sandwichensis) Chipping Sparrow ( S p i z e l l a passerina) U n i d e n t i f i e d Sparrows B l a c k b i r d s : -Red-winged B l a c k b i r d (Agelaius phoeniceus) Brown-headed Cowbird (Molothrus ater) Western Meadowlark ( S t u r n e l l a neglecta) Others: McCowans Longspur (Rhynchophanes mccownii) Lark Bunting (Calamospiza melanocorys) Pine S i s k i n (Spinus pinus) Cedar Waswing (Bombycilla cedrorum) Eastern K i n g b i r d (Tyrannus tyrannus) Red Phalarope (Phalaropus f u l i c a r i u s ) Spotted Sandpiper ( A c t i t i s macularis) U n i d e n t i f i e d Shorebirds Richardson's Ground s q u i r r e l (juv) ( C i t e l l u s r i c h a r d s o n i i ) Other Rodents (Family Crecidae-mice and/or v o l e s ) 67 TABLE 14. PAIR DENSITY AND SPECIES ABUNDANCE OF GRASSLAND BIRDS IN GRASSLAND AND AGRICULTURAL REGIMES (Based on Owens, 1971) Ha b i t a t # Pairs/100A Species Most Abundant Grassland (Undisturbed) Grassland (Grazed and Mowed) 54.5 27.4-41.5 Sparagues P i p i t B a i rd*s Sparrow Savannah Sparrow Western Meadowlark Clay-coloured Sparrow C h e s t n u t - c o l l a r e d Longspur Western Meadowlark C u l t i v a t e d (Seeded and Fallow) 6.3-7.5 Vesper Sparrow Horned Lark 68 Owens (197.1) a l s o i n d i c a t e s that Horned Larks, and C h e s t n u t - c o l l a r e d Longspurs . are found abundantly on grazed n a t i v e grasslands, h a b i t a t s w i t h the l e a s t amount of cover (Table 15). The preponderance of Horned Larks, C h e s t n u t - c o l l a r e d Longspurs and Vesper Sparrows i n M e r l i n prey i n d i c a t e d the preference of M e r l i n s f o r open h a b i t a t over which to hunt. This i s to be expected since i t i s here where prey would f i n d the l e a s t amount of escape cover. Despite the higher breeding d e n s i t y of small b i r d s i n undisturbed grassland t h i s h a b i t a t would probably be used l e s s by hunting M e r l i n s because of the escape cover a v a i l a b l e . I n my study area most of the grassland were grazed. The h a b i t a t most p r e f e r r e d by Horned Larks, next to c u l t i v a t e d land, i s grazed g r a s s l a n d (Owen 1971). Weins (1973) s t a t e s , "Again, however, the response of i n d i v i d u a l species to grazing e f f e c t s was more c l e a r cut. Horned Lark d e n s i t y was greater i n grazed p l o t s and, at Pawnee, i n p l o t s subjected to heavy summer gr a z i n g . Western Meadowlarks and Grasshopper Sparrows, on the other hand, were more numerous on ungrazed p l o t s . . . " This preference of Horned Larks f o r open h a b i t a t would i n p a r t account f o r the h i g h per-centage of Horned Larks found i n the M e r l i n s ' d i e t . A second f a c t o r accounting f o r heavy u t i l i z a t i o n of Horned Larks and C h e s t n u t - c o l l a r e d Longspurs i s that of the behaviour of these b i r d s . Cody (1968) i n h i s work on the Pawnee IBP Grassland Study Area has devised a s c a l e of sawtooth curves r e p r e s e n t i n g feeding behaviour i n fourteen North American species s t u d i e d ( F i g . 