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Ecological studies of marten (Martes Americana) in Algonquin Park, Ontario Francis, George Reid 1958

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ECOLOGICAL STUDIES OE MARTEN (MARTES AMERICANA) IN ALGONQUIN PARK ONTARIO by George Reid Francis B.A., University of Toronto, 1956 A Thesis Submitted in Partial Fulfillment of The Requirements for the Degree of Master of Science in the Department of Zoology We accept this thesis as conforming to the required standard The University of British Columbia April, 1958 i A b s t r a c t . L i v e - t r a p s were set f o r max5! n i n a g r i d system covering about f i v e square miles i n Algonc-^a Park, Ontario. Marten were tagged and recaptured during p a r t s o f the summers of 1954 and 1955, and throughout the e n t i r e summers of 1956 and 1957; 452 captures of 55 marten were obtained. Fewer marten were captured i n mid-summer than i n l a t e s p r i n g or l a t e summer. The most complete recapture data I n d i c a t e t h a t s i x minimum for a g i n g ranges of males ranged from 0.50 to 1.05 square miles averaging 0.74 square m i l e s , and were occupied from 8 to 41 days. Males i r r e g u l a r l y s h i f t e d range, overlapping f o r a g i n g ranges of other marten, but moved independently of one another; throughout a summer two males moved over an area of at l e a s t 1.68 and 1.53 square m i l e s , composed of three and f o u r f o r a g i n g ranges r e s p e c t i v e l y . Females occupied d i s c r e t e ranges, perhaps t e r r i t o r i e s , f o r months or maybe yea r s , and t r a v e l l e d them more thoroughly. Four such ranges averaged 0.29 square m i l e s . Two nest dens were found, one among boulders and the other i n a hollow cedar l o g . Immature marten s t a r t e d to disperse through the area i n August. The r e s i d e n t p o p u l a t i o n of marten was probably two per square mi l e (one male and one female) but the t o t a l number of marten found on the given area was f o u r or f i v e per square m i l e ; these l a t t e r were adjacent a d u l t s and d i s p e r s i n g young. Summer l i v e - t r a p p i n g showed no s i g n i f i c a n t d i f f e r e n c e among captures i n d i f f e r e n t f o r e s t types. I n w i n t e r , t r a c k censuses showed t h a t marten p r e f e r r e d c o n i f e r f o r e s t s ! t h i s i i p reference c o i n c i d e d w i t h shallower snow there and the gr e a t e s t occurrence of Clethrionomya as food. Hence, Marten seemed to concentrate t h e i r a c t i v i t y i n c o n i f e r f o r e s t s i n winter and spread through a l l adjacent f o r e s t s i n summer. S h e l t e r from c l i m a t i c extremes appeared to be the most l i k e l y basis f o r h a b i t a t s e l e c t i o n . Analyses of 1427 summer scats and 191 winter and e a r l y s p r i n g scats suggest t h a t food s e l e c t e d depended on i t s a v a i l a b i l i t y . Small mammals formed the major p a r t of the d i e t . Small mammal trapping gave an e v a l u a t i o n of the r e l a t i v e abundance o f d i f f e r e n t s p e c i e s ; although t h i s was not s t r i c t l y comparable to an e v a l u a t i o n by marten p r e d a t i o n , there was no reason to suppose that p a r t i c u l a r species of s m a l l mammals were s p e c i f i c a l l y hunted by marten. The d i e t was h e a v i l y supplemented- w i t h n e s t i n g b i r d s and r i p e b e r r i e s during the seasonal abundance of these; many other items were sampled. The extent of the fo r a g i n g range i n summer appeared to be independent of f o r e s t type and a v a i l a b l e food. In presenting t h i s t h e s i s i n p a r t i a l f u l f i l m e n t of 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 permission f o r extensive copying of t h i s t h e 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 e p r e s e n t a t i v e . I t i s understood tha t copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n permission. Department of Zoology The U n i v e r s i t y of B r i t i s h Columbia, Vancouver B, Canada. Date A p r i l , 1958. i i i TABLE OP CONTENTS PAGE L i s t of t a b l e s v i L i s t of graphs, maps and f i g u r e s v i i i Acknowledgements . . . . . . . . . . x I n t r o d u c t i o n 1 B r i e f H i s t o r y of the Algonquin Marten Study 2 The Study Area 5 A. B r i e f D e s c r i p t i o n 5 B. Mapping of the Forest Cover 6 LIVE-TRAPPING STUDY ; 9 Methods Employed 9 A. Trapping . • • 9 B. Handling and Tagging Marten v,'. . . 11 C. Information Recorded 12 1. Sex and Weight 12 2. Age and Aging . . 12 Assumptions of Method . 15 I n t e r p r e t i n g , Data f o r Range 16 Trapping E f f o r t - E f f e c t i v e Trap Nights' 17 Completeness of Record . . . . . . . 17 A. Percent of Maximum Captures 18 B. C o n t i n u i t y of Record 18 C. D i s t r i b u t i o n of Recapture I n t e r v a l s 21 Resu l t s 2 1 A. Range Size 2 1 B. Movements W i t h i n the Range 2% i v PAGE C. D i s t r i b u t i o n of Captures W i t h i n The Range . . . . 28 D. C i r c u i t s and Scent Posts . . . . . . . . . . . . 28 E. Homing 31 P. T r a v e l l i n g Behaviour 31 G. Den S i t e s 31 H. S p a c i a l R e l a t i o n s Among Male Marten 35 I . Range D i s t r i b u t i o n of Female Marten 37 J . Movements of Male to Female Marten i n Breeding Season 39 K. Di s p e r s a l ' of Immature Marten 41 Populations 42 A. D e n s i t i e s 42 B. Sex Rat i o s 44 Seasonal A c t i v i t y . 45 R e l a t i o n of Marten to Forest Cover 46 A. Winter .- 46 B. Summer 47 FOOD ANALYSES 51 Methods 51 Res u l t s 53 A. Food Items 53 1. Mammals . . . . . . 53 2. B i r d s and B i r d Eggs . 59 3. R e p t i l e s and Amphibians 60 4. Insects and Other Invertebrates . . . . . . . 60 5. B e r r i e s 61 6. Duff Debris 62 V PAGE B. Pood Trends 63 C. Pood Preference of Captive Animals 63 D. R e l a t i o n to A v a i l a b i l i t y of Small Mammal Pood U t i l i z e d 64 Management I m p l i c a t i o n s . . 68 L i t e r a t u r e C i t e d 71 v i LIST OF TABLES TABLE PAGE I Development of the Summer Marten P r o j e c t i n Algonquin Park 3 I I Component Forest Types of the Marten Study Area . 7 I I I T o t a l Animal Captures i n Marten Traps, 1954 to 1957 10 IV Extent of Information on Seventeen Selected Marten 19 V D i s t r i b u t i o n of Recapture I n t e r v a l s f o r Male and Female Marten 21 VI Range Size and Time of Occupancy Arranged by Decreasing C o n t i n u i t y o f Record 22 V I I R e l a t i v e Amount of Movement of Marten Va'ithin Their Range ( D a i l y Movement Index) . 27 V I I I Traps Capturing Marten W i t h i n Their Ranges . . . . 29 IX S p a t i a l R e l a t i o n s Between. Two Male Marten i n Overlapping Ranges 36 X Capture D i s t r i b u t i o n o f Two Males In Overlapping Ranges • 37 XI I n t e r v a l f o r Re-occupation of Vacated Ranges -Adult Female Marten 38 X I I Appearance of New Marten on the Study Area -1956 and 1957 41 X I I I Marten Occurring on 4.5 Square M i l e s of Forest -Summers Only 43 XIV Occurrence of Marten Tracks i n B a s i c Forest Types . 46 v i i TABLE PAGE XV Marten Captures i n Basic F o r e s t Types 48 XVI A n a l y s i s of Marten Scats - Summer, 1955 54 XVII A n a l y s i s o f Marten Scats - Summer, 1956 55 X V I I I A n a l y s i s of Marten Scats - Summer, 1957 56 XIX A n a l y s i s of Marten Scats - Winter and E a r l y Spring 57 XX Analyses of Marten Food - Percent Frequency of Occurrence . . . . . . . . . . . . 67 : v i i i LIST OP GRAPHS, MAPS AND FIGURES Graph 1 V a r i a t i o n s i n Season A c t i v i t y of Marten as I n d i c a t e d by Observed versus Expected Captures. Graph 2 Winter Marten A c t i v i t y Related to Snow Depth i n Major Forest Types. Graphs Seasonal V a r i a t i o n i n Major Pood Types of Marten. 3 - 1 4 Graph 15 Small Mammals Snap-trapped and Small Mammals -Eaten by Marten. Map 1 Marten Study Area and Small Mammal Trap-Lines. Map 2 , Minimum f o r a g i n g range, male No. 55, a d u l t . Map 3 Minimum f o r a g i n g range, male no. 58, a d u l t . Map 4 Minimum fo r a g i n g range, female no. 37, a d u l t . Map 5 Minimum fo r a g i n g range, female no. 41, a d u l t . Map 6 D i s t r i b u t i o n o f male marten showing range o v e r l a p , 1956. Map 7 D i s t r i b u t i o n of male marten showing range o v e r l a p , 1957. Map 8 D i s t r i b u t i o n of female marten - J u l y , August 1955 and May 1956. Map 9 D i s t r i b u t i o n of female marten - June to August 1956. Map 10 D i s t r i b u t i o n of female marten - May to August 1957.. F i g . 1 Winter a e r i a l photograph o f a p o r t i o n of the study area. Co n t r a s t i n g c o n i f e r and hardwood f o r e s t s can be seen; two hemlock r i d g e s show prominently. F i g . 2 Spring a e r i a l photograph of a p o r t i o n of the study area. F i g . 3 D i s t r i b u t i o n of f o r e s t types on the study area. P i g . 4 View over a p o r t i o n of the study area i n the foreground i n d i c a t i n g the r o l l i n g topography of the r e g i o n . i x F i g . 5 Mixed c o n i f e r f o r e s t b ordering O l i v e Lake. P i g . 6 Hemlock f o r e s t - summer. P i g . 7 Hemlock f o r e s t - w i n t e r . F i g . 8 White b i r c h , white spruce and pine f o r e s t - summer. F i g . 9 Sugar maple, yellow b i r c h , hemlock f o r e s t type - w i n t e r . X Acknowledgements; This study was financed by the D i v i s i o n of Research of the Ontario Department of Lands and Forests to whom the w r i t e r i s indebted f o r employment throughout the p e r i o d . The co-op e r a t i o n , i n t e r e s t and suggestions of Mr. R.O. S t a n d f l e l d , D i r e c t o r of the W i l d l i f e Research S t a t i o n i n Algonquin Park, g r e a t l y f a c i l i t a t e d the programme's o p e r a t i o n . The success of the study can l a r g e l y be a t t r i b u t e d to the s u b s t a n t i a l a s s i s t a n c e of Messrs. David Johnston, Douglas B r o d i e , Michael DanieL, Roger H a n s e l l , Ronald Paxton and o t h e r s , a l l of whom at times c o n t r i b u t e d t o approximately 4000 miles of h i k i n g necessary to operate the t r a p l i n e s and c o l l e c t m a t e r i a l f o r the food analyses. A s s i s t a n c e on other aspects o f the work was given on innumerable occasions by Mr. David Bates and was most s i n c e r e l y a p preciated. Acknowledgement i s a l s o due to Mr. Gerard van Tets, various s t a f f members of the Department of P a r a s i t o l o g y , Ontario Research Foundation, and D i v i s i o n of Research, Department of Lands and F o r e s t s , f o r help and suggestions. The w r i t e r a l s o wishes to thank Dr. I . MoT. Cowan, Head Department of Zoology, U n i v e r s i t y of B r i t i s h Columbia f o r advice during the p r e p a r a t i o n o f t h i s r e p o r t . 1 I n t r o d u c t i o n ; In recent decades, a marked decrease i n the economic value of marten i n eastern Canada, co i n c i d e s w i t h a steady d e c l i n e I n numbers and un p r e d i c t a b l e f l u c t u a t i o n s i n f u r demands. Although current Canadian f u r trade problems are p r i m a r i l y economic, the management of a-depleting f u r resource might w i s e l y aim at i n s u r i n g a supply of f u r should vagaries of f u r fashions p r e c i p i t a t e a sudden demand. Pine f u r s p e c i e s , such as marten, r e q u i r e the most c a r e f u l management (Edwards and Cowan, 1957). In O n t a r i o , de Vos (1952) showed the d r a s t i c r e d u c t i o n o f Marten to t h e i r major centres of abundance i n Algonquin Park and the Chapleau Game Reserve. Elsewhere, marten e x i s t e d In r a t h e r i s o l a t e d and w i d e l y s c a t t e r e d groups. Since then, marten have shown some i n d i c a t i o n s of i n c r e a s i n g w h i l e at the same time, the D i v i s i o n of P i s h and W i l d l i f e of the Ontario Department of Lands and Forests has undertaken r e s t o c k i n g programmes. Marten from the Algonquin and Chapleau reserves are r e l e a s e d i n p a r t s of Ontario where the o r i g i n a l p o p u l a t i o n has e i t h e r been exterminated or s e r i o u s l y reduced. A subsequent i n t e r e s t i n management procedures f o r marten demanded i n f o r m a t i o n on the h a b i t s of the species i n Ontario, and was, i n p a r t , r e s p o n s i b l e f o r the present study. Previous f i e l d work has been done on marten* M a r s h a l l (1951b) obtained i n f o r m a t i o n from t r a c k i n g marten i n Boise N a t i o n a l F o r e s t , Idaho. L i v e - t r a p p i n g s t u d i e s i n c l u d e p r e l i m i n a r y ones by de Vos and Guenther (1952) at Chapleau, Ontario and Twin Lakes, Washington; by Lensink (1953) i n 2 i n t e r i o r A l a s k a ; and by M i l l e r e t . a l . (1955) i n Wells Gray Park, B r i t i s h Columbia. Newby and Hawley (1954) and Hawley and Newby (1957) g i v e r e s u l t s of the most Intensive study, from G l a c i e r N a t i o n a l Park, Montana. Lensink e t . a l . (1955) summarized most major food analyses of marten. Except f o r the i n i t i a l Chapleau p r o j e c t , previous work on marten ecology has been done i n montane regions of western North America. Hagmeier's (1955) summary of present marten d i s t r i b u t i o n i n America suggests t h a t the Algonquin p o p u l a t i o n Is one of the l a s t which remains, to some extent, i n the Deciduous Forest Biome. The r e s u l t s of the present study may In d i c a t e h a b i t s o f marten i n former populations of north-eatern North America and provide a b a s i s f o r comparison w i t h areas f u r t h e r west. B r i e f H i s t o r y of the Algonquin Marten Study: Algonquin Park i s a p r o v i n c i a l l y administered area o f about 2700 square miles i n s o u t h - c e n t r a l Ontario. I t was set aside from 1893 to 1914 p r i m a r i l y to p r o t e c t headwaters and t r i b u t a r i e s of four r i v e r s , provide a p u b l i c park, and p r o t e c t w i l d l i f e . I n 1944, a 30 square m i l e Wilderness Area was closed to the p u b l i c and reserved e x c l u s i v e l y f o r w i l d l i f e research. The Ontario Department o f Lands and Forests has si n c e developed the W i l d l i f e Research S t a t i o n , to provide f a c i l i t i e s f o r f i e l d r e s e a r c h , on Lake Sasajewun a t the south-east corner of t h i s Wilderness Area. The present marten study was c a r r i e d on i n f i v e square miles adjacent to Lake Sasajewun. 3 Some marten l i v e - t r a p p i n g was done by s t a f f o f the W i l d l i f e Research S t a t i o n from 1951 to 1953 to o b t a i n p r e l i m i n a r y i n f o r m a t i o n on t r a p p i n g techniques and b a i t preference. The w r i t e r was f i r s t engaged i n l a t e summer, 1954, to continue t h i s . Prom 1955 on, the marten study became a main res e a r c h endeavor and t r a p - l i n e s were expanded to t h e i r f i n a l form by mid-summer 1956. By that time they i n c l u d e d a l l previous areas that had been trapped. The g r i d system of t r a p p i n g then i n operation', enabled range s i z e s o f marten to be determined; a l l previous trapping could at best i n d i c a t e movements by l i n e a r distances between captures. Table I i n d i c a t e s the development of the l i v e - t r a p p i n g programme. Results presented i n t h i s r e p o r t are based on t r a p p i n g i n August 1954, J u l y and August 1955, and the summers (May to August) of 1956 and 1957. TABLE I : Development of the summer marten p r o j e c t i n Algonquin Park l e a r Maximum no. of traps Maximum miles o f t r a p - l i n e Approximate t o t a l t r a p - n i g h t s T o t a l marten captures No. of i n d i v i d u a l marten 1951-53 42 5 6,150 99 31 1954 41 4 1,000 32 14 1955 68 8 3,400 85 18# 1956 142 20 11,250 203 17# 1957 152 20 13,500 139 22# T o t a l 35,300 556 # i n c l u d e s some i n d i v i d u a l s from previous years. 