5 ) . H i s graph shows the distance moved over a given time span, and the number and d u r a t i o n of stops, made i n feeding behaviour. I t i s i n t e r e s t i n g to note that the two most " a c t i v e " b i r d s shown are Horned Larks and C h e s t n u t - c o l l a r e d Longspurs. Perhaps these b i r d s ' 69 TABLE 15 AVERAGE NUMBER OF BIRDS, OF SOME SELECTED SPECIES, RECORDED AT ROADSIDE STOPS OF DIFFERENT LAND-USE SUB-TYPES. LAND-USE SPECIES p w pq Pi & EH CO M P ta t=> P w EH CO H P P W N P i o EH 1=3 CJ \, P W N as p w S 3 > M EH !=> a CO o ta H P M ta PQ Number of stops Native species 5 3 11 10 10 6 Sprague's P i p i t 13.6 10.6 4.5 .4 0 0 Baird's Sparrow 12.6 9.6 4.2 .8 0 0 Savannah Sparrow 9.0 5.0 6.0 6.8 8.5 5.6 Western Meadowlark 8.4 10.6 8.4 7.6 7.0 5.6 Ch e s t n u t - c o l l a r e d Longspur 3.4 6.6 12.3 .4 .2 .5 Vesper Sparrow .6 3.6 3.9 6.7 5.1 6.6 Cl a y - c o l o r e d Sparrow 2.0 2.0 1.1 4.6 1.5 3.0 Horned Lark 1.6 1.6 4.4 2.4 5.5 3.5 Barn Swallow .6 0 .4 2.8 1.0 4.8 Introduced species Ring-necked Pheasant 0 0 0 .8 .5 .6 S t a r l i n g 0 0 0 14.2 11.8 20.6 House Sparrow 0 0 0 4.9 0 7.0 Rock Dove 0' 0 0 2.1 .3 3.1 Notes: N e i t h e r Roadside Count route passed any r e c e n t l y mowed n a t i v e g r a s s l a n d , so there i s no column f o r the "mowed''sub-type of land-use, (from Owens, 1971) To face page 70 70 Figure 5. Feeding Behaviour of Grassland B i r d s (from Cody, 1968) . Sawtooth curves r e p r e s e n t i n g feeding behaviour i n fourteen North American species s t u d i e d . The h o r i -z o n t a l p a r t of a "tooth" i s the d u r a t i o n of the average stop i n seconds; the r a t i o : between t h i s i n t e r v a l and that between successive " s t a r t s " i s the p r o p o r t i o n of the time spent s t a t i o n a r y during a feeding sequence; the slope of the l i n e common to a l l teeth i n a curve i s p r o p o r t i o n a l to the average speed of progression f o r the species during feeding. 71 more a c t i v e feeding behaviour makes them more n o t i c e a b l e to M e r l i n s and the r e f o r e e x p l a i n s i n pa r t the higher r a t e of pred a t i o n on them. 72 APPENDIX IV COMMON PLANTS OF THE STUDY AREA (SPECIMENS LODGED IN THE BOTANY DEPARTMENT HERBARIUM) UNIVERSITY OF BRITISH COLUMBIA A. SOUTH SASKATCHEWAN RIVER GRASSES Boutelous g r a c i l i s (HBK.) l a g . S t i p a comata T r i n . and Rupr. K o e l e r i a c r i s t a t a (L.) Pers. Oryzopsis hymenoides (Roem. & Sc h u l t . ) Bromis inermis Leyss. Agropyron c r i s t a t u m (L.) Gaertn. S i t a n i o n h y s t r i x (Nutt.) J.G. Smith Festuca spp. Poa sp. FORBS Plantage p u r s h i i R & S Ranunculus glaberrimus Hook. Smi l a c i n a s t e l l a t a (L.) Desf. Thermopsis r h o m b i f l o r a (Nutt.) Richards Lygodesmia .juncea (Pursh) D. Don A l l i u m t e x t i l e Nels. & Macbr. V i o l a n u t t a l i (Pursh) Blue grama Spear grass June Grass Indian r i c e grass Awnless brome Crested