4 In a d d i t i o n to t r a p p i n g , a r a t h e r i n t e n s i v e food study was completed from 1955 to 1957 and a d e s c r i p t i o n of f o r e s t types enabled some assessment of marten to t h e i r h a b i t a t . Supplementary winter observations were made i n 1956 and 1957. 5 The Study Area: A. B r i e f D e s c r i p t i o n * The study area l i e s w i t h i n the Algonquin-Laurentides S e c t i o n (L4) of H a l l i d a y ' a (1937) Great Lakes-St. Lawrence Forest Region. J a r v i s (1956) subdivides H a l l i d a y ' s c l a s s e s f u r t h e r such t h a t the Algonquin area of J a r v i s ' Algonquin-Pontiac Forest S e c t i o n l i e s q u i t e close to the W i l d l i f e Research S t a t i o n . This area i s c h a r a c t e r i z e d by J a r v i s (op. c i t . ) as uplands, 1200 to 1700 f t . above sea l e v e l , w i t h shallow dumped t i l l covering the moderately r o l l i n g bedrock h i l l s of g r a n i t e and g r a n i t e - g n e i s s , and f l u v i a l d e p o s i t s occuning c h i e f l y as narrow deposits between the h i l l s . The climate i s f a v o r a b l e to t o l e r a n t hardwoods but the co l d winters and l a t e s p r i n g f r o s t s r e s t r i c t some s p e c i e s . Sugar maple i s a dominant s p e c i e s , forming pure stands on the h i l l s except on some r i d g e s where hemlock forms pure stands. Other c o n i f e r s , considered e s s e n t i a l l y as i n t r u s i v e s from the Boreal Forest Region ( H a l l i d a y , 1937) occur around lake shores or i n pure or mixed stands on some other s i t e s . White pine at one time occurred throughout the r e g i o n but logg i n g i n the l a t t e r p a r t of the nineteenth century system-a t i c a l l y removed p r a c t i c a l l y a l l stands. Where pine occurred as a climax on sand p l a i n s , the s i t e s now are occupied by various s e r a i stages composed o f white b i r c h , aspen, balsam f i r and white spruce. The a c t u a l marten study area appears l i t t l e a f f e c t e d by t h i s e a r l y c u t t i n g s i n c e i t was f o r the most p a r t q u i t e 6 selective. The area has numerous old tote-roads meandering through i t ; many have been overgrown to some extent. B. Mapping of the Forest Cover. Two sets of aerial photographs were taken on an area of nine square miles centering on the study area. Data pertaining to these photos are as follows: 1) air speed: 100 m.p.h.; 2) altitude: 4500 f t . ; 3) photographic equipment: 3-l/4 x 4-1/-4 Speed Graphic, Kodak Super XX, exposed l/200 sec. at f l 6 ; 4) intervals between pictures: 15 sees. (Enough for an overlap but not enough for stereoscopic interpretation). One set was taken In late winter when the ground was snow covered and hardwood forests contrasted markedly with conifer forests, and the second set was taken i n late spring, after the snow had melted but before buds had burst on the hardwoods; this enabled birches to be distinguished well from other hardwood species. Figures 1 and 2 are sample photographs from these two sets. Forest types were outlined on the photographs and the outlines transferred to a base map with a sketchmaster. The following four major forest types were distinguished, on the basis of c r i t e r i a given by the Canadian Institute of Forestry (1956): Forest type Composition C - conifer up to 25$ hardwoods CH - mixed, predominantly conifer _ 25% to 50% hardwoods HC - mixed, predominantly hardwood 50% to 75% hardwoods H - hardwood 75% to 100$ hardwoods. 7 The four major types were then subdivided on the basis of species composition i n the o v e r s t o r y , i n t o ten component f o r e s t types. I d e n t i f i c a t i o n of ove r s t o r y species could be done l a r g e l y from winter photographs using c r i t e r i a such as those given by Spurr (1948). D i f f i c u l t y was encountered i n d i s t i n g u i s h i n g , w i t h a e r i a l photographs, understory from o v e r s t o r y c o n i f e r In some stands, hence ground checks were made to v e r i f y t h e i r c l a s s i f i c a t i o n . Table I I shows the r e l a t i v e extent of each of these types on the a c t u a l marten study a r e a , w i t h i n the l i m i t s of t h i s area a r b i t r a r i l y s e t . (Map 1 ) . F i g . 3 shows the d i s t r i b u t i o n of the f o r e s t types i n nine square miles c e n t e r i n g on the study a r e a , and Figures 4 to 11 i l l u s t r a t e examples o f some types. TABLE I I : Component f o r e s t types of the marten study area Forest type A r e a , i n acres# Percent of t o t a l area (approx.) CONIFER (C) Hemlo ck White p i n e , r e d pine Black spruce Mixed - Balsam, white spruce, black spruce, white p i n e , hemlock 357 47 22 104 12 2 1 4 MIXED PREDOMINANTLY CONFIER (CH) White spruce, white p i n e , balsam, white b i r c h , aspen Hemlock, sugar maple, yellow b i r c h 93 676 3 23 MIXED PREDOMINANTLY HARDWOOD (HC) White b i r c h , aspen, white spruce, balsam, white pine Sugar maple, yellow b i r c h , hemlock 57 429 2 15 HARDWOOD (H) Sugar maple White b i r c h 1120 24 38 1 $ measured w i t h a dot g r i d superimposed on a map of the f o r e s t cover. 8 The remainder of the r e p o r t i s presented i n two p a r t s , d e a l i n g w i t h the l i v e - t r a p p i n g programme and food s t u d i e s . 9 LIVE-TRAPPING STUDY Methods Employed: A. Trapping. C o l l a p s i b l e wire t r a p s , s i z e s 6" x 6" x 24" and 9" x 9" x 32", produced by the N a t i o n a l L i v e Trap Company, Tomahawk, Wisconsin, were used throughout the study. Both captured marten r e a d i l y and the l a r g e r traps were used to supplement the a v a i l a b l e s m a l l "marten" t r a p s - I n 1956 and 1957, any l a r g e r traps used, were set one i n every four along the l i n e . T r a p - l i n e s were set on a g r i d p a t t e r n as much as topography and p r a c t i c a l c o n s i d e r a t i o n s allowed. The g r i d was constructed from an o r i g i n a l r e c t a n g u l a r l i n e surrounding Kathlyn Lake i n 1953; and was presumed to g i v e a ramdom sample of f o r e s t types; t r a p - s i t e s along these l i n e s were chosen s u b j e c t i v e l y as the best s i t e s - f o r t r a p p i n g marten, and once chosen were r e t a i n e d throughout the course of the study. Map 1. shows t h e i r l o c a t i o n s . A l l but two t r a p s were s e t by f a l l e n l o g s . Each was c a r e f u l l y enclosed w i t h bark, brush and other d u f f d e b r i s . T r a i l s were cut along the t r a p - l i n e s and three d i s t i n c t r o u t e s , e i g h t , seven and f i v e m i l e s long were arranged so th a t a l l were v i s i t e d d a i l y before noon to i n s u r e r e g u l a r r e l e a s e o f marten. F i g s . 12 and 13 show a t y p i c a l t r a p - s i t e and a marten captured i n a l i v e - t r a p . Traps v a r i o u s l y b a i t e d during the f i r s t years of the study (1951 to 1953 and p a r t of 1955) i n d i c a t e d that a v a r i e t y 10 of f i s h , meats, sweets and scented o i l may a t t r a c t marten but meats tended to dessicate, r o t , or a t t r a c t animals other than marten. The greater b a i t effectiveness of kippered herring and other f i s h scent was noted by Newby and Hawley (1954) whereas M i l l e r et. a l . (1955) and Lensink (1953) captured marten successfully with a number of meats and scents. Lensink (1953) states that some trappers use only bright, shiny objects without scent and Loucks (1955) reported a few marten captured i n l i v e -traps "baited" with shiny tags. Raspberry jam and Rexand Poultry O i l - ( f i s h extract) were used as b a i t from 1954 to,.1957; both items were eagerly accepted by captured marten. Jam and f i s h o i l were r e l a t i v e l y inexpensive, easy to obtain, and easy to handle In large quantities. Traps were rebaited at three day i n t e r v a l s and af t e r every disturbance. Table I I I summarizes animal captures from 1954 to 1957 and shows the selectiveness of t h i s trapping technique for marten. TABLE I I I : T o t a l Animal Captures i n Marten Traps, 1954 to 1957 Species Number of 1954 1955 captures 1956 1957 Total Percent of t o t a l Marten 32 78# 203 139 452 71 Red S q u i r r e l 13 1 27 98 139 22 Raccoon 2 2 14 16 34 5 Eastern Chipmunk 1 3 2 6 -Fisher 2 1 2 5 -Snowshoe Hare 3 1 4 -# excluding trapping o f f present study area. Fewer raccoons would have been captured had small traps been used exclusively. Chipmunks were seldom retained i n traps 11 but f r e q u e n t l y sprung them. B. Handling and Tagging Marten. Several methods of handl i n g marten were t r i e d . Animals were e n t i c e d from l i v e -traps Into a wire cone then held f o r examination and tagging; others were anaesthetized i n the cone f i r s t , as suggested by L l e w e l l y n (1953). T r a n q u i l i z e r s were administered to two marten but c o r r e c t dosages and p o s s i b l e harmful s i d e e f f e c t s were too p o o r l y understood, and the treatment too time consuming to adopt. I t was found that one person wearing a stout p a i r of mink handling gloves could grasp a marten, e x t r a c t i t from the trap and h o l d i t while a second person examined and tagged i t . I t Is p o s s i b l e f o r one person to accomplish t h i s alone but the danger o f l o s i n g the animal seemed g r e a t . Hence, i n t h i s study, a l l animals were brought back to the l a b o r a t o r y when f i r s t caught, and a f t e r the r e q u i r e d i n f o r m a t i o n was recorded, r e l e a s e d at the p o i n t o f capture the f o l l o w i n g day. When subsequently recaptured, the animals were r e l e a s e d immediately at the t r a p - s i t e . Of various ear tagsjused, s t y l e 1005, s i z e No. 3, of the N a t i o n a l Band and Tag Company, Newport Kentucky, proved most s a t i s f a c t o r y . These were r e t a i n e d best by marten when placed at the base of the ear r a t h e r than the t i p . Although some tags l a s t e d f o r n e a r l y two years, most were l o s t e v e n t u a l l y ; i t was thought t h a t w i n t e r f r e e z i n g and subsequent decay of t i s s u e immediately i n contact w i t h the tag was r e s p o n s i b l e f o r some of t h i s , others merely p u l l e d through the t h i n ear t i s s u e . 12 From 1954 to 1957, a l l marten were tattooed on one ear. Tattoo model 101, Ketchum Manufacturing Company, Ottawa, On t a r i o , and green tatoo i n k served best. This i n s u r e d p o s i t i v e i d e n t i f i c a t i o n of a l l study animals-when replacement of the e a s i l y read ear tags was necessary. Sketches of t h r o a t patch p a t t e r n were done e a r l y i n the study and l a t e r , photographs of patterns were taken on some anaesthetized marten. Although throat patterns vary w i d e l y , the i d e n t i f i c a t i o n of i n d i v i d u a l s from them so f r e q u e n t l y proved d i f f i c u l t and u n c e r t a i n , t h a t such records were di s c o n t i n u e d soon a f t e r t a t t o o s were a p p l i e d . Marten which died during the study were preserved as museum study s k i n s and deposited at the S o u t h e r n Research S t a t i o n , Maple, O n t a r i o . C. Information Recorded. 1. Sex and weight. Marten are r e a d i l y aexed by n o t i n g the presence of e i t h e r a baculum or a v u l v a , although immature animals sometimes r e q u i r e - c l o s e i n s p e c t i o n . Marten were weighed to the nearest quarter pound when f i r s t captured but t h i s was discontinued i n 1957. Some i n d i v i d u a l s can change weights r a p i d l y (Newby and Hawley, 1954). 2. Age and aging. Hodgson (1956) records a female marten which l i v e d on an A l b e r t a f u r farm f o r s i x t e e n years. Markley and Bassett (1942) c i t e s l o n g e v i t y records up to,eleven years and one r e c o r d of at l e a s t f i f t e e n years f o r a female Martes martes. They f u r t h e r found from observations on a f u r farm, that the youngest marten to produce a l i t t e r was three 1 3 years o l d and the youngest s e x u a l l y mature male to s i r e a f i r s t l i t t e r was a l s o three years o l d j many attempted matings between the f i r s t and second years were noted. Walker (1929) found no evidence of breeding i n two male and two female marten r a i s e d to twenty-seven months, but Hodgson (op. c i t . ) records one female which was s u c c e s s f u l l y bred at about f i f t e e n months. Thus i t appears that marten under three years of age can u s u a l l y be considered immature. Tentative c r i t e r i a have been set up f o r aging marten on s k e l e t a l m a t e r i a l . The s i z e o f the s a g i t t a l c r e s t i s a f u n c t i o n of age and sex ( M a r s h a l l , 1951). M a r s h a l l p o s t u l a t e s that male marten w i t h a c r e s t over 20 mm. l o n g , and females w i t h any c r e s t at a l l were a d u l t s ; males and females w i t h c r e s t s 30 mmi. and 20 mm. r e s p e c t i v e l y were considered to be very o l d . Lensink (1953) set c r i t e r i a which aged male marten up to about f i v e years on height and l e n g t h of c r e s t s and females on e i t h e r minimum s e p a r a t i o n of temporal muscles or subsequent l e n g t h of c r e s t s . He concluded that males up to three years of age have c r e s t s up to 40 mm. long and females up to three years have c r e s t s up to 20 mm. Newby and Hawley (1954), r e f e r r i n g to known age marten, found t h a t " a d u l t males had a pronounced enlargement o f the corpora cavernosa and b a s a l p o r t i o n of the baculum w h i l e j u v e n i l e males lacked t h i s development". Lensink (op. c i t . ) found t h a t marten up to three years of age had a baculum weighing up to 260 mg. and M a r s h a l l (op. c i t . ) concluded that the baculum of immature marten weighed under, and of mature marten over, 100 mg. dry weight. 14 Such d i f f e r e n c e s appear inherent i n d i f f e r e n t p o p u l a t i o n s . Hagmeier (1955) i n h i s taxonomic study on marten s t a t e d ; "One r e s u l t of the present study has been to show tha t marten from d i f f e r e n t l o c a l i t i e s possess bacula of s i g n i f i c a n t l y d i f f e r e n t weights at mat u r i t y and acquire s a g i t t a l c r e s t s of d i f f e r e n t s i z e s a t d i f f e r e n t ages". Any attempt to apply extant c r i t e r i a to the Algonquin, or other Ontario p o p u l a t i o n s , must be done w i t h r e s e r v a t i o n . Aa y e t , no s a t i s f a c t o r y method of aging l i v e marten has been devised. Broad, t e n t a t i v e age c r i t e r i a were adopted f o r the present study and are as f o l l o w s : A d u l t s : - females w i t h a s a g i t t a l c r e s t or w i t h a pronounced l o n g i t u d i n a l depression on top of the head (caused by growth of temporal muscles l a t e r a l l y before the confluence of the temporal r i d g e s to form the c r e s t ) and/or w i t h conspicuous mammae i n d i c a t i n g they had l i t t e r e d . - males with a broad face and conspicuous s a g i t t a l c r e s t ; some l a r g e males w i t h pronounced c r e s t s and n o t i c e a b l y broken and worn t e e t h were designated " o l d a d u l t " . Immatures: - males and females w i t h smooth heads and i n d i s t i n c t i n d i c a t i o n s of the temporal ridges.; mammae of females very inconspicuous; males w i t h narrow face and small t e s t e s . - males w i t h narrow faces and sma l l s a g i t t a l were cl a s s e d as " y e a r l i n g s " , i . e . between j u v e n i l e and a d u l t . - j u v e n i l e marten (young of the year) during the summer had a woolly pelage because of the s p a r c i t y of guard h a i r s , and r e t a i n e d a r a t h e r small k i t - l i k e f a c e . C l e a r l y , these characters are so s u b j e c t i v e t h a t they must be accepted w i t h due r e s e r v a t i o n s . 15 Assumptions of Method. L i v e - t r a p p i n g data can on l y - g i v e the l o c a t i o n o f animals at c e r t a i n times. The i n t e p r e t a t i o n o f such data-to evaluate the s i z e of fo r a g i n g ranges, n e c e s s i t a t e s s e v e r a l assumptions, summarized as f o l l o w s J the animal w i l l be caught i n a l l major pa r t s of i t s range; trapping w i l l not g r e a t l y a l t e r the animal's h a b i t s and p o i n t s of capture do i n f a c t , represent the animal's chosen a c t i v i t y area. Of these, the p o t e n t i a l e f f e c t on the animal's h a b i t s r a i s e s most d i f f i c u l t i e s . Such e f f e c t s may a c t d i r e c t l y on i n d i v i d u a l s , or i n d i r e c t l y through changed p o p u l a t i o n f a c t o r s . Animals may l e a r n to avoid traps a l t o g e t h e r or r e t u r n to c e r t a i n ones f r e q u e n t l y . C h i t t y (1937) concluded that i t was d e s i r a b l e to recapture s m a l l mammals as i n f r e q u e n t l y as necessary to o b t a i n i n f o r m a t i o n . This helped avoid some e f f e c t s o f , a) induced unnatural concentrations of animals w i t h i n an area, b) changes i n p r e d a t i o n r a t e , c) spread;: of i n f e c t i o n by contaminated food and repeated use of t r a p s , d) changes i n feeding h a b i t s , e) r e d u c t i o n of chance matings and f ) death of young through prolonged absence of mother. In four summers, nineteen of f i f t y - f i v e - marten d i e d i n l i v e t r a p s , most f o l l o w i n g repeated captures. Two of these were l a c t a t i n g females and des p i t e i n t e n s i v e searching, n e i t h e r l i t t e r was found. Thus,trapping may have e f f e c t e d the study p o p u l a t i o n by i n c r e a s i n g the m o r t a l i t y r a t e and inducing untimely s h i f t s 'of range. Otherwise there was no evidence t h a t other f a c t o r s given by C h i t t y a p p l i e d e x t e n s i v e l y to marten. 16 In the same four summers, four hundred and f i f t y two captures of these f i f t y - f i v e marten give s u f f i c i e n t evidence that marten do not h a b i t u a l l y avoid t r a p s , although probably some enter more r e a d i l y than o t h e r s . Two a d u l t males were caught f i f t y - s i x and f i f t y - f i v e times respectively;, and a female was caught f o r t y times. Hawley and wewby (1957) caught one male seventy-three times. Analyses of one hundred and s i x t e e n recaptures on successive nights showed that s t r a i g h t l i n e d i s t a n c e s between p o i n t s cf capture ranged from 0 to l - l / 2 miles (p. 27). This suggests that t r a p p i n g d i d not r e s t r i c t the extent of movement, but merely delayed i t . Consequently, i n t h i s study, i t has been assumed that analyses based on the l i v e - t r a p data correspond w e l l w i t h normal marten a c t i v i t y . I n t e r p r e t i n g data f o r Range. Hayne (1949) l i s t e d three ways of i n t e r p r e t i n g such data. 1. The longest distance between recaptures i s taken as the major a x i s of an e l l i p t i c a l range or the diameter of a c i r c u l a r one. This t a c i t l y i m p l i e s a symmetrical range and seems too great an a b s t r a c t i o n based on too l i t t l e data. 2. A c t u a l captures are mapped and a boundary zone i s added on the outer p e r i p h e r y to compensate f o r a l a c k of traps t h e r e . This i n t e r p r e t a t i o n seems r e s t r i c t e d to data obtained from r e g u l a r co-ordinate t r a p p i n g . 3. A range, e n c l o s i n g an area i n which the animal d e f i n i t e l y was known to occur, i s o u t l i n e d by j o i n i n g outer p o i n t s of capture. 17 Burt (1943) p o i n t s out t h a t ranges may be amoeboid _in o u t l i n e and d e l i n e a t i o n of o u t l y i n g points may accentuate the amount of o v e r l a p w i t h adjacent ranges and give f a l s e impressions o f s i z e . Another somewhat analogous argument s t a t e s that i t i s extremely u n l i k e l y t h a t any range co i n c i d e s w i t h the t r a p - l i n e p a t t e r n , so r e a l range must extend beyond the outer p o i n t s of capture (Hayne, op. c i t ) . However, t h i s "minimum range" i n t e r p r e t a t i o n has been adopted by the w r i t e r because i t seemed most conservative and o b j e c t i v e . Trapping E f f o r t - E f f e c t i v e Trap Nights. For comparative purposes, trapping r e s u l t s were evaluated on a per t r a p - n i g h t b a s i s . One t r a p - n i g h t i s defined as one t r a p set f o r one n i g h t , or more p r e c i s e l y , the i n t e r v a l between successive d a i l y v i s i t s to the t r a p . Traps were f r e q u e n t l y sprung by various s m a l l animals and, at times, many were t o r n out of t h e i r sets by bears. Since i t i s never determined e x a c t l y when such traps were d i s t u r b e d , i t was assumed that each was e f f e c t i v e f o r capturing marten f o r one h a l f of the t r a p - n i g h t i n t e r v a l . Hence, the term " e f f e c t i v e t r a p - n i g h t " (ETN) i s used, and i s defined as the t o t a l t r a p -nights minus one h a l f of the number of disturbances. Completeness of Record. The records of 10 male and 7 female marten have been s e l e c t e d f o r i n f o r m a t i o n on range s i z e . An e v a l u a t i o n of the completeness of these records seemed necessary and two a r b i t r a r y standards were set up. 18 A. Percent of Maximum Captures. One capture per day i s d e f i n e d as the maximum number f o r a given animal. The t o t a l number of captures of a given animal i s expressed as a percentage of the t o t a l number of days i t s area was trapped. B. C o n t i n u i t y of Record. One capture per week i s def i n e d as the minimum requirement f o r a continuous r e c o r d of a given animal's whereabouts. The number of a l l recaptures o f an animal i n i n t e r v a l s of seven days or l e s s i s expressed as a percent o f a l l captures of the same i n d i v i d u a l . A record c o n t i n u i t y of 100$ i n d i c a t e s that a l l captures of the animal occurred w i t h i n t e r v a l s of a week or l e s s , although a t the extreme, t h i s need o n l y be 14$ (one-seventh) of the maximum p o s s i b l e captures. Thus, a hig h r a t i n g f o r these two evaluations i n d i c a t e s a continuous record throughout the t r a p p i n g season. High c o n t i n u i t y and low percent of maximum captures i n d i c a t e s a continuous r e c o r d f o r o n l y a p a r t of the season; low c o n t i n u i t y and consequent low percent of maximum i n d i c a t e s an incomplete r e c o r d throughout the t r a p p i n g season. Again, a r b i t r a r i l y , h i g h c o n t i n u i t y o f r e c o r d i s defined as 75$ or over, whereas h i g h percent of maximum need be 14$ or over. 19 Table IV summarizes recapture i n f o r m a t i o n of seventeen s e l e c t e d marten. TABLE IV: Recapture Information on Seventeen Selected Marten MALES: Age T o t a l T o t a l days area Percent of Record captures was trapped maximum c o n t i n u i t y No. 40 ad. 1956 23 106 22 86$ 1957 23 106 22 82$ No. 53 ad. 1955 14 44 28 85$ No. 55 o l d ad. 1955 10 39 20 89$ 1956 106 Range 1 17 38 100$ Range 2 18 43 Range 3 10 45 100$ No. 56 juv. 1955 10 39 20 89$ No. 58 ad. 1955 ' 5 41 5 56$ 1956 106 Range 1 12 27 83$ Range 2 13 42 92$ Range 3 8 42 100$ Range 4 7 64 100$ 1957 10 106 9 30$ No. 67 o l d ad. 1956 15 70 19 1957 9 103 9 71$ No. 76 ad. 1957 '7 105 7 100$ No. 86 juv. 1957 5 103 6 100?' 'o No. 88 y e a r l i n g 1957 9 101 9 88$ No. 91 y e a r l i n g 1957 6 103 6 100$ 20 TABLE IV: continued FEMALES: Age T o t a l T o t a l days area Percent of Record captures was trapped maximum c o n t i n u i t y No. 37 1954 1956 1957 No. 41 1954 1955 1956 3 y r s . i n 1957 ad. 1 25 14 1 12 3 17 106 106 14 21 8 23 13 25 54$ No. 64 1955 1956 ad. 3 2 19 7 No. 65 1956 1957 No. 66 1956 1957 No. 68 1956 1957 No. 71 1956 ad. ad. ad. 2 6 16 10 17 11 2 yrs . ? 106 32 106 105 77 103 76 17 15 9 22 11 100$ 78$ 21 G- D i s t r i b u t i o n o f Recapture I n t e r v a l s . Table V summarizes marten recaptures a f t e r varying i n t e r v a l s . I t shows t h a t most occur w i t h i n a p e r i o d of one week. Of these, almost 40$ occur on successive days, and about 80$ of male, and 70$ of female recaptures occur i n i n t e r v a l s up to three days. TABLE V: D i s t r i b u t i o n o f Recapture I n t e r v a l s f o r Male and Female Marten I n t e r v a l s i n Days Frequency o f Recaptures, males Percent of Recaptures W i t h i n One Week a l l Frequency of Recaptures, FemaDas Percent of a l l Recaptures W i t h i n One Week 1 74 39 33 36 2 52 28 18 19 3 27 14 13 14 4 18 10 12 13 5 9 5 8 9 6 5 3 6 7 7 3 .1 2 2 R e s u l t s : A. Range s i z e . The f o r a g i n g range of marten, h e r e a f t e r r e f e r r e d to as range, i s defined i n t h i s study to be any area i n which a marten concentrates I t s a c t i v i t y f o r a week or more. This i s d i s t i n g u i s h e d from a summer range, and an annual range, which may c o n s i s t of a s e r i e s of fo r a g i n g ranges occupied a t d i f f e r e n t i n t e r v a l s . Maps 2 to 10 show the s i z e of f o r a g i n g ranges of marten, occupied during the study. The area of each was determined w i t h 22 a dot g r i d superimposed on the range maps. S i z e and l e n g t h o f occupancy, o f a gi v e n range v a r i e s w i d e l y w i t h male marten. Table VI gives such records f o r s i x ranges determined w i t h captures of 100$ c o n t i n u i t y , and nine others w i t h h i g h c o n t i n u i t y . TABLE V I : Foraging Range Size and Time o f Occupany Arranged by Decreasing C o n t i n u i t y of Record  Male Record T o t a l Minimum Dates o f ! Minimum No. C o n t i n u i t y number len g t h of occupancy range of occupancy s i z e , captures (days) sq.miles 55 100% 17 41 May 18-June 28/56 0.64 55 100$ 10 12 Aug. 11-23/56 1.05 58 100$ 8 18 Aug. 1-19/56 0.89 58 100$ 7 10 Aug. 20-30/56 0.58 91 100$ 6 9 Aug. 10-19/57 0,81 86 100$ 5 8 Aug. 1-9/57 0.50 o Average 0.74 55 94$ 18 34 J u l y 7-Aug.10/56 0.84 58 92$ 13 23 J u l y 5-28/56 0.57 55 89$ 10 42 J u l y 15-Aug.26/55 0.22 56 89$ 10 36 J u l y 20-Aug.25/55 0.17 88 88$ 9 24 Aug. 5-29/57 0.30 40 86$ 23 68 June 24-Aug.31/56 0.70 53 85$ 14 42 J u l y 14-Aug.25/55 0.16 58 83$ 12 40 May 18-June 27/56 0.36 40 82$ 23 85 May 20-Aug.13/57 0.41 The f i r s t s i x minimum ranges average 0.74 square miles w i t h e xtremes of 0.50 to 1.05 square m i l e s , f o r periods ranging from e i g h t to forty-one days, of the l e s s complete r e c o r d s , 23 three i n d i c a t e range s i z e s w i t h i n these l i m i t s . The remainder had ranges below one h a l f square mi l e w i t h i n the t r a p l i n e s , and the s l i g h t d i s c o n t i n u i t y of record may i n d i c a t e they ranged beyond. Males o f t e n change range at short and i r r e g u l a r i n t e r v a l s . Maps 2 and 3 show that t h i s s h i f t more f r e q u e n t l y i s a gradual moving away from a previous range r a t h e r than an abrupt change from one to the other. The dates o f s h i f t i n these instances were recorded a r b i t r a r i l y a t a time when the animal was f i r s t captured w e l l outside of i t s previous known range. The same male w i l l f r e q u e n t l y occupy an o l d range at a l a t e r date but the r e t u r n to former areas does not occur w i t h r e g u l a r p e r i o d i c i t y . Male No. 55 occupied a d i f f e r e n t range i n August,1956 from that of August 1955, although there was some ove r l a p . Male No. 40 occupied a range i n J u l y and August 1956 and from May to J u l y 1957 which overlapped a p o r t i o n of the range I t occupied In l a t e summer, 1954; i t was absent from the re g i o n throughout the l a t t e r p a r t of the summer of 1955. Male No. 58 occupied a range i n the f i r s t of August 1956, part of which was subsequently used i n August 1957, wihereas i t s range during the l a t t e r p a r t of August 1956 overlapped p a r t of i t s known range of August 1955. Male No. 38 occupied a range during the summer of 1955 which was two miles from p a r t o f i t s range used i n August 1954. Male No. . 6 7 occupied a range i n June and August 1958 from which i t appeared absent i n J u l y , and i n 1957 i t was absent from the same range from mid-June to mid-August. An a c c i d e n t a l capture of t h i s i n d i v i d u a l was made one mile from the outer edge o f the study area soon a f t e r i t l e f t i n June. 24 Thus, the f o r a g i n g range of a male marten at any given time i s only a p a r t o f the e n t i r e summer range which may be only a p a r t o f i t s annual range, and t h i s too may vary y e a r l y . The t o t a l minimum summer range o f male No. 55 i n 1956 was 1.68 square miles composed of three c o n s t i t u e n t ranges averaging 0.84 square m i l e s , and of male No. 58, 1.53 square miles o f four c o n s t i t u e n t ranges averaging 0.60 square m i l e s . By c o n t r a s t , female marten are qu i t e sedentary. Complete c o n t i n u i t y o f record was d i f f i c u l t to o b t a i n perhaps because of a d i f f e r e n t i a l t r a p a b i l i t y between males and females. S i x r e s i d e n t female marten were caught an average of nineteen times each, whereas f o u r r e s i d e n t males averaged f o r t y - s i x captures each. Maps 4 and 5 and 8 to 10 show ranges of female marten. Female No. 41 d i e d i n May 1956 i n the range where i t was f i r s t found i n August 1954. Complete c o n t i n u i t y of re c o r d was obtained f o r a l i m i t e d t r a p p i n g p e r i o d w i t h i n i t s range i n August 1955 and i n May 1956 u n t i l i t d i e d ; t h i s e s t a b l i s h e d a minimum range s i z e o f 0.24 square m i l e s . Female No. 66 occupied a minimum range of o n l y 0.09 square miles f o r f i f t y -one days i n l a t e summer 1956 and was present w i t h i n the same area at varying times i n 1957. Female No. 37 was captured as a j u v e n i l e i n August 1954 at the peri p h e r y of an area subsequently found, by more extensive trapping-,--to be i t s range i n 1956 and 1957; p o s s i b l y i t had l i v e d i n the same area f o r four summers. The s i z e o f i t s range throughout the summer of 1956 was 0.54 square m i l e s , and at l e a s t 0.41 square miles during the summer 25 of 1957 although r e c o r d c o n t i n u i t y was low (54$). Map 4 shows that the l o c a t i o n of t h i s marten's range had s h i f t e d s l i g h t l y between 1956 and 1957. Female No. 68 occupied a range of 0.29 square miles f o r f i f t y - o n e days i n 1956 and was captured throughout the same area i n 1957. The average o f these four determinations o f range s i z e i s 0.29 square m i l e s . Female No. 64 d i e d i n May 1956 i n a range where i t was f i r s t found i n August 1955, and No. 65 died i n June 1957 where i t was f i r s t found i n 1956. Hence, evidence suggests t h a t females occupy a range f o r a r e l a t i v e l y long time. I t i s much smaller than the areas ranged by males i n equal times, although ranges occupied by males f o r o n l y a few days or weeks may be no l a r g e r than a female's range over the e n t i r e summer. Only Hawley and Newby (1957) obtained l i v e - t r a p data on ranges of marten. In Montana, they found that ranges o f s i x male marten averaged 0.92 square miles (extremes of 0.34 to 1.68) over periods ranging up to 788 days and ranges of f i v e female marten averaged 0.27 square miles (extremes o f 0.03 to 0.70) over periods ranging up to 660 days. These data agree remarkably w e l l w i t h those presented above and i n d i c a t e consistency i n h a b i t s of marten i n w i d e l y v a r y i n g h a b i t a t s . M a r s h a l l (1951b) estimated from winter t r a c k evidence that one marten covers from one h a l f to two square miles d a i l y i n Idaho. I n an e n t i r e winter season one marten was considered to move over a ten to f i f t e e n mile square area, covering c e r t a i n p o r t i o n s i n t e n s i v e l y n i g h t by n i g h t and r e t u r n i n g t o favoured 26 spots f o r hunting at i n t e r v a l s of two to three weeks, de VSos and Guenther (1952) c i t e a previous statement of Marshall's that "the p a t t e r n o f movements i s an adjacent or overlapping s e r i e s ; o f s m a l l areas that change more or l e s s each day. In a complete c y c l e the animal tends to r e t u r n to- i d e n t i c a l favourable hunting spots but the "tracks by no means always f o l l o w those of, -previous t r i p s " . D u l k e i t , as c i t e d by Lensink (1953) found that the e n t i r e winter range of s a b l e , Martes z i b e l l i n a , on Great Shantar I s l e , U.S.S.R. g e n e r a l l y ranged from 1.9- to 7.7 square miles with- exceptions up to 11.5 square m i l e s . This range however, was made up o f hunting sections averaging 0.6 square miles i n s i z e . One male spent t h i r t y - s e v e n days during January and February i n an area of o n l y 0.1 square kilometers and one other male i n 0.3 to 0.4 square miles from November 5th through to March. Seven other sable remained i n hunting s e c t i o n s varying from 0.05 to 1.0 square miles f o r at l e a s t a month each. A female l i v e d i n 1.5 to 1.9 square miles from October 25th to February 18th. This i n f o r m a t i o n suggests that the ranging h a b i t s as noted f o r Algonquin marten i n summer, f o l l o w the same p a t t e r n I n d i c a t e d f o r the species the year round i n western montane areas and corresponds remarkably w e l l to h a b i t s o f Martes z i b e l l i n a . This l a t t e r i s a l l the more i n t e r e s t i n g i n the l i g h t of Hagmeier's (1955) contention that M. z i b e l l i n a and M. americana are c o n s p e c i f i c . D i f f e r e n c e s i n magnitude of areas recorded may, to some extent, be due to d i f f e r e n t methods of o b t a i n i n g the i n f o r m a t i o n . 