Wheatgrass S q u i r r e l t a i l Fescue Blue grass Pursh's P l a n t a i n Buttercup Star-flowered solomon's s e a l Golden bean Skeletonweed P r a i r i e onion N u t t a l ' s v i o l e t APPENDIX IV (Continued) SOUTH SASKATCHEWAN RIVER FORBS Lithospermum canescens (Michx.).Jehm. Tragopogon dubius Scop. Malvastrum coccineum (Pursh) A. Gray Erysimum asperum (Nutt.) DC. Pentstemon n i t i d u s Dougl. P o t e n t i l l a anserina L. Lomatium v i l l o s u m Raf. Comandra p a l l i d a A.D.C. Astragalus p e c t i n a t u s Dougl. T h l a s p i arvense L. A c h i l l e a m i l l e f o l i u m L. Cleome s e r r u l a t a Pursh L i s t r i s punctata Hook Astragalus spp.  Hedysarum spp. SHRUBS & TREES  Populus a u g u s t i f o l i a James Symphoricarpos o c c i d e n t a l i s (Hook) S a l i x spp. Rosa spp. Populus spp. Hoary puccoon Y e l l o w goat's beard S c a r l e t Mallow Western W a l l f l o w e r Smooth blue beardtongue Silverweed H a i r y - f r u i t e d p a r s l e y Pale comandra Narrow-leaved m i l k - v e t c h Stinkweed Yarrow S p i d e r f l o w e r , Pink cleome Dotted b l a z i n g s t a r M i l k - v e t c h Sweet-broom Narrow-leaved c o t t o n wood Wolfberry, Buckbrush Willow Rose Poplar 74 APPENDIX IV (Continued) B. HANNA-YOUNGSTOWN, ALBERTA AREA GRASSES Festuca s c a b r e l l a Torr. Rough Fescue A g r o s t i s scabra W i l l a Northern bentgrass Agropyron g r i f f i t h s i i S c r i b n . & Smith A. dasystachyum (Hook.) Agropyron s m i t h i i Rydb. Western wheatgrass Agropyron subsecundum (Link ) K i t c h c . Bearded wheatgrass Boutelous g r a c i l i s (HBK.) Lag. Blue grama K o e l e r i a c r i s t a t a (L.) Pers. June grass Bromus inermis Leyss. Awnless brome Hordeum iubatum L. Wi l d b a r l e y Poa i n t e r i o r Rydb. Inland blue grass Poa p r a t e n s i s L. Kentucky blue grass Poa p a l u s t r i s L. Fowl blue grass Beckmannia syzigachne (Steud.) Fern. Slough grass S t i p a comata T r i n . & Rupr. Spear grass Agropyron c r i s t a t u m (L.) Gaertn. Crested wheatgrass S t i p a v i r i d u l a T r i n . Green needlegrass Calamagrostis montanensis S c r i b n . P l a i n s reedgrass Calamagrostis canadensis (Michx.) Beauv. Marsh reedgrass D i s t i c h i l i s s t r i c t a (Torr.) Rydb. A l k a l i grass Calamagrostis n e g l e c t a (Ehrh.) Gaertn. Narrow reed grass Trisetum spicatum (L.) R i c h t . Spike t r i s e t u m Muhlenbergia sp. Muhly 75 APPENDIX IV (Continued) B. HANNA-YOUNGSTOWN. ALBERTA AREA GRASSES S t i p a spartea T r i n . C a l a m o v i l f a l o n g i f o l i a (Hook.) S c r i b n . Poa ampla Merr. H e l i c t o t r i c h o n h o o k e r i (Scrion.) - Avena Hookeri S c r i b n . FORBS Ranunculus glaberrimus Hook V i o l a n u t t a l l i Pursh Ranunculus abortivus L. Cerastium arvense L. A r n i c a fulgens Pursh Zygadenus venenosus Wats. - Z. gramineus Rydb. Petalostemon purpureum (Vent.) Rydb. G r i n d e l i a squarrosa (Pursh) Dunal Solidago decumbens Greene G u t i e r r e z i a sarothrae (Pursh) B r i t t . & Rusby Orthocaropus lutens Nutt. A r t e m i s i a f r i g i d a W i l l d . Oenthera n u t t a l l i i Sweet Anaphalis margaritacea (L.) C B . C l a r k Campanula r o t u n d j f o l i a L. Solidago g r a m i n i f o l i a (L.) S a l i s b . Penstemon procerus Dougl. Porcupine grass Sand grass B i g bluegrass Hooker's oat grass Buttercup N u t t a l l ' s v i o l e t Buttercup F i e l d chickweed A r n i c a Death camas Purple p r a i r i e c l o v e r Gumweed Mountain goldenrod Broomwood, Matchbrush Owl c l o v e r S i l v e r sage White evening primrose P e a r l y e v e r l a s t i n g B l u e b e l l , H a r e b e l l Flat-topped goldenrod Slender beardtongue 76 APPENDIX IV (Continued) B. HANNA-YOUNGSTOWN, ALBERTA AREA FORBS Penstemon g r a c i l i s Nutt. Galium boreale L. Antennaria p a r v i f l o r a Nutt. Polygonum convulvulus L. Anemone m u l t i f i d a P o i r . A r t e m i s i a sp. Chamaerhodos e r e c t a (L.) Bunge Aster pansus (Blake) Cronq. Oxytropis s e r i c e a Nutt = ). macounii Z i z i a aptera (A. Gray) Fern. Erysimum inconspicuum (S. Wats.) MacM. Medicago s a t i v a L. Cleome s e r r u l a t a Pursh C a s t e l l e j a coccinea (L.) Spreng. A c h i l l e a m i l l e f o l i u m L. G a i l l a r d i a a r i s t a t a Pursh Penstemon erianthus Pursh Tragopogon dubius Scop. V i o l a nephrophylla Greene Antennaria campestris Rydo. Anemone patens (Bess.) Koch Phlox h o o d i i Richards Androsace s e p t e n t r i o n a l i s L. V i o l a adunca J.E. Smith L i l a c - f l o w e r e d beardtongue Northern bedstraw Pussytoes Goosefoot Cut-leaved anemone Wormwood Chamaerhodos White p r a i r i e a s t e r Locoweed Meadow parsnip Small-flowered rocket A l f a l f a S p i d e r f l o w e r , Pink cleome S c a r l e t paintbrush Yarrow Great-flowered g a i l l a r d i a Crested beardtongue Yellow goat's-beard Northern bog v i o l e t P r a i r i e e v e r l a s t i n g Crocus anemone Moss phlox Pygmyflower E a r l y blue v i o l e t 77 APPENDIX IV (Continued) B. HANNA-YOUNGSTOWN, ALBERTA AREA FORBS P o t e n t i l l a sp. Plantago p u r s h i i R. & S. Carex e l e o c h a r i s B a i l e y TREES AND SHRUBS Acer negundo L. Caragana arborescens Lam. S a l i x exigua Nutt. Thermopsis r h o m b i f o l i a (Nutt.) Richards Elaeagnus commutata Bernh. Symphoricarpos o c c i d e n t a l i s Hook. Rosa woodsii L i n d l . C i n q u e f o i l Pursh's p l a n t a i n Low sedge Manitoba maple Common caragana Willow Golden bean S i l v e r b e r r y , Wolf Willow Wolfberry, Buckbrush P r a i r i e rose 78 APPENDIX V MERLIN POPULATION AND NEST DATA KEY Data on f i l e i n L i b r a r y Copy, Main L i b r a r y , U n i v e r s i t y of B.C. A bland equals missing or unknown value. Column 1. 2,3 4. 5,6,7. 8,9,10,11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. Region S i t e I d e n t i f i c a t i o n # Year Area Designation L o c a t i o n # Eggs Analysed Occupied # Eggs Layed # Eggs hatched # Young fledged # Pre-hatched Eggs Removed # V i s i t s to S i t e Beta Backscatter Great Horned Owl Flushed Nest Type 1. Hanna 2. South Saskatchewan R. 3. K i n d e r s l e y 1. -85 1. 