27 B. Movements w i t h i n the range* S t r a i g h t l i n e d i s t a n c e s between captures on successive days were assumed to show the r e l a t i v e amount of movement of marten w i t h i n t h e i r range. Such distances were measured on a map and c a l l e d " D a i l y Movement Index". TABLE V I I summarizes t h i s i n f o r m a t i o n . Adults moved twice as f a r as immatures i n a given time perhaps because immature marten do not always s e l e c t d e f i n i t e ranges (p. 41) and females moved about two-thirds (63$) the distance that males d i d i n equal i n t e r v a l s . TABLE V I I : R e l a t i v e Amount of Movement of Marten W i t h i n Their Range ( D a i l y Movement Index) Adult Immature A l l Adult Immature A l l males males - males females females females No. of observations 58 16 74 34 8 42 Average and 0.60 0.30 0.54 0.38 0.14 0.34 extreme m i l e s . ' (0-1.4) (0-1.2) (0-1.0) (0-0.4)  A l l a d u l t s : 92 observations, averaging 0.52 m i l e s . A l l immatures: 23 observations averaging 0.25 m i l e s . Hawley and iNfewby (1957) obtained f o r t y captures on successive days throughout the year on one ad u l t male marten, and found a range from 0 to 1.6 miles w i t h an average of 0.44; t h i s c o i n c i d e s w e l l w i t h the above data. However, a r e p r e s e n t a t i o n of data obtained by M i l l e r e t . a l . (1955) gives the average of seven successive recaptures of male marten i n B r i t i s h Columbia as one m i l e , ranging from 0 to 4.5 m i l e s , and one s i m i l a r recapture o f a female a t 1.25 m i l e s . 28 C l e a r l y , the above data depend l a r g e l y on t r a p spacing, e s p e c i a l l y to o b t a i n the gr e a t e r distance r e c o r d s , but M i l l e r e t . a l . (op. c i t . ) mention the other p o s s i b i l i t y that such d i f f e r e n c e s are r e a l , contingent on h a b i t a t or seasonal f a c t o r s . Such c o n s i d e r a t i o n s seem however, not to i n v a l i d a t e the co n c l u s i o n t h a t male marten u s u a l l y t r a v e l f u r t h e r than females at any given time. C. D i s t r i b u t i o n of captures w i t h i n rang^e. For marten whose h i g h c o n t i n u i t y of record enabled a determination of range, the number of traps i n which each was caught was compared as a percent of the number of traps a v a i l a b l e w i t h i n the range. This e v a l u a t i o n was assumed to i n d i c a t e the extent that each animal covered i t s range. TABLE V I I I gives t h i s i n f o r m a t i o n , and shows t h a t although the number of captures of males averaged o n l y s l i g h t l y fewer than females, ,male:marten were captured i n about one t h i r d of the a v a i l a b l e traps and females i n about one h a l f . This may suggest t h a t females more thoroughly cover the range they s e l e c t . D. C i r c u i t s and scent p o s t s . The range maps show that marten were captured one or more times i n some traps and not a t a l l i n o t h e r s . The consistency of the trapping method makes i t reasonable to assume t h a t a l l traps are e q u a l l y e f f e c t i v e f o r catching marten, thus, the capture d i s t r i b u t i o n may i n d i c a t e some r e s t r i c t i o n i n marten movements w i t h i n a given range. This has been e x e m p l i f i e d i n TABLE X comparing capture d i s t r i b u t i o n of two males i n overlapping ranges (p. 3 7 ) . 29 " D a i l y Movement Index" determinations show tha t marten can e a s i l y t r a v e r s e t h e i r e n t i r e range i n one day so l i v e - t r a p p i n g TABLE V I I I : Traps Capturing Marten W i t h i n Their Ranges Record No. o i % of Record No.of % of c o n t i n u i t y captures traps con- cap- traps i o b t a i n i n g t i n u i t y tures o b t a i n i n g captures captures Males: Males: No. 55 89 10 46 No. 40 86 23 33 100 17 39 82 23 41 94 18 34 No. 53 85 14 61 100 10 18 No. 56 89 10 61 No. 58 83 12 42 No. 86 100 5 22 92 13 30 No. 91 100 6 17 100 8 14 No. 88 88 9 39 100 7 25 No. 67 78 15 47 Average • • 13 35 Females : 100 Females • • No. 41 15 62 No. 68 88 17 43 No. 66 100 16 60 No. 37 90 25 50 Average • * 18 54 r e s u l t s cannot be r e l i e d on to give i n f o r m a t i o n on c i r c u i t s . Female No. 66 was caught nine times i n a t r a p beside a l o g i n a s m a l l spruce bog. This may i n d i c a t e a more p r e c i s e route s e l e c t e d through wet areas. Marshall's (1951b) t r a c k i n g s t u d i e s showed that the d a i l y movement of one marten ranged from an estimated two to nine m i l e s . W i l l i a m s (1947) sta t e s t h a t marten "have r e g u l a r runs that take i n a twenty to twenty-five mile c i r c u i t which they cover p e r i o d i c a l l y " , and Remington (1950), r e f e r r i n g to marten i n Colorado, s t a t e d that the d a i l y range i s approximately one to four m i l e s . Schmidt (1943) c i t e s observations on three species of European marten which note re g u l a r c i r c u i t routes w i t h i n 30 e s t a b l i s h e d hunting ranges. Martes f o i n a i s a l l e g e d to have more r e s t r i c t e d routes than M. martes or M.~ z i h e l l i n a . Routes are marked w i t h scent posts t h a t are reputed to be r e g u l a r l y r e - v i s i t e d . D e p o s i t i n g scent appears to be an innate a c t i o n i n which the abdominal gland i s r a p i d l y rubbed on any sm a l l p r o j e c t i n g s u r f a c e , p a r t i c u l a r l y stones or s t i c k s . Observations are c i t e d on eighteen c a p t i v e marten and sable l i v i n g together, each of which r e t a i n e d i t s own scent p o s t s , r e g u l a r l y r e - v i s i t e d . Marten r e a d i l y rescent " f o r e i g n " scent posts of other I n d i v i d u a l s i n t h e i r range. Rate of d e p o s i t i n g scent was highest d u r i n g the r u t and lowest i n mid-winter. The motions of scent d e p o s i t i o n s t a r t i n young at the age of three months, but i t i s l a t e r before f u n c t i o n a l scent p i l e s are e s t a b l i s h e d . L i t t l e i s extant i n North American l i t e r a t u r e on marten scents. H a l l (1926) describes i n d e t a i l the prominent abdominal gland, and draws a t t e n t i o n to the f a c t that i t s occurrence had been p r e v i o u s l y overlooked. Both H a l l ( o p . c i t . ) and Markley and Bassett (1942) re p o r t observations on captives rubbing t h e i r b e l l i e s across p r o j e c t i n g limbs and other o b j e c t s . The same has been observed by the w r i t e r i n c a p t i v e marten. Remington (1950) c i t e s from track evidence i n Colorado, a s m a l l spruce protruding s i x to e i g h t inches above the snow t h a t was used as a u r i n a t i n g or "scent post" by one marten and i n v e s t i g a t e d l a t e r by another which deviated from a l i n e o f t r a v e l about twenty f e e t to i n v e s t i g a t e i t . Thomas(1952) opines that scent e s t a b l i s h e s " i n v i s i b l e t r a i l s " which are "followed by many martens i n the area". 31 E. Homing> On the b a s i s of work a t Chapleau, Ontario, de Vos and Guenther (1952) f i n d "...some I n d i c a t i o n that animals t r a n s p l a n t e d over a distance v a r y i n g from one to twelve m i l e s tend to r e t u r n to the place where they were f i r s t captured." Male No. 33 escaped at Lake Sasajewun on August 26, 1954 and was recovered w i t h i n i t s range at K a t h l y n Lake the f o l l o w i n g morning at a s t r a i g h t l i n e distance of one m i l e . Marten No. 51 escaped from the same place on May 24, 1955 and was recovered two and one h a l f miles away three days l a t e r beside i t s o r i g i n a l p o i n t of capture. Although not tagged, I d e n t i f i c a t i o n was rendered q u i t e c e r t a i n by a d i s t i n c t b r i g h t pelage and moult p a t t e r n . C h i t t y (1937), commenting on repo r t e d homing i n Peromy3cus, notes the p o s s i b i l i t y t h a t r e t u r n s from shorter distances may be explained by the mice having l a r g e r home ranges than expected; the same p o s s i b i l i t y may apply to marten. P. T r a v e l l i n g Behaviour. Numerous short observations on marten a f t e r r e l e a s e from t r a p s , and winter t r a c k s t u d i e s , r e v e a l some methods of hunting and t r a v e l l i n g commonly used by marten. , The number of marten seen i n , or c l i m b i n g , trees was reporded f o r three summers. Of f o u r hundred and sixty-one marten o b s e r v a t i o n s , most of the animals j u s t r e l e a s e d from t r a p s , o n l y nine concerned marten i n t r e e s . In s p i t e of a greater p r o b a b i l i t y of observing marten on the ground than i n t r e e s , t h i s s t i l l i n d i c a t e s t h a t most time i s spent on the 31 ground. Several o f the nine observations were of marten f r i g h t e n e d i n t o trees from which they soon descended. Winter t r a c k i n g i n d i c a t e d frequent climbing through or along d e a d f a l l s i n c o n i f e r f o r e s t s but there was no c l e a r evidence of extended movement through t r e e s . One marten tracked f o r over two miles climbed trees f o u r times; i n three cases i t crossed to a second tree w i t h i n ten f e e t of the f i r s t then jumped i n t o the snow, i n one instance probably from a height of about f i f t e e n f e e t . This same marten jumped about f i f t e e n f e e t over a small ledge i n t o s o f t snow below. No other marten which were followed showed t h i s behavious but some climbed around the base of l a r g e t r e e s and leaped forward two or three f e e t i n t o the snow. One marten a c t u a l l y floundered through deep snow r a t h e r than t r a v e l through t r e e s i n hardwoods. Tracks have i n d i c a t e d that marten, when t r a v e l l i n g through s o f t snow over a c r u s t , w i l l sometimes s l i d e down slopes on t h e i r b e l l y . Repeated observations of marten i n summer show that f a l l e n logs are used f o r runways and a route i s a c t u a l l y s e l e c t e d through the f o r e s t which allows f u l l e s t use of l o g s . In w i n t e r , marten seemed o n l y to r u n down l a r g e logs i f the l o g l a y v i r t u a l l y p a r a l l e l to the d i r e c t i o n they were going. They d i d not detour f a r to get to one, hence d i d not show the same s e l e c t i o n as i n summer. P o s s i b l y , snow cover makes t r a v e l e a s i e r than i t i s through tangled summer growth. Observations and t r a c k s i n d i c a t e that marten have three d i s t i n c t g a i t s . These can be described as: Bounding, most 32 f r e q u e n t l y used i n loose s o f t snow or through denser ground v e g e t a t i o n ; G a l l o p i n g , while t r a v e l l i n g on crust covered snow or open ground; and Walking, when movement becomes d i f f i c u l t or something i s being i n v e s t i g a t e d . Hunting appears to be of a haphazard nature. Movement, though tending i n a g i v e n d i r e c t i o n i s q u i t e e r r a t i c , w i t h many t w i s t s and t u r n s , most f o r no apparent reason. I n w i n t e r , no tendency was noted f o r marten to f o l l o w deer t r a i l s , snowshoe t r a i l s or other marten tr a c k s f o r more than a short d i s t a n c e . Remington (1950) tracked eleven marten i n Colorado and found that a l l t r a v e l l e d s i n g l y , seldom climbed a t r e e , and when they d i d , u s u a l l y jumped back out of i t . He notes that "marten thoroughly i n v e s t i g a t e downed l o g s , stumps, brush p i l e s and holes i n or under rocks when hunting,...also d i g to the ground l e v e l and below i n deep snow; i f w i n d f a l l s a l l o w , w i l l t r a v e l as much as f i f t y yards under these l o g s " . He c i t e d two instances of marten having dug b i t s of h a i r from holes s i x to t h i r t y inches deep i n snow. Ma r s h a l l (1951b) i n Idaho, spent nine days t r a c k i n g two marten oyer an estimated seventeen and one h a l f m i l e s , d u r i n g which n e i t h e r animal climbed a t r e e . A t h i r d marten, tracked f o r twenty-one miles i n s i x days, climbed trees only e i g h t times; each t r e e had a dense crown and u s u a l l y "witches brooms". Marten u s u a l l y jumped out of these t r e e s , l a n d i n g t h i r t y to f o r t y f e e t from the trunk. He noted that marten hunt from one snag, w i n d f a l l or dense tangled clump of low brush to another, o f t e n t r y i n g to approach such f e a t u r e s by 33 long leaps from the u p h i l l s i d e . This was o s t e n s i b l y to s u r p r i s e prey s p e c i e s . x f i t f a i l e d , then the animal searched every hole or c r e v i c e . He f u r t h e r s t a t e s , that of t r a c k s f o l l o w e d , one hundred and f o r t y - s i x were of a s i n g l e animal, twenty-nine were of two, f i v e o f three, two of four and one of s i x animals probably t r a v e l l i n g together. Confluence o f marten tracks was o f t e n noted i n Algonquin but no way was found of determining which, i f any, were made simultaneously. de Vos (1951) remarks t h a t a marten tunnels q u i t e f r e q u e n t l y under the snow, w h i l e a f i s h e r does so r a r e l y i n Ontario, de Vos and Guenther (1952) b e l i e v e marten r e s t r i c t t h e i r movements to the v i c i n i t y of main watersheds. The topography of Algonquin i s such that marten a r e never f a r from water. Popular and semi-popular w r i t i n g s (e.g. Cross and Dymond, 1929; B u r t , 1946; H i g g i n s , 1948) commonly s t r e s s predominating a r b o r e a l h a b i t s of marten. The most extreme of such p r e s e n t a t i o n s , convey impressions of marten r a c i n g through t r e e t o p s , eagerly k i l l i n g s q u i r r e l s , and denning e x c l u s i v e l y i n hollow t r e e s . The European pine marten (Martes  martes), i s c h i e f l y a r b o r e a l (Schmidt, 1943). I t would be of I n t e r e s t , although d i f f i c u l t , to a s c e r t a i n the extent to which h a b i t s of t h i s species were u n c r i t i c a l l y a s cribed t o Martes  americana because of the s u p e r f i c i a l morphological s i m i l a r i t i e s of the two. 34 G. Den S i t 6 3 . Two nest dens were found on the Algonquin study area by Mr. David Johnston. A l i v e - t r a p p e d k i t marten was followed to one a f t e r i t s r e l e a s e on J u l y 28, 1955. The entrance was among bou l d e r s , beside a s m a l l spring i n c o n i f e r f o r e s t , w i t h i n a hundred yards of Kathlyn Lake; numerous scats were by the entrance. The second, was found on June 8, 1957 a f t e r a marten was seen running i n t o a hollow l o g and a female, observed to have l a c t a t e d , was f r i g h t e n e d from the l o g i n t o a l i v e - t r a p . Three young were subsequently e x t r a c t e d and on the b a s i s of c r i t e r i a given by Brassard and Bernard (1939), were estimated to be e i g h t weeks of age; they were r a i s e d i n c a p t i v i t y . The l o g was a f a l l e n cedar on top of a hemlock r i d g e (near Jack Lake) and the p o r t i o n used as a nest was approximately twenty f e e t from the entrance In a s e c t i o n of about ten inches diameter, w i t h a r o t t e n centre four inches in.diameter, f u l l of r o t t e d wood powder and f l a k e s . -Some knotholes opened from the s i d e . A small p i l e of scats was a t the den entrance and others were on a nearby .log l e a d i n g away from the den l o g . In the nest m a t e r i a l i t s e l f , t a i l s of one Peromyscus, one Tamias and two Tamiasciurus and the l e g s , f e e t and f e a t h e r s o f a b i r d , probably a t h r u s h , were found. The adult female and a l l three young had numerous t i c k s (Ixodes ?) i n t h e i r ears. Figures 14 and 15 i l l u s t r a t e t h i s l a t t e r den s i t e and a winter day den. Four winter day den s i t e s were found. A l l were underground with an entrance o n l y l a r g e enough f o r a marten to 35 enter but these openings could be enlarged by digging the snow away. Two e x i s t e d under l a r g e boulders, one under a hummock formed by t r e e roots and one under a f a l l e n l o g completely covered w i t h snow. In summer, s e v e r a l marten re l e a s e d from traps were seen e n t e r i n g hollow logs which may also serve as dens f o r short p e r i o d s . Winter evidence showed that one den was used on two consecutive days during a s i x t e e n day p e r i o d i n January ( P i g . 