1968 2. 1969 3. 1970 4. 1971 5. 1972 6. 1973 7. 1974 eg. SNW, YTE. eg. F10, L 1 0 2 (7 = zero) 1. Yes 2. No (7 (8 (9 0) Yes) NO) 1. Yes 2. NO 1. Yes 2. No 1. Open (Crow or Hawk) 2. Enclosed (Magpie) 79 APPENDIX V KEY (Continued) 22. Nest Height 1. 0 - 5 ' 2. 6' - 10' 3. 11' - 15' 4. 16' - 20' 5. .21' - 25' 6. More than 25' 23. Nest Tree Type , 1. Acer 2. Populus 3. S a l i x 4. Caragana 24. Nest Tree Height 1. 0 - 9' 2. 10' - 19' 3. 20' - 29' 4. 30' - 39' 5. More than 39' 25. L i v e Branches Above Nest 1. 0 - 5 ' 2. 6' - 10' 3. More than 10' 26. L i v e Branches Below Nest 1. 0 - 5 ' 2. 6' - 10' 3. More than 10' 27. P o s i t i o n of Nest 1. Against Trunk 2. In Crotch 3. Out on Limb 28. S i t e Tree Type 1. Acer-Populus 2. A c e r - S a l i x 3. Po p u l u s - S a l i x 29. S i t e Tree Height 1. 0 - 9 ' 2. 10* - 19' 3. 20' - 29' 4. 30' - 39' 5. More than 39' 30. S i t e O r i g i n 1. Upland Grove 2. Ri v e r Grove 3. Windbreak 80 APPENDIX V KEY (Continued) 31. Undercover 32. Density of S i t e 33. Size of S i t e ( d i a . or L.) 34. Bare Branches Above Foilage 35. Bare Trunk Benearth Foilage 36,37,38 % Grassland 1 mile Radius 39,40,41 7o Hay (Cult.) 1 mile Radius 42,43,44. 7o C u l t i v a t e d 1 mile Radius 45. Distance to Roads 46. Distance to Occupied Bldgs. 47. Distance to Water 1. Bare Ground 2. Less than 6" 3. 6" - 2' 4. More than 2' 1. Impenetrable 2. See Through 3. Bare 1. Less than 100 yd. 2. 100 - 300 yd. 3. More than 300 yd. 1. Yes 2. No 1. Less than 2' 2. 2' - 6' 3. More than 6' 1. Less than k 2. % mile - % mile 3. \ mile - 1 mile 4. More than 1 mile 1. Less than \ mile 2. \ mile - % mile 3. % mile - 1 mile 4. More than 1 mile 1. Less than % mile 2. % mile - % mile , 3. % mile - 1 mile 4. More than 1 mile 48. 49. 50. Male Behaviour 1st v i s i t Male Behaviour Egg v i s i t Male Behaviour Banding v i s i t 1. 2. 3. Aggressive Noisy Quiet . 81 APPENDIX V KEY (Continued) 51. Female Bhvr. 1st v i s i t 52. Female Bhvr. Egg v i s i t 53. Female Bhvr. Banding v i s i t 54. Male Trapped 55. Male Already Banded 56. Male Years Since Banding 57. Male Age at Banding 58. Male Region of Banding 59,60 Male S i t e # Where Banded 61,62,63 Male M i l e s from Banding 64. Female Trapped 65. Female Already Banded 66. Female Years Since Banding 67. Female Age at Banding 68. Female Region of Banding 69,70. Female S i t e # Where Banded 71,72,73. Female M i l e s from Banding 74. Nest A v a i l a b l e ( K i n d e r s l e y ) 1. Aggressive 2. Noisy 3. Quiet 1. Yes 2. No 1. Yes 2. No 1. Hanna-Youngstown 2. South Saskatchewan R i v e r 1 - 8 5 1. Yes 2. No 1. Yes 2. No 1. Hanna-Youngstown 2. South Saskatchewan R i v e r 1 - 8 5 1. Yes 2. No 82 APPENDIX VI Merlin P e s t i c i d e Data Key Data on f i l e i n L i b r a r y Copy, Main Libr a r y , U n i v e r s i t y of B.C. Column ' 1. Region 1. Hanna-Youngstown 2. South Saskatchewan River 3. Kindersley 2,3. 4. Si t e I d e n t i f i c a t i o n # Year 5,6,7. 