15)j others appeared to be used f o r only one day at a time. Quick (1955) found one den s i t e i n northern B r i t i s h Columbia which was used by a marten f o r about a month i n e a r l y w i n t e r . No evidence was obtained i n Algonquin of marten spending time i n hollow t r e e s . In Colorado, Remington (1952) found a nest den i n a rock p i l e i n a moist open draw next to a w i l l o w t h i c k e t , and noted remains of a hare and a woodpecker at the s i t e , i t was close to a highway and p u b l i c camp ground. M a r s h a l l (1951b) found t h i r t e e n w i n t e r den s i t e s i n downed logs and three i n holes i n stumps, and Remington (195C) r e p o r t s winter dens underground i n o l d stumps or under r o c k s j some evidence was obtained that marten may use t h i c k l y branched trees and those w i t h hollows i n them f o r r e s t i n g s i t e s . H. S p a c i a l R e l a t i o n s Among Male Marten. Maps 6 and 7 g i v e a l l occurrences of male marten on the study area f o r the summers of 1956 and 1957. I t i s r e a d i l y seen t h a t considerable overlapping of areas occurs i n the i n t e r v a l of an e n t i r e summer. However the p r o p e n s i t y of males f o r s h i f t i n g and occupying o n l y p o r t i o n s of a t o t a l range at one time i s such that not a l l of 36 these ranges overlap simultaneously. Adult males No. 55 and No. 58 had ranges which overlapped c o n s i d e r a b l y throughout most o f the summer of 1956. S t r a i g h t l i n e distances between points of capture of both marten on the same day were c a l c u l a t e d and are summarized i n TABLE IX. The d i s t r i b u t i o n o f such distances appears random and suggest t h a t the movements of one marten were independent of the movements of the other. The a l t e r n a t e p o s s i b i l i t y was of one male f o l l o w i n g the other along the same route a f t e r a c e r t a i n time l a p s e . TABLE IX: S p a t i a l R e l a t i o n s Between Two Male Marten i n Overlapping Ranges T o t a l number of captures (1956) Male Male No.55 No.58 Captures of both marten on the same day 45 40 Greatest d i s t a n c e between extremes 2.3 or range ( m i l e s ) . 2.2 Distances between No.of captures per captures ( m i l e s ) , distance c l a s s . 0 - 0.5 7 0.5 - 1.0 4 1.0 - 1.5 5 1.5 - 2.0 1 Mean dis t a n c e and extremes: 0.7 (0 - 1.6) miles The record of i n t e r e s t was the capture of both marten i n the same t r a p , one of the l a r g e r model. Yifrien found by Mr. David Johnston, both were c o n s t a n t l y moving over and under each other w i t h i n the t r a p ; each f l e d d i f f e r e n t ways when r e l e a s e d . The most c r e d i b l e explanation holds that one chased the other i n t o the trap and immediately an (innate?) escape d r i v e replaced the former aggressive behaviour. 37 TABLE X gives the d i s t r i b u t i o n of captures r e l a t e d to a v a i l a b l e traps i n the combined ranges of the two males. TABLE X: Capture D i s t r i b u t i o n of Two Males i n Overlapping Ranges T o t a l traps i n combined ranges; 90  Traps capturing e i t h e r one or Traps capturing both males the other male •  No.of traps Captures per trap No.of traps Captures one male:other male 39 1 3 1 1 13 2 4 2 1 2 3 1 2 3 1 4 1 3 1 2 5 1 4 1 1 5 1 Percent of t o t a l traps capturing e i t h e r Percent of traps capturing male; 51 both males : 12  Whereas only one h a l f of the a v a i l a b l e traps captured e i t h e r marten, twelve percent captured both at l e a s t once. This suggests that although one appeared to move independently of the other, movements of both were confined to c e r t a i n p o r t i o n s of the area. I . Range D i s t r i b u t i o n of Female Marten. Maps 8 to 10 give the d i s t r i b u t i o n of female marten on the study area, and r e v e a l that summer ranges are q u i t e d i s c r e t e . Only two p o s s i b l e instances of overlapping were recorded In four summers. In 1955, female No. 61, an immature, occurred i n p a r t of a range subsequently occupied by adult No. 41, but d i e d a day a f t e r the f i r s t 1955 capture of No. 41 was obtained. In 1956, immature No. 72 was captured at one end of the known range of adult No. 68 on the same day that No. 68 was captured at the other end. S i x 38 other immature female marten, recorded from 1955 to 1957, were a l l found i n areas not occupied by other females. Among adult females, f o u r d i e d during the l i v e - t r a p p i n g . The ranges each p r e v i o u s l y h e l d , were subsequently re-occupied, but only a f t e r a lapse of 19 to 61 days (average 36 days) (TABLE X I ) . Female No. 65 was captured once at the edge of No. 64's former range, seven days a f t e r No. 64 had d i e d . TABLE X I : I n t e r v a l 7 B e f o r e Re-occupation of Vacated Ranges Adult Female Marten Marten which d i e d Marten re-occupying range I n t e r v a l i n days No. 95 No. 64 33 No. 41 No. 66 24 Nov 64 No. 71 61 No. 71 No. 65 19 Average: 36 No. 65 then disappeared and No. 71 subsequently re-occupied No. 64's range. Nineteen days a f t e r No. 71 d i e d , No. 3 5 waa recaptured there again. This occurred at the end of the trapping season i n 1956 and No. 65 was found i n the same area at the s t a r t of the tr a p p i n g i n 1957. When No. 65 d i e d , No. 37 was f i r s t i captured i n a sm a l l p a r t of No. 65's range s i x days l a t e r . Such data t a c i t l y imply that female marten occupy t e r r i t o r y , t hat i s , defended range. Vacated ranges may be t e n t a t i v e l y i n v e s t i g a t e d s e v e r a l days a f t e r abandonment by other females and l a t e r , completely.r-e-occupied. Some s p e c u l a t i o n may be allowed on the r o l e of scent posts i n 39 d e l i m i t i n g such areas. I f so, the delay i n re-occupation of a vacated range may be r e l a t e d to time necessary f o r o l d scent posts to fade. Hawley and Newby (1957) noted an even d i s t r i b u t i o n of marten ranges on t h e i r study area w i t h l i t t l e overlap except i n one case of two adult males. J . Movements of Male to Female Marten i n Breeding Season. I t i s c l e a r from the range maps presented that ranges of male marten overlap those of females. Marten have one breeding season a year i n J u l y and August (Ashbrook and Hanson, 1927). Markley and Bass e t t (1942) note that extreme l i m i t s of the mating season during seven years on a f u r farm were J u l y 10th to September 7 t h , ranging from twenty-four to f o r t y - s i x - d a y s i n a given year. C e r t a i n males are "extremely rapacious" and females are a n t a g o n i s t i c to each o t h e r , so f a t a l i t i e s occurred during t h i s p e r i o d . Enders and Leekley (1941) give c r i t e r i a f o r determining the e s t r a l stage i n female marten, on s i z e , shape and colour of the v u l v a . Most matings occur during maximum s w e l l i n g which l a s t s s i x to t h i r t y - n i n e days w i t h s l i g h t r e c e s s i o n s . Nearly a l l males are h i g h l y polygamous (Markley and B a s s e t t , op. c i t . ) . On J u l y 24, 1956, vulva examination of female No. 71 showed i t was i n heat. When i t was f i r s t found i n the l i v e -t r a p , a second marten thought to have been a male, was f r i g h t e n e d from the v i c i n i t y . On August 1 s t the same female was caught by Mr. David Johnston and as soon as i t was r e l e a s e d , two other marten, 40 which looked l i k e males, were observed. One passed about s e v e n t y - f i v e f e e t from the female which remained under a l o g . The second appeared almost immediately and attacked the female which ran i n t o a hollow l o g a f t e r i t freed i t s e l f . The male p e r s i s t e n t l y dug at a crack i n the l o g while the female sna r l e d w i t h i n . Observer disturbances s e v e r a l times f r i g h t e n e d the male about f i v e f e e t up a nearby t r e e , where i t growled, scolded and f l i c k e d i t s t a i l i n s q u i r r e l - l i k e f a s h i o n before r e t u r n i n g to the l o g . When the female f i n a l l y r a n out, i t was chased by the male i n t o a t h i c k e t , a f t e r which, the growling g r a d u a l l y diminished. Next day t h i s female die d i n a l i v e - t r a p and examination showed i t to be receding from f u l l e s t r u s . On J u l y 4, 1957, Mr. Douglas Brodie r e l e a s e d female No. 37 from a t r a p and i t was soon attacked by a l a r g e r marten which looked l i k e a male. I t dug i t s claws i n t o the female's back but I t screamed and escaped. The ensuing chase was punctuated by. screams, and observed f o r s e v e r a l minutes. Soon a f t e r , presumably the same male, was seen l y i n g on a l o g nearby. . The female had disappeared; i t i s not known i f i t was i n heat a t the t ime. In an a r b i t r a r i l y chosen p e r i o d of one week before and a f t e r observed estrus i n a given female, the l o c a t i o n o f males overlapping i t s range was noted during i n t e r v a l s of two days from each capture of t h i s female while i n heat. In 1956, distances t h a t male3 were captured from females Nos. 37 and 71 i n heat averaged 0.5 miles and ranged from 0.1 to 1.4 miles around a female i n heat, t h e r e f o r e i t seems that although males 41 do come to females then, they do not s t a y w i t h , or f o l l o w them f o r l o n g . K. D i s p e r s a l o f Immature Marten. In the summer of 1956 only four immature marten but eleven a d u l t s were captured on the study area, and i n 1957, ten immature and twelve a d u l t s were captured; e i g h t of these twelve a d u l t s were present on the area i n 1956. TABLE X I I approximately dates the occurrence o f new untagged marten on the t r a p p i n g area i n 1956 and 1957 and summarizes them according to sex and age. I t shows th a t of twenty-four marten which entered the area a f t e r t r a p p i n g had been c a r r i e d on f o r a week, f i f t e e n were immature animals and of these, about h a l f were thought to be young of the year. Fourteen of the f i f t e e n appeared i n August and most during the f i r s t two weeks. TABLE X I I : Appearance of New Marten on the Study Area -1956 and 1957 Male Female T o t a l adult Immature adult immature  W i t h i n one week a f t e r t r apping s t a r t e d . 4 2 6 May 24 - 31 1 1 June 1 - 1 5 1. 1 June 16 - 30 2 2 4 J u l y 1 - 1 5 1 1 J u l y 16 - 31 1 1 1 3 August 1 - 1 5 6 2 8 August 16 - 31 3 3 6 T o t a l 9 9 6 6 30 Maps 6 to 10 give the l o c a t i o n s of these animals. Three of the males appeared to e s t a b l i s h a range f o r short p e r i o d s ; No. 86 f o r e i g h t days, No. 91 f o r nine days and No. 88 f o r about 42 e i g h t out o f a twenty-four day i n t e r v a l . None of the other twelve appeared to occupy a range that was c l e a r l y d i s c e r n i b l e . The e i g h t l a s t seen a l i v e were caught an average of twice each, but f i v e of them were caught only once. Some showed tendencies to r e t u r n to one t r a p - s i t e f o r a few days. Male No. 87 was caught i n one trap f i v e out o f seven times i n eight days, male No. 73 three of four times i n seven days, and male No. 96 three of four times i n ten days. I t appears that a d i s p e r s a l of immature marten occurs during l a t e summer. This i n c l u d e s not o n l y j u v e n i l e young o f the year but marten i n the " y e a r l i n g " c l a s s as w e l l . Some may s e l e c t a temporary range approximating the s i z e used by a d u l t s , while others seem to move at complete l i b e r t y , although sometimes remaining i n s m a l l r e s t r i c t e d areas f o r a few days at a time. Hawley and Newby (1957) r e c o r d , as t r a n s i e n t s on t h e i r study area i n March 1954, two j u v e n i l e males which were captured s i x m iles away i n August 1954 and A U g U S t 1955 r e s p e c t i v e l y . One other j u v e n i l e ma-le, b e l i e v e d born on t h e i r area was captured about twenty-five miles away i n October. The former two " j u v e n i l e s " seem to b e . i n the " y e a r l i n g " group as defined i n t h i s study, and the i n f o r m a t i o n suggests-that d i s p e r s a l of immature marten may occur at any time i n the year. Perhaps more f i x e d ranges are not occupied by marten u n t i l they reach r e p r o d u c t i v e m a t u r i t y . P o p u l a t i o n s : A. D e n s i t i e s . Map 1 i n d i c a t e s an area a r b i t r a r i l y d e l i m i t e d which was assumed to be that i n which captured marten 43 l i v e d . T o t a l f o r e s t e d area w i t h i n these l i m i t s i s 4.5 square m i l e s . TABLE X I I I summarizes the number of i n d i v i d u a l marten captured there i n 1956 and 1957. TABLE X I I I : Marten Occurring i n 4.5 Square M i l e s of Forest -Summers Only  1956 male female 1957 male female Adults r e s i d e n t most of summer 4 3 4 A d u l t s o c c a s i o n a l l y captured 2 1 4 1 Immatures d i s p e r s i n g through the area 1 3 7 3 T o t a l 7 8 14 8 Approximate p o p u l a t i o n d e n s i t y per square mile 4 5 # excludes two former r e s i d e n t females had ranges re-occupied. which d i e d i n May and This suggests that the r e s i d e n t p o p u l a t i o n of marten was about two per square mile but the t o t a l number of marten us i n g the area was four to f i v e per square mile i n a g i v e n year. Hawley and Newby (1957) l i v e - t r a p p e d e i g h t y - f i v e i n d i v i d u a l marten i n two and one h a l f years i n 6.2 square miles of Montana f o o t h i l l s . Of these, f o r t y - s e v e n were captured over a p e r i o d not longer than a week and were classed as t r a n s i e n t s . Of the other