8,9,10. 11. 13-18. 20-25. 27-32. 34-39. 41-44. 45-49. 51-55. 57-60. Area Designation S i t e Location Designation E'gg.# DDE Residue D i e l d r i n Residue Heptachlor Epoxide Residue Mercury (Hg) Residue R a t c l i f f e Index (Rl) Random or Dead Egg Nest Hatched or Nest F a i l e d Condition of Egg 1-85 1. 1968 2. 1969 3. 1970 4. 1971 5. 1972 6. 1973 7. 1974 eg. BOW, RDR, SSR, YTE eg. B3, 015, 24A 83 APPENDIX VII dichloro-diphenyl-trichloroethane endo-exo isomer of 1,2,3,4,10,10-hexachloro-6,7-epoxy-l,4,4a,5,6,7,8, 8a,octahydro-1,4-5,8-dimethanonaphthalene (HEOD) 1,2,3,4,10,10-hexachloro-l,4,4a,5,8,Sa-hexahydro-l, 4-exo-5,8-endo-dimethanonaphthalene (HHDN) 1,2 ,'3,4,10,10-hexachloro-6,7-epoxy-l ,4,4a,5,6,7,8, 8a-octahydro-l,4-exo-5,8-exo-dimethanonaphthalene 1,2,4,5,6,7,10,10-octachloro-4,7,8,9-tetrahydro-4, 7-endo-methyleneindane 1,4,5,6,7,10,10-heptachloro-4,7,8,9-tetrahydro-4, 7-endomethyleneindene f i 14il-IA!".B151718847 j. 481 SNHH07 :l 7 J 5 5 5 7 3 1 4 2 H A F B i 5 i ? 3 8 8 2 7 1 0 5 2 C P W J 0 5 1 7 1 i 232HAWriO':.; 11 j 8 i V 1 3 3 2 8 U F F 0 1 1 1 1 4 11 ' 1 <)42DYfc'I..C)i:J 1 1 1 5 4 4 7 2 1. 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A2- 0. 0.1 o. 30 o 70 0. 33 :i 50 RAN* .1 N E A I L ER8H 1 72 RAWli NH ! 0H RE I N 1 04 RANI! NH i i.;H ER8H 1 i. 2 RANl.,1 N H ! !7H 4QR F 1 37 RftNi.i N H i CH ER8H .1 10 OifAi.i NH fHH 4HRT 1. 1 0 7!: AO NH i CH 8HR f 1 04 o!-" AO NH i CH 4H!-Ei 1. 01 RAi\iO Ni--AH. ER8H 0. 94 RANi i NEA1!.. ADDS. 1. 06 RANf.i N!" A 1 !. R'i i'N . ! . 7 7 RAM..! N! 1 ! <. E, FRSH 2 6 RAW)..* Nil i E.i i ER8H 0. 8 7 RANK Ni- A:I !.. 1 EH-; t 1 . 07 RANi.i NH ! 8H ER8H 0. 9 7 RANK NH ! .El ER8l-i 0. 90 01:.AH Nil ! 7 rl A DDL 1 . 00 BEAU NH H7H R'I 1 N 1 . 08 RANU NH r i.;H E RSH 1 . RANM o'i o.H I-R8H 0. 8 A DK: AD Ni- Al l . RT I N 0. 90 DEAD Nl- A i E R i i'N 0. 9A RANU NI-'A.H. 1 m r 1. 30 ERE. H j . 30 ERE;!-! 1. 2 1 RAND N H ICE; 1.HRI" 1. 1 0 DEAD NEAR. lQR'f 1 07 DEAD Ni-AH. :.-';!.-!!-.: t'' o. 1.0 DEAD Ni ; All.. 4HR'i 0. .i. O RANU Nl- A 11.. 2WRT DEAD Nf AH.. 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H i 23 2087B0W52B1 2) 1 7SSR02A1 2 % 8 7SSR051 2187SSR06i: 223798R08A 1. 2297S8R 10(77 250788R20F1 2627SSR74A1 2A:i77S3l7.<Aiy; 2697SSR26P.1 274 788R29A1 2787SSR29F ;! 2 3 3 7 8 8 R 1 l U i 2587S8R23B1 221 788R0 /(.: i 2297SSR1088 1.7; RAN!.! NP iP 00 RANU iMH i i., 07 RANU NH i C 04 RANU NN (' 8 05 DFAU Nil! C .1 8 BkAU _ NHl'C 08 UP A)I Nidi.;: 10 DFAU Ni-i I C 1.3 BFAU NH IV, A9 RANu Nl-NC 25 RANU NH •'!.: 05 RANU Ni- Al. 93 RANU Ni- A1. -14 RANU NH8 Di-AiJ NHH RANU Ni-i I (.: RANU Nf-I i C X / 1. 1. 15 KANi > Nl- Ai 1 03 RANU Nl-Al 16 RANU Nl-A) 02 RANU NHIC 21 RANU Ni-Al 00 RANU NH |'i.;: HP A* > NH 07 Ui-.A'U NHl'C A. H IHR'f H FRSH H AUDI... H 1 HR" ! H I CR i H 1HRT ! i 4-HR i" :H 4HR"I" H 4HRT H 8w^T if 2HR| I... 3QRT I.. X H R T H 2G,i\'l i i :-:i.-!i-7i R 7 Hi-; I II 2UR1 i. 3HRT i 3QRT i. 3QRI' il 8HR"i i. 4QRT i-l 4 HR l H AU)'M. H 1.URT 

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