t h i r t y - e i g h t , no more than twenty-seven r e s i d e d on the area at any one time. This completely r e s i d e n t p o p u l a t i o n f l u c t u a t e d between twenty-seven and fourteen i n d i v i d u a l s , or considering o n l y those l i v i n g completely w i t h i n the area boundaries, eleven or three i n d i v i d u a l s r e s p e c t i v e l y . D e n s i t i e s 44 computed on t h i s b a s i s are 4.4 or 1.8 marten per square mile at the h i g h p o p u l a t i o n and 2.3 or 0.5 marten per square mile at the low p o p u l a t i o n . The l a r g e s t l o s s was among the a d u l t females, and evidence of lower mean weights, lowered r e p r o d u c t i o n and s u s c e p t i b i l i t y to s t r e s s was obtained. The p o p u l a t i o n drop of marten was preceded by an observed drop i n small mammal abundance. B. Sex R a t i o s . Pur returns o f t e n show a preponderance of male marten, e s p e c i a l l y among animals trapped at the f i r s t p art of the w i n t e r . (Yeager, 1950, Douglas, 1953). Pur farm l i t t e r s c o n s i s t e n t l y show an average one to one sex r a t i o . (Markley and B a s s e t t , 1942, R i t c h i e , 1953) and the above l i v e -t r a pping s t u d i e s i n d i c a t e the same i n r e s i d e n t w i l d populations. Yeager ( o p . c i t ) a s c r i b e s the o s t e n s i b l e m a j o r i t y of males to t h e i r wider ranging h a b i t s which increases t h e i r chance of being caught. Q u a n t i t a t i v e data on p o p u l a t i o n d e n s i t i e s i s apparently r e s t r i c t e d to the Montana and Algonquin samples and comparison i s most t e n t a t i v e because d i f f e r e n t p o p u l a t i o n c o n d i t i o n s are i n v o l v e d . However, both i n d i c a t e that any g i v e n area has a r e l a t i v e l y small r e s i d e n t p o p u l a t i o n compared to the t o t a l number of animals found. Such a sampled area w i l l o c c a s i o n a l l y o b t a i n adjacent r e s i d e n t s and t r a n s i e n t i n d i v i d u a l s . Perhaps, from the more conservative estimates from the Montana area together w i t h one from Algonquin, a r e s i d e n t d e n s i t y of about two marten per square m i l e , u s u a l l y one male and one female, represents a norm. 45 Seasonal A c t i v i t y : Trapping r e s u l t s s t r o n g l y suggested that fewer marten were captured i n mid-summer than i n l a t e s p r i n g or l a t e summer. The summer was a r b i t r a r i l y d i v i d e d i n t o h a l f monthly periods and f o r each, a comparison of captures obtained w i t h the number expected was made. The expected number was determined from the ETN accumulated i n one pe r i o d compared to tha t necessary f o r each capture; t h i s i n t u r h was the average ETN per capture f o r the e n t i r e summer. Graph 1 compares the observed and expected captures f o r 1956 and 1957 and shows that f o r at l e a s t one month i n mid-summer,"observed captures are below the expected, suggesting that marten a c t i v i t y has lessened. During the low p e r i o d i n 1956 (June and e a r l y J u l y ) , eleven " D a i l y Movement Index" observations on four male marten averaged 0.7 m i l e s , the same average as t h i r t y - f o u r other observations on the same marten throughout the r e s t of the summer. This suggests that movements do not d i m i n i s h i n e x t e n t , but perhaps only i n number, i n mid-summer. I t i s d i f f i c u l t to evaluate a l l the c o n t r i b u t i n g f a c t o r s to such seasonal v a r i a t i o n . P o s s i b l y , the increase i n a v a i l a b l e food a f t e r l a t e s p r i n g reduces the necessary a c t i v i t y , and i t may a l s o reduce b a i t e f f e c t i v e n e s s . - The onset of the r u t i n l a t e summer may then increase a c t i v i t y and i n t e n s i f y the r e a c t i o n of marten to scented b a i t . Yeager and Remington (1956) noted that 46 of 66 (70$) of marten observations i n Colorado occurred i n June, J u l y and August; t h i s may have i n d i c a t e d more d i u r n a l a c t i v i t y 46 a c t i v i t y during the breeding season. R e l a t i o n of Marten to Forest Cover: A. Winter. From December 29 to January 3, 1956-57, a l l the marten t r a p - l i n e s were t r a v e l l e d and the l o c a t i o n of every marten tr a c k c r o s s i n g , or coming w i t h i n f i v e f e e t e i t h e r side of the t r a i l , was recorded and assigned to i t s c o r r e c t f o r e s t type. This was assumed to give an e v a l u a t i o n of h a b i t a t preference. TABLE XIV compares t r a c k occurrences i n the four b a s i c f o r e s t types w i t h expected occurrences, i f the tra c k s were e q u a l l y d i s t r i b u t e d . This shows th a t there are s i g n i f i c a n t l y more t r a c k s found I n c o n i f e r and s i g n i f i c a n t l y TABLE XIV: Winter Occurrence of Marten Tracks i n Ba s i c Forest Types. Forest C CH HC H Miles censused 3.3 2.5 2.2 7.9 Tracks observed 19 12 9 16 Tracks expected 11.7 8.8 7.8 28.1 S i g n i f i c a n t d i f f e r e n c e 4.6 ', 5.2 fewer I n hardwoods, and that mixed f o r e s t s comprised predominantly of c o n i f e r have s l i g h t l y more tr a c k s than those w i t h more hardwoods• On January 9, 1957, twent y - f i v e snow depth measurements i n each o f the fo u r types were made; l o c a t i o n s were pre-s e l e c t e d on a map. Fiv e measurements of t o t a l snow depth and l e v e l of f i r s t c r u s t were taken at each l o c a t i o n ; four measurements were made at r i g h t angles s i x f e e t away from a c e n t r a l measurement. 47 Graph 2 gives the snow depth and an index of marten a c t i v i t y based on t r a c k s per mile f o r each f o r e s t . I t suggests that an i n v e r s e r e l a t i o n e x i s t s such t h a t winter marten a c t i v i t y c oincides w i t h heavier cover and r e s u l t a n t shallower, s l o w l y accumulating snow. F i e l d observations showed most marten a c t i v i t y i n hemlock stands. Only some spruce-balsam showed marten a c t i v i t y ; i n such areas, snow i s deeper but there i s considerable wind s h e l t e r and sometimes numerous openings under blowdown, logs and stumps. In mixed f o r e s t s , observations i n d i c a t e d that marten move c h i e f l y near the c o n i f e r p o r t i o n s . Track occurrences show that marten are not continuously present i n the favoured c o n i f e r stands but tend to move from one to the other i r r e g u l a r l y . One marten l i v e d i n the pantry and a t t i c of the W i l d l i f e Research S t a t i o n during p a r t o f March 1956 and o l d marten scats were c o l l e c t e d i n the deserted Minnesing Lodge a t t i c i n August 1956. B. Summer. The number of captures of marten i n any given f o r e s t type was assumed to be i n d i c a t i v e of the extent of a c t i v i t y i n , and preference shown f o r , the type. TABLE XV compares captures i n the f o u r major types f o r 1956 and 1957, to that expected from each summer's average. The comparison between captures before and a f t e r mid-July gives only three instances where the observed and expected captures d i f f e r s i g n i f i c a n t l y but these appear u n r e l a t e d , and are b e l i e v e d due to f a c t o r s other than a s e l e c t i o n of f o r e s t s . One such d i f f e r e n c e r e s t s p r i m a r i l y on nine recaptures of one marten i n a small b l a c k spruce bog. 4.8 When considered over the e n t i r e summer f o r each year, there i s no s i g n i f i c a n t d i f f e r e n c e among captures i n d i f f e r e n t f o r e s t s , and i t i s concluded that marten do not show s e l e c t i o n f o r one or another type. TABLE XV: Marten Captures i n Basic Forest Types. Forest C 1956 CH HC H C CH 1957 HC H May 16-July 15 captures 16 15 11 34 23 16 10 22 Expected 22.3 16.1 15.1 44.7 17.0 13.1 7.9 27.0 S i g n i f i c a n t d i f f e r e n c e - - - - - -J u l y 16-Aug.31 captures 39 22 25 41 7 24 9 27 Expected 23.8 18.6 12.1 44.5 16.9 16.9 7.1 30.3 S i g n i f i c a n t d i f f e r e n c e yes yes yes E n t i r e summer May 16-Aug.31 captures 55 37 ,36 75 30 40 19 49 Expected 46.1 34.8 27.2 89.8 31.7 30.1 15.0 57.3 S i g n i f i c a n t d i f f e r e n c e — — — — — — — — Although the winter data are few, i t seems that marten concentrate t h e i r a c t i v i t y i n c o n i f e r f o r e s t s i n winter and spread through a l l adjacent f o r e s t s i n summer. She l t e r from c l i m a t i c extremes appears a more important b a s i s f o r s e l e c t i o n than e i t h e r -the s t r u c t u r e or f l o r a l c o n s t i t u e n t s of -the f o r e s t i t s e l f . Hawley and Newby (1957) noted that home ranges of marten sometimes c o i n c i d e d w i t h the edge o f l a r g e open meadows and 49 burns and a l s o observed that on numerous occasions marten seemed to avoid e n t e r i n g areas that l a c k overhead cover. Williams (1957) observed one marten which refused to f o l l o w a hare i n t o a f o r e s t c l e a r i n g . Records o f marten i n human h a b i t a t i o n s may p a r t i a l l y support t h i s view o f the importance of adequate s h e l t e r . In Ontario, de Vos and Guenther (1952) r e p o r t that favoured winter h a b i t a t i n Chapleau Is cedar swamp r a t h e r than s p r u c e - b i r c h , jack.pine or b l a c k spruce f o r e s t s . MacFie (1954) notes that marten i n the extensive timbered muskeg o f extreme north-western Ontario l i v e on spruce, tamarack r i d g e s r a t h e r than i n the muskeg i t s e l f . I n the Rocky Mountains, marten i n winter use dense a l p i n e f i r ( M a r s h a l l , 1951b) or Engelraann spruce, a l p i n e f i r at e l e v a t i o n s ranging from 9500 to 11,500 f e e t (Remington, 1950). In Washington, winter preference was shown f o r Douglas f i r , cedar and hemlock (de Vos and Guenther, l o c . c i t . ) . In i n t e r i o r A l a s k a , marten occur only i n areas dominated by spruce f o r e s t "which i s apparently the b a s i c element of t h e i r h a b i t a t " . (Lensink, 1953). M a r s h a l l (op. c i t . ) noted that marten are l e s s r e s t r i c t e d i n summer months when they are then report e d f o r a g i n g i n open rock s l i d e s , brush s l i d e s and meadows as w e l l as i n f o r e s t s . A l t i t u d i n a l migrations are suspected i n some mountain areas. As c i t e d by Lensink (op. c i t . ) such movements are reported from mountains to c o a s t a l beaches i n south-eastern A l a s k a , from summer ranges at 4000 to 5500 f e e t to winter ranges below 3400 f e e t i n the Cascade Mountains and, apparently, there 50 i s evidence of an upward winter movement i n the Uinta Mountains. Hawley and Newby. (1957) found no seasonal movements of r e s i d e n t marten i n G l a c i e r Park, Montana. I t seems c l e a r from t h i s evidence that marten h a b i t s are r e l a t e d to f o r e s t cover i n a most s u p e r f i c i a l way o n l y , w i t h no c l e a r l y d e f i n e d c o n t r o l l i n g f a c t o r inherent i n the f o r e s t i t s e l f . 51 PART I I : POOD ANALYSES Several reports on food of marten are a v a i l a b l e , the most extensive and s y n o p t i c being by L e n s i n k e t . a l . (1955). An attempt was made i n the present study to do a more i n t e n s i v e survey to f i n d seasonal v a r i a t i o n s i n major food types. Methods: Fresh marten scats were c o l l e c t e d d a i l y while t r a p - l i n e s were operated i n the summers of 1955, 1956 and 1957. At the beginning of the summer tra p p i n g season, p a r t i c u l a r l y i n 1957, many o l d scats were obtained which probably gave a sample of winter and e a r l y s p r i n g foods. Marten scat c h a r a c t e r i s t i c s are perhaps best described by Murie (1954)j f r e s h m a t e r i a l has a p e c u l i a r nauseating odour. In s i z e , marten scats tend to range between those of mink and f i s h e r . The presence of f i s h e r on the study area introduced a p o t e n t i a l source of e r r o r i n r e c o g n i z i n g some s c a t s , but a l l dubious m a t e r i a l was r e j e c t e d to reduce m i s i d e n t i f i c a t i o n e r r o r s to a minimum. Scats were d r i e d i n the l a b o r a t o r y then analysed f o r occurrence o f d i f f e r e n t items. D i r e c t comparison w i t h known m a t e r i a l enabled i d e n t i f i c a t i o n of b e r r y seeds to genus and sometimes s p e c i e s . Seeds from raspberry jam b a i t could be d i s t i n g u i s h e d from w i l d Rubus species on the b a s i s of s i z e and shape. Mammal h a i r of those species l i v i n g on the study area, could best be I d e n t i f i e d by d i r e c t comparison to known h a i r 52 samples. S i z e , colour and proportions of various h a i r s are d i a g n o s t i c i n most species and the i d e n t i f i c a t i o n of some samples can be confirmed by d i s t i n c t i v e t e e t h , and o c c a s i o n a l l y r e l a t i v e s i z e s of f e e t and claw, remains. In two groups, t e n t a t i v e d i a g n o s t i c characters were used. The two jumping mice were separated on r e l a t i v e d i f f e r e n c e i n colour gradient on the l a r g e b l a c k guard h a i r s . Zapus shows a gradual blending from black t i p to white base, whereas Napaeozapus has a comparatively more abrupt t r a n s i t i o n from b l a c k to white at the middle of the h a i r . Napaeozapus- has few b l a c k h a i r s l a t e r a l l y so t h a t t u f t s o f f u r there appear b r i l l i a n t orange. These characters overlap to some extent and are used w i t h r e s e r v a t i o n . Frequency o f occurrence of jumping mice i n marten sc a t s i s given c o l l e c t i v e l y under Zapodidae; subsequent d i f f e r e n t i a t i o n , based on the above c h a r a c t e r s , Is given p a r e n t h e t i c a l l y . There appears to be no c o n s i s t e n t e x t e r n a l d i f f e r e n c e among shrew h a i r . Comparison o f c u t i c u l a r s c a l e patterns was made, using Williamson's (1951) method of impressing h a i r s i n p o l y v i n y l acetate. Again no d i f f e r e n c e s appeared i n sc a l e patterns among Sorex ciner.eus, Sorex fumeus and Microsorex h o y i , but B l a r i n a  brevicauda u s u a l l y had a more i r r e g u l a r arrangement of s cales on the middle p o r t i o n of the h a i r s , compared to a r e g u l a r b l o c k -l i k e p a t t e r n of Sorex spp. Since there was some overlap i n t h i s character between B l a r i n a and Sorex spp. i d e n t i f i c a t i o n based e n t i r e l y on h a i r impressions are recorded p a r e n t h e t i c a l l y beside those confirmed by other remains i n the s c a t . 53 Results t TABLES XVI, XVII, and XVIII i t e m i z e dated summer m a t e r i a l from 1955 to 1957 and TABLE XIX gives dated winter items and undated m a t e r i a l . Graphs 3 to 14 compare the seasonal occurrence of major food items. A l l comparisons were made f o r h a l f month i n t e r v a l s except June 1956 when fewer data were obtained. A. Food Items. 1. Mammals. S o r i c i d a e . In 1955 and 1956, shrews formed o n l y a s m a l l p a r t of t o t a l food but i n 1957 they c o n s t i t u t e d about twelve percent of a l l summer items and at times t o t a l l e d about a t h i r d of the s m a l l mammals eaten. Evidence from winter i n d i c a t e s t h a t shrews c o n s t i t u t e up to twenty percent of the d i e t . This i s of some i n t e r e s t compared to the l a c k of shrews i n marten food i n Alaska (Lensink e t . a l . 1955), B r i t i s h Columbia (Cowan and Mackay, 1950) and Washington (Newby 1951) a t t r i b u t e d to an o s t e n s i b l e a v e r s i o n to the smell of shrew musk glands. Captives eagerly accepted a l l shrews, although one p a r t i c u l a r l y musky B i a r i n a was l e f t uneaten by one marten. Two captives h e l d by Remington (1952) accepted shrews. C r i c e t i d a e . Scats presumed to be from winter or l a t e s p r i n g contained Clethrionomys more than any one other animal, but fewer are found i n summer. Snap-traps obtained s i x t y - f i v e percent of the C l e t h r i o n o m y 3 i n pure or predominantly c o n i f e r stands (p. 6 3 ) ; f o r t y percent were from pure c o n i f e r f o r e s t s . The high number eaten by marten i n winter may be explained by the preference marten show f o r the same h a b i t a t then. 54 TABLE XVI: A n a l y s i s of Marten Scats - Summer, 1955 Based on dated m a t e r i a l : 240 items i n 192 scats June J u l y August 16-30 1-15 16-31 1-15 16 -31 No. of scats 2 2 54 17 17 No. of items 4 2 150 49 35 OCCURRENCES Sorex sp. 1 (1) Clethrionomys 2 4 Microtus 1 5 3 4 Peromyscus 5 1 2 Zapodidae 1 12 3 4 (Napaeozapus) (1) (9) (3) (4) (Zapus) (3) Tamias 3 3 Tamiasciurus 1 Lepus 1 Odocoileus 1 T o t a l mammals: 1 1 29 12 15 B i r d sp. 1 1 17 5 1 B i r d egg sp. 1 1 2 Hymenoptera: Vespoidea 1 2 2 2 Pormicoidea 1 Coleoptera spp. 1 1 Insect sp. 1 A r a l i a N u d i c a u l i s 31 2 5 Vaccinium spp. 38 12 3 Rubus spp. 22 9 1 Cornus canadensis 2 Prunus s e r o t i n a 1 2 Seed sp. 2 Duff debris (Rubus b a i t t o t a l l e d 8 items) 1 3 55 TABLE XVII: A n a l y s i s o f M a r t e n Scats - Summer, 1956 Based on dated m a t e r i a l : 511 items i n 261 scats # see text May June J u l y August 16-31 1-15 16-30 1-15 16-31 1-15 16-31 No. of scats 2 11 10 34 72 81 51 No. of items 4 23 20 57 140 153 114 Occurrences Sorex sp. (1) (1) (1) (2) (3) B l a r i n a (1) Clethrionomys 5 2 2 6 Mlcrotus 4 3 3 28 19 10 Synaptomys 1 2 Peromyscus 1 1 2 1 5 7 Zapodidae 1 2 17 16 1 (Napaeozapus) (1) (2) (16) (14) (1) (Zapus) (1) (1) Tamias 3 5 5 1 Tamiasciurus 1 1 1 1 Lepus 2 1 Odocoileus 1 Alces# 1 T o t a l mammals: 2 11 10 12 55 46 29 B i r d sp. 3 1 28 32 22 4 B i r d eggs sp.. 1 2 2 2 3 1 Snake sp. 2 Amphibian sp. 1 Hymenoptera: Vespoidea 2 17 24 12 Coleoptera: S i l p h i d a e 2 spp. 1 2 4 7 5 1 D i p t e r a : Mycetophilidae 1 Insect pupa 1 1 1 3 Insect, sp. 1 3 3 1 Gasteropoda: Zonitoides S t r o b i l o p s Discus Ghilopoda: L i t h o b i u s type P l a n t s : A r a l i a n u d i c a u l i s Vaccinum spp. Rubus spp. Seed sp. Gramineae Duff d e b r i s " T a r - l i k e " d e b r i s 2 1 1 1 1 2 1 1 9 2 27 11 5 3 1 7 1 34 9 18 3 3 (Rubus b a i t t o t a i l e d 43 items) 56 TABLE X V I I I : A n a l y s i s of Marten Scats - Summer, 1957. Based on dated m a t e r i a l : 614 items i n 349 scats May June J u l y Augus t 16-31 1-15 16-30 1-15 16.-31 1-15 16-31 No. of scats 42 65 30 33 56 71 52 No. of items 60 101 49 55 106 151 92 OCCURRENCES Sorex sp. 1 (1) (9) 7 (3) 2 (9) 2 (4) 6 (1) 3 B l a r i n a (1) 4 (3) 5 1 (3) (1) 2 6 (1) 1 Parascalops 1 Clethrionomys 7 5 2 4 3 4 12 Microtus 10 7 7 9 4 11 14 Syhaptomys 1 1 Peromyscus 12 7 4 6 9 7 4 Zapodidae 4 5 2 3 14 20 13 (Napaeozapus) (3) (3) (2) (3) (12) (15) (8) (Zapus) (1) (2) (2) (4) (5) Tamias 4 19 7 5 1 3 Tamiasciurus 2 5 2 Glaucomys 1 Lepus 2 1 Odocoileus 2 1 T o t a l mammals: 60 751 26 36 45 60 50 B i r d sp. 3 9 3 7 10 8 1 I i r d e ( p i n feathers) 1 13 3 6 7 1 B i r d egg sp. 1 3 3 2 1 Snake sp. 2 1 Amphibian sp. 1 2 Hymenoptera: Vespoidea 3 3 2 2 Coleoptera sp. 1 1 S i l p h i d a e 1 Emphemeroptera sp • 1 Insect sp. 1 2 2 2 1 Gasteropoda sp. 1 Zonitoides 1 1 Discus 1 Polygyridae 1 P l a n t s : A r a l i a n u d i c a u l i s 4 13 Rubus spp. 2 21 48 29 Vaccinium spp. 4 8 4 Prunus s e r o t i n a 1 Streptopus roseus 1 1 Taxus canadensis 1 Seed sp. 2 4 2 Duff - Debris 6 7 1 1 " T a r - l i k e " d e b r i s 1 Rubus b a i t t o t a l l e d 31 items. 57 TABLE XIX: A n a l y s i s of Marten Scats - Winter and E a r l y Spring Dated m a t e r i a l : 5 items i n 4 scats ( e a r l y January 1957) and 1 item from February 8, 1956. Undated m a t e r i a l : 184 items from 140 scats No. of scats No. o f items OCCURRENCES Sorex sp. B l a r i n a Clethrionomys Microtus Synaptomys Peromyscus Zapodidae (Napaeozapus) (Zapus) Tamias Tamiasciurus Sciurus Myotis sp. Lepus Odocoileus Dated winter m a t e r i a l 5 6 1 1 1 1 Undated m a t e r i a l 1956 1957 28 35 6 (1) 2 14 6 2 1 1 2 112 151 2 (16) 8 (3) 23 20 2 12 11 (8) 15) 18 4 1 1 8 11 T o t a l mammals: 35 140 B i r d sp. Snake sp. Hymenoptera: Vespoidea Coleoptera: S i l p h i d a e Insect sp. Duff d e b r i s 1 1 2 1 Microtus c o n s i s t e n t l y appeared as a food item I n winter and summer. Numerous t o t e roads and small openings i n the f o r e s t probably c o n t r i b u t e d most i n d i v i d u a l s . Synaptomys i s considered a r a t h e r scarce species i n the re g i o n and occurred i n f r e q u e n t l y i n marten s c a t s . Although Peromyscus comprised f o r t y percent of mammals snap-trapped i n 1956, they formed only 58 twelve percent of the items i n scats i n 1956 and fourteen percent i n 1957. Perhaps the r a t h e r marked preference shown by Peromyscus f o r hardwoods and s t r i c t l y n o c t u r n a l h a b i t s , expose them l e s s to marten p r e d a t i o n . In the winter and s p r i n g of 1957, they comprised only eight percent of the items; marten, i t i s r e c a l l e d , showed an avoidance of hardwoods i n w i n t e r . Other small mammals. Both jumping mice occurred, Napaeozapus more f r e q u e n t l y than Zapus. One mole, Parascalops, was recorded, and one b a t , Myotis sp. from an undated, probably winter s c a t , may have been found h i b e r n a t i n g . S c i u r i d a e . Red s q u i r r e l s occurred as marten food q u i t e uncommonly. In three summers they accounted f o r no more than one percent of the items and i n the winter and s p r i n g samples, only about three percent were s q u i r r e l s . This i s of i n t e r e s t because of a former a l l e g e d importance of r e d s q u i r r e l s as marten food ( M a r s h a l l , 1946, 1951b). One r e c o r d of a b l a c k s q u i r r e l i n 1957 coincided w i t h s e v e r a l s i g h t observations f o r the species that year; Algonquin i s at the p e r i p h e r y of the species range. Only one record was obtained fo r . a f l y i n g , s q u i r r e l . A scat c o l l e c t e d f r e s h on January 10, 1957, contained chipmunk f u r . This suggests that h i b e r n a t i n g animals are a p o t e n t i a l food source i n w i n t e r . Chipmunks are eaten r e g u l a r l y i n summer and In June 1957 they accounted f o r over one quarter of the mammals eaten and formed about one h a l f of the items found i n scats by a marten den discovered a t the same time. This c o i n c i d e d w i t h an observed abundance of the species i n hardwood f o r e s t s . 59 Larger mammals. Snowshoe hares occurred r a r e l y i n summer food but In winter and e a r l y s p r i n g c o n s t i t u t e d about f i v e percent of a l l items. Deer, undoubtedly c a r r i o n , a l s o occurred more frequently, i n winter scats but never formed over seven percent of food items. Marten tracks i n winter showed that wolf k i l l s of deer were i n v e s t i g a t e d by marten, but not thoroughly; no tendency to s t a y around c a r r i o n was- observed. M a r s h a l l (1951b) records one marten which remained four days around an e l k c a r c a s s , and Yeager and Remington (1956) record a marten feeding on v i s c e r a of an e l k k i l l ; t h i s l a t t e r i n d i v i d u a l was thought to be the same one seen at the k i l l during the two succeeding days. One occurrence o f moose h a i r i n 1956 was based on a few h a i r s i n a scat which a l s o contained remains of b i r d eggs. This s t r o n g l y suggests the h a i r s were part of the nesit l i n i n g . 2. B i r d s and B i r d Eggs. B i r d s appear most s u s c e p t i b l e to marten p r e d a t i o n during the n e s t i n g season. In the f i r s t p art of J u l y 1956, b i r d s comprised one h a l f of marten food; numerous q u i l l sheathes suggested t h a t most were n e s t l i n g b i r d s . In 1957, b i r d s formed one t h i r d of marten food towards the end of June, and of s i x t e e n items, t h i r t e e n showed the sheathed feathers of n e s t l i n g s . The p r o p o r t i o n of b i r d s eaten increases q u i c k l y from s p r i n g to the n e s t i n g season then decreases over the r e s t of the summer. B i r d eggs are taken r e a d i l y and comprise up to ten percent of t o t a l marten food j u s t b e f o r e , or during the peak occurrence of b i r d food items. Incubating b i r d s , p a r t i c u l a r l y ground n e s t e r s , are perhaps more vulnerable to 60 marten p r e d a t i o n . Evidence i n one case pointed s t r o n g l y to a marten having taken an i n c u b a t i n g thrush and i t s eggs; s e v e r a l other occurrences of blue egg fragments suggest p r e d a t i o n on thrushes. In one other i n s t a n c e , l a r g e white fragments, p o s s i b l y of grouse eggs, were found. 3. R e p t i l e s and Amphibians. S i x scats i n three summers, contained p o r t i o n s of snake s k i n and s c a l e s . The s i z e of b e l l y acutes and keeled l a t e r a l s c a l e s suggested that most were g a r t e r snakes, the most common l o c a l s p e c i e s . A l l but one occurred i n s p r i n g or e a r l y summer when most main foods of marten were r e l a t i v e l y s c a rce. Amphibian remains were not always easy to determine. Four were noted, and were most probably f r o g s . Captive marten r e a d i l y accepted leopard f r o g s . 4. Insects and Other I n v e r t e b r a t e s . B e e t l e s and wasps predominated i n i n s e c t remains found i n s c a t s . In 1956, beetles occurred i n the f i r s t p a r t of the summer c o i n c i d e n t w i t h an abun-dance of Carabid b e e t l e s i n maple f o r e s t s during the f i r s t two weeks of June. In s e v e r a l instances i n e a r l y summer, i d e n t i f i a b l e remains of S i l p h i d b e e t l e s , genus N eorophorus, give f u r t h e r i n d i c a t i o n s of c a r r i o n as food. Beetles cease to be of any importance as a food item by the end of J u l y . During the l a t t e r p a r t of the summer, e s p e c i a l l y i n 1956, wasps g r a d u a l l y became the main component of i n s e c t food, and i n the f i r s t p a r t of August 1956, comprised s i x t e e n percent of t o t a l food. The most complete remains of wasps i n scats were s u f f i c i e n t to i d e n t i f y the species as a member of the Vespdidea. I t i s not 61 known i f marten a c t u a l l y s e l e c t wasps as food, or i f wasps are eaten while the animalr i s f o r a g i n g near wasp n e s t s . Remains of i n s e c t pupa cases appear i n scats o c c a s i o n a l l y and suggest that marten forage i n ground l i t t e r to a c e r t a i n extent. On August 24, 1954, a marten was observed i n v e s t i g a t i n g c r e v i c e s i n the rough bark of a yellow b i r c h ; perhaps i t was a f t e r i n s e c t s . Other i n v e r t e b r a t e s appear i n f r e q u e n t l y i n marten s c a t s . Eleven occurrences of s n a i l s w i t h i n s c a t s , and not under them, i n d i c a t i n g they had been eaten, revealed i d e n t i f i a b l e remains of at l e a s t three s p e c i e s . One centipede was recorded. 5. B e r r i e s . With the r i p e n i n g of b e r r i e s towards the end of summer, marten d i e t s changed over c o n s i d e r a b l y and scats c o n t a i n i n g nothing but b e r r y seeds appeared. Quite p o s s i b l y t h i s food has a l a x a t i v e e f f e c t on marten since there was a decided increase i n the number of scats found during b e r r y season. A r a l i a n u d i c a u l i s appeared to be a p r e f e r r e d species i n a l l three years. B l u e b e r r i e s (probably both Vaccinium  a u g u s t i f o l i u m and V. m y r t i l l o i d e s ) were r e a d i l y taken, and i n l a t e J u l y and e a r l y August 1955, c o n s t i t u t e d about on quarter of the t o t a l food. Raspberries (both Rubus idaeus and R. pubescens) i n the same p e r i o d i n 1957 comprised one quarter of t o t a l food, hence i t appears that extent of u t i l i z a t i o n depends on a v a i l a b i l i t y . Although d e f i n i t e preference appears to be shown f o r the above s p e c i e s , o c c a s i o n a l occurrences o f seeds of Cornus 62 canadensis, Frunus serotina-, Streptopus roseus and Taxu3  canadensis i n d i c a t e d sampling of various other b e r r i e s as w e l l . The eagerness w i t h which c a p t i v e marten accept sweet substances may i n d i c a t e t h a t s e l e c t i o n of species of f r u i t s depends l a r g e l y on t h e i r sweetness. On J u l y 22, 1955, a marten r e l e a s e d from a l i v e - t r a p stopped, then detoured a few f e e t to eat a r i p e c l u s t e r of A r a l i a n u d i c a u l i s b e r r i e s before continuing on; i t ignored c l u s t e r s of r i p e Cornus canadensis b e r r i e s a l s o present. On August 9, 1957, Mr. Douglas Brodie observed a marten, r e l e a s e d from a l i v e - t r a p , run about f i f t y f e e t away and climb on to a l o g about two and one h a l f f e e t h i g h . I t then jumped on a r a s p b e r r y bush and devoured r i p e b e r r i e s while weighting the p l a n t down. This' procedure was repeated f o r a l l raspberry bushes about the l o g . 6. Duff d e b r i s . Duff debris ( c o n i f e r needles, l e a v e s , decayed wood etc.) was recorded only i f one h a l f or more of a s c a t was composed of i t . Almost a l l scats contained d e b r i s to some extent and some were composed e n t i r e l y o f i t . Scats containing a considerable p r o p o r t i o n of d e b r i s f u r t h e r i n d i c a t e that marten feed on items i n the ground l i t t e r . P o s s i b l y some of the d u f f m a t e r i a l i s eaten f o r mineral content, perhaps s a l t s from animal u r i n e . A few scats contained b l a c k t a r - l i k e m a t e r i a l , p o s s i b l y congealed blood. One scat of t h i s s o r t came from a l i v e - t r a p p e d marten which was reported to have appeared s i c k . Two scats contained only f r e s h green g r a s s . 63 B. Food Trends. Small mammals are the major marten food. The few dated winter scats contained them almost e n t i r e l y and undated m a t e r i a l presumably r e p r e s e n t i n g winter and e a r l y s p r i n g food, contained at l e a s t e i g h t y percent small mammals. Graphs 3 to 14 show v a r i a t i o n s i n small mammal occurrences i n summer marten d i e t s . In e i t h e r l a t e June or e a r l y J u l y , depending on the year, b i r d s predominate over mammals and i n l a t e summer at any time from the end of J u l y through to the end of August, again varying from year to year, b e r r i e s r e p l a c e mammals as a predominant food. Insects r e g u l a r l y c o n s t i t u t e p a r t of summer food but may vary i n the p r o p o r t i o n from year to year. C. Food Preference of Captive Animals. Captive marten In Algonquin r e a d i l y accepted a l l mammals, b i r d s , eggs, raw meat and sweets o f f e r e d . C r a y f i s h , f i s h , peanut b u t t e r , cooked potatoes and oranges, o f f e r e d to two, were r e f u s e d . Unnatural foods marten are known to eat i n c l u d e canned salmon, r a i s i n s (Walker ,1929), m i l k , bread, bo-iled r i c e , cornmeal, graham mush, cooked prunes, (Anonymous, 1936), pancakes, gravy, c e r e a l , f i s h , l e t t u c e , c e l e r y , s p a g h e t t i , (Remington, 1952), grease, (Hawbecker, 1946), c a r r o t s , cabbage and t u r n i p s i n a c e r e a l w i t h meat, poached eggs, (Anonymous, 1952), candy, cheese, peanut b u t t e r , (Lensink e t . a l . 1955), and even f i g newtons, (Orsborn, 1953). Remington ( l o c . c i t . ) notes they ignored salamanders and toads. C l e a r l y , l i t t l e preference i s shown, although on the b a s i s of observations i n Algonquin, meats and sweets are most eagerly accepted. 64 D. R e l a t i o n to A v a i l a b i l i t y o f Small Mammal Food U t i l i z e d . In 1956, an attempt was made to compare small mammals eaten by marten w i t h those obtained by snap-trapping. Two tra p p i n g methods were employed. One set twelve trapping s t a t i o n s f i f t y f e e t apart on each of two p a r a l l e l l i n e s two hundred f e e t a p a r t . Each s t a t i o n had f i v e snap-traps, one by the stake marking the l o c a t i o n and the others at r i g h t angles, f i v e f e e t from the stake. B a i t used was a mixture o f peanut b u t t e r , r o l l e d o a t s , bacon and crushed walnut. These l i n e s sampled f o r e s t types C (hemlock and mixed c o n i f e r ) , CH (hemlock, sugar maple), HC ( w h i t e - b i r c h , spruce) and H (sugar maple) f o r three n i g h t s each i n e a r l y June, J u l y and August. Locations of these are given on Map 1. Information was provided by the Ontario Research Foundation on snap-trapping which sampled f o r e s t types HC ( w h i t e - b i r c h , spruce), CH (spruce, white b i r c h ) and H (sugar maple) f o r three n i g h t s a month from May to September. These l i n e s were l o c a t e d no f u r t h e r than four miles from the marten study area. Four l i n e s of one hundred traps each, i n c l u d i n g some r a t t r a p s , were i n groups o f f i v e at r i g h t angles to the l i n e at i n t e r v a l s v a r y i n g from f i f t e e n to f i f t y f e e t . They were b a i t e d w i t h e i t h e r d r i e d a p r i c o t s , f i g s , prunes, or a mixture of peanut b u t t e r and r o l l e d o a t s . R e s u l t s from a l l these l i n e s were combined, to present a general impression of p r o p o r t i o n a l abundance of small mammals during_the summer. A percent composition of such a p o p u l a t i o n , based on 359 captures of s m a l l mammals i n over 20,000 t r a p - n i g h t s , i s compared i n Graph 15 to a percent of d i f f e r e n t species i n 148 65 occurrences i n marten scats through the same summer. There i s a considerable d i f f e r e n c e between the two evaluations of Microtus p r i m a r i l y because no area trapped was optimum Microtus h a b i t a t . In August, about one hundred t r a p -night s obtained s i x Microtus pensylvanicus i n a grassy tote road and along p a r t of a grassy creek edge; t h i s seemed to i n d i c a t e they were r e a d i l y a v a i l a b l e i n such areas. Discrepancies between the two values f o r Peromyscus and Napaeozapu3 seem most l i k e l y due to h a b i t a t s e l e c t i o n of these s p e c i e s . About s i x t y percent o f Peromyscus trapped were i n sugar maple f o r e s t s and two t h i r d s of the Napaeozapus found were i n various hardwood or mixed predominantly hardwood f o r e s t s . Since marten move r e a d i l y through a l l f o r e s t types, these species would not be continuously a v a i l a b l e . There i s no consistency i n the r e l a t i v e number of any one species eaten through the v a r i o u s periods of one summer, or from one summer to the next. D i f f e r e n c e s probably e x i s t i n the v u l n e r a b i l i t y of c e r t a i n s m a l l mammals to carnivore p r e d a t i o n , and i t i s known that -t r a p - l i n e s are not a f u l l y r e l i a b l e means of measuring r e l a t i v e abundance of d i f f e r e n t species ( S t i c k e l , 1948). These and other f a c t o r s may a l t e r d i f f e r e n c e s between a trapping and a p r e d a t i o n e v a l u a t i o n of species abundance, so that data g i v e n on Graph 15 probably are not s t r i c t l y comparable. Nevertheless, i t does suggest that the mammals eaten by marten probably are s e l e c t e d on the b a s i s o f a v a i l a b i l i t y ; t h i s , w i t h the observed randomness of hunting behaviour i n marten, gives reason to suppose t h a t no species of s m a l l mammal i s s p e c i f i c a l l y hunted. 66 TABLE XX summarizes food stud i e s reported to date, and i s adapted i n part from Lensink e t . a l . (1955). Composition i s expressed as a percent o f frequency occurrence. Results of the present study are summarized f o r comparison. G e n e r a l l y , the same types of food occur i n a l l the samples; v a r i a t i o n s i n r e l a t i v e abundance depends on the a v a i l a b i l i t y o f such foods i n each area, and, p a r t i c u l a r l y i n the summer, on the d i s t r i b u t i o n of the sample taken. A l l tend to agree on the marten's dependence on small mammals, although i n some instances s q u i r r e l s seemed to a t t a i n importance. However, i n one i n s t a n c e , M a r s h a l l (1951b) l a t e r thought that the importance of s q u i r r e l s as marten food was "overemphasised". Quick (1955) found one marten l i v i n g on hares which were l o c a l l y abundant. I t i s g e n e r a l l y agreed that marten food h a b i t s are governed c h i e f l y by a v a i l a b i l i t y of the various foods (Lensink e t . a l . 1955). These l a s t w r i t e r s note "the tremendously important p s y c h o l o g i c a l r o l e t h a t food g e t t i n g has i n the marten's a c t i v i t i e s " and b e l i e v e t h a t marten movements are food c o n t r o l l e d . Marten are found remaining f o r s e v e r a l days i n s m a l l areas, o f t e n around l u s h b e r r y patches or choice c a r r i o n . M a r s h a l l (1951b) concluded that a v a i l a b i l i t y of food probably i s the most important f a c t o r a f f e c t i n g d a i l y range. Although t h i s may w e l l be true i n w i n t e r , the w r i t e r doubts t h a t , i n Algonquin at l e a s t , marten have to range the extent they do i n summer j u s t to get food. 67 TABLE XX: Analyses of Marten Pood - Percent Frequency of Occurrence Alaska B.C. ## B.C. ## Washington ## Lensink e t . Cowan and Quick, 1955 Newby, 1951 a l . 1955 MacKay 1950 Summer Winter Summer Winter Winter Winter Summer Win1 Scats 374 28 112 85 78 17 D i g e s t i v e t r a c t s 64 3 1 16 Items 469 107 202 104 127 -Shrews 8 6 Mice and voles 74 68 59 80 40 56 13 27 S q u i r r e l s and chipmunks 2 2 1 13 10 41 Hares 1 2 5 11 5 3 Ca r r i o n 1 B i r d s 8 19 6 1 54 13 3 8 Insects 8 55 B e r r i e s 18 9 6 4 8 3 Other 5 5 Montana Montana ## Ontario M a r s h a l l Thompson ( t h i s study) 1946 1949 Winter Winter Winter Summer Winter and e a r l y s p r i n g Scats 64 18 46 745 144 D i g e s t i v e t r a c t s Items 77 20 57 1427 191 Shrews 6 20 Mice and voles 14 15 14 30 50 S q u i r r e l s and 12 chipmunks 59 55 60 4 Hares 9 20 5 1 5 C a r r i o n 5 5 1 7 Bi r d s 7 5 8 14 1 Insects 1 2 9 2 B e r r i e s 27 Other . 10 7 8 4 data taken to nearest whole percent 68 Hawley and Newby (1957) noted a drop i n s m a l l mammal abundance preceding a r e d u c t i o n of a high marten p o p u l a t i o n ; i t was concluded that food was probably a l i m i t i n g f a c t o r under such c o n d i t i o n s . I t i s r e c a l l e d t h a t the extent of i n d i v i d u a l marten's ranges I n t h i s h i g h p o p u l a t i o n was e s s e n t i a l l y the same as those i n Algonquin. Management I m p l i c a t i o n s : A decrease i n marten abundance i s g e n e r a l l y a t t r i b u t e d to o ver-trapping or unfavourable h a b i t a t manipulation. Marten are considered to be a climax or near climax f o r e s t species ( M i l l e r e t . a l . 1955), and appear to su r v i v e adequately i f the f o r e s t remains t h a t way. Yeager (1950) has shown the continuing abundance of marten i n B r i t i s h Columbia and Yukon as contrasted to considerable reductions i n eastern Canada. I t has been noted i n s e v e r a l instances (e.g. W i l l i a m s , 1947) that marten are l e f t at higher a l t i t u d e s i n montane areas; these inaccessable areas cannot be logged or h e a v i l y trapped whereas eastern areas are qui t e a c c e s s i b l e to both. The exceedingly wide range o f mature f o r e s t types i n which marten are found s u f f i c i e n t l y i n d i c a t e s t h a t f l o r a l c o n s t i t u e n t s have l i t t l e importance, but as i n d i c a t e d i n the present study, s u f f i c i e n t s h e l t e r may. Yeager ( l o c . c i t . ) notes that continued p r o d u c t i v i t y of marten occurred i n areas o f Colorado which were grazed by sheep. The Algonquin observations come from an area of extreme d i v e r s i t y of f o r e s t s which are so s i t u a t e d to form an extensive mosaic of small stands. In areas of the province covered by vast t r a c t s of even age stands of one 69 s p e c i e s , e.g. jack p i n e , marten may show c o n s i s t e n t and marked f o r e s t type preference. In O n t a r i o , much marten range i s under timber management. The e f f e c t t h i s has, i f any, on marten p r o d u c t i v i t y has yet to be evaluated. Marten are a r e l a t i v e l y l o n g - l i v e d and slow reproducing fur-bearer so that p o p u l a t i o n turnover may be comparatively low. Females must s u r v i v e two trapping seasons before they produce young, and a f t e r m a t u r i t y they do not n e c e s s a r i l y l i t t e r every year. This has been noted f o r captives (Hodgson, 1956) and i n females observed during the present study. L i t t e r s average only three i n number (Yerbury, 1947, Hodgson, op. c i t . ) . Hence, i f an area does a c t u a l l y get h e a v i l y over-trapped or otherwise decimated, r e p o p u l a t i o n of i t may be qu i t e slow. The i n d i c a t e d two marten per square mi l e r e s i d e n t p o p u l a t i o n would be a u s e f u l base f o r s e t t i n g t r a p quotas only i f a r e l i a b l e census technique f o r marten could be dis c o v e r e d , to r e v e a l the excess. Such has not been devised; frequency of occurrence of tracks cannot be r e l i e d on. I d e a l l y , t r a p p i n g should o n l y take surplus animals but i n d i c a t i o n s are that t r a p - l i n e s normally do t h i s (e.g. Quick, 1956). Areas between l i n e s s e r v e as reserves. Lensink (1953) noted, again i d e a l l y , that a marten refuge could be s m a l l , w i t h a diameter at l e a s t twice the " c r u i s i n g r a d i u s " of the species (up to f i f t e e n m i l e s ) . The e f f e c t i v e radius of d i s p e r s a l i s then considered to be up to t h i r t y m i l e s , hence refuges should not be separated by a dis t a n c e more than twice t h i s . 70 In areas of Ontario which have been e x t e n s i v e l y burned, or logged to the extent t h a t most of the f o r e s t i s i n e a r l y s e r a i stage, some c o n s i d e r a t i o n could be given to r e s e r v i n g remanent smaller d i s j u n c t areas of mature f o r e s t f o r marten, i f enforcement problems were not too great. 71 L i t e r a t u r e C i t e d : Anonymous. 1936. R a i s i n g martens i n c a p t i v i t y . W i l d l i f e L e a f l e t No. 63, U.S. Department of the I n t e r i o r . Anonymous. 1952. Marten r a i s i n g . Pur Trade J o u r n a l of Canada, 30214- October. Ashbrook, F.G. and K.B. Hanson. 1927. Breeding marten i n c a p t i v i t y . J . o f H e r e d i t y , 18(11):499-503. Brassard, J.A. and R. Bernard. 1939. 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A c t i v i t y of the S i e r r a pine marten. J . of Mammalogy, 26:435. Hawley, V.D. and P.E. Newby. 1957. Marten home ranges and p o p u l a t i o n f l u c t u a t i o n s . J . of Mammalogy, 38:174-184. Hayne, D.W. 1949. C a l c u l a t i o n of s i z e of home range. J . o f Mammalogy, 30:1-18. Higgins , R.J. 1948. Lords of the t r e e t o p s . Fauna, 10:43-45. Hodgson, R.G. 1956. The R i t c h i e martens. Who's Who i n f u r s , 7:20-24. J a r v i s , J.M. 1956. An e c o l o g i c a l approach to t o l e r a n t hardwood s i l v i c u l t u r e . Can. Dept. Northern A f f a i r s and Nat. Resources - Forest Res. Div. Tech. Note No. 43. Lensink, C.J. 1953. An i n v e s t i g a t i o n of the marten i n i n t e r i o r A l a s k a . Unpublished M.S. t h e s i s . U n i v e r s i t y of Alaska. Lensink, C.J., R.O. Skoog and J.L. Buckley. 1955. Food h a b i t s of marten i n i n t e r i o r Alaska and t h e i r s i g n i f i c a n c e . J . o f W i l d l i f e Management, 19:364-368. L l e w e l l y n , L.M. 1953. Tips on t r a p p i n g , tagging and handling s m a l l game and f u r animals. Paper presented to the Northeastern Section of the W i l d l i f e S o c i e t y . September, 1953. Loucks, M.R. 1955. Report f o r the Gogama D i s t r i c t - 1955 Sudbury Fur Advisory Committee. Ontario Dept. of Lands and F o r e s t s . A p r i l . MacFie, J.A. 1954. Notes on the b i r d s , mammals and f i s h of extreme northwestern Ontario. F i s h and W i l d l i f e Management Report, Ontario Dept. of Lands and F o r e s t s , No. 16. Markley, M.H. and C.F. Ba s s e t t . 1952. Habits of captive marten. Am. Mid. N a t u r a l i s t , 28:604-616. M a r s h a l l , W.H. 1946. Winter food h a b i t s of the pine marten i n Montana. J . o f Mammalogy, 27:83-84. 1951. An age determination method f o r the pine marten. J . of W i l d l i f e Management, 15:276-283. 73 1951b. Pine marten as a f o r e s t product. J . of F o r e s t r y , 49:899-905. M i l l e r , R.G., R . W . R i t c e y and R.Y. Edwards. 1955. L i v e -trapping marten i n B r i t i s h Columbia. The M u r r e l e t , 36:1-8. Murie, C.J. 1954. A f i e l d guide to animal t r a c k s . Boston. Newby, F.E. 1951. Ecology of the marten i n the Twin Lakes area. Chelan County, Washington. Unpublished MS t h e s i s . State College of Washington. ( C i t e d by Lensink, 1953). Newby, F.E, and V.D. Hawley. 1954. Progress on a marten l i v e -t rapping study. Trans. 19th N. Am. W i l d l i f e Conference. 452-460. Orsborn, E.V. 1953. More on marten r a i s i n g . Fur Trade J o u r n a l of Canada, 31:14- December. Quick,;, H.F. 1955. Food h a b i t s of marten (Martes americana) i n northern B r i t i s h Columbia. Can. F i e l d N a t u r a l i s t , 69:144-147. 1956. E f f e c t s of e x p l o i t a t i o n on a marten p o p u l a t i o n . J . of W i l d l i f e Management, 20:267-274. Remington, J.D. 1950. Ecology and economics of the Rocky Mountain marten. Quart, r e p o r t . Colorado Coop. W i l d l i f e Research U n i t , 3:21-27. 1952. Food h a b i t s , growth and behaviour of two captive pine marten. J . of Mammalogy, 33:66-70. R i t c h i e , J.W. 1953.. Marten r a i s i n g f o r twenty-four years. Fur Trade Journal of Canada. .30:10- March. Schmidt, F. 1943. Naturgeschichte des baum und des steinmarders. Mono. Der Wildsaugeterre Band X:1-258. Spurr, H.S. 1948. A e r i a l Photographs i n F o r e s t r y . Ronald Co.N.Y. S t i c k e l , L.F. 1948. The t r a p l i n e as a measure of mammal pop u l a t i o n s . J . of W i l d l i f e Management, 12:153-161. Thomas, E.M. 1952.. The f u r bearing mammals of Wyoming. Wyoming W i l d l i f e , 16:12-17. Thompson, W.K. 1949. A study of marten i n Montana. Proc. Ann. Conf. Western Assn. State Game and F i s h Comm. 29:181-188. 74 de Vos, A. 1951. Tracking of f i s h e r and marten. S y l v a , 7:15-20. 1952. Ecology and management o f f i s h e r and marten i n Ontario. Technical B u l l e t i n , Ontario Dept. of Lands and F o r e s t s . de Vos, A. and S.E. G-eunther. 1952. P r e l i m i n a r y l i v e - t r a p p i n g s t u d i e s of marten. J . of W i l d l i f e Management, 16(2) :207-2l4. Walker, E.P. 1929. Evidence of the g e s t a t i o n p e r i o d of martens. J . of Mammalogy, 10:206-209. W i l l i a m s , CM. 1947. Marten i n Colorado". Colorado Conservation Comments, 10:12-13. W i l l i a m s , T.R. 1957. Marten and hawk harass snowshoe hare. J . of Mammalogy, 38(4):517-518. W i l l i a m s o n , V.H.H. 1951. Determination of h a i r s by impressions. J. o f Mammalogy, 32:80-84. Yeager, L.E. 1950. I m p l i c a t i o n s on some harvest and h a b i t a t f a c t o r s on pine marten management. Trans. 15th N. Am. W i l d l i f e Conference:319-334. ^eager, L.E. and J.D. Remington. 1956. Sight observations of Colorado martens. J . of Mammalogy, 37(4):52l-524. Yerbury, H. 1947. R a i s i n g martens i n c a p t i v i t y . Fur Trade J o u r n a l of Canada, 25: September. GRAPH 1: V a r i a t i o n s i n Seasonal A c t i v i t y of Marten as Indicated by Observed versus Expected Captures. 40 ra <D U 0 -P ft COi o o 0 1956 J Ont. standard deviaf ion Observed May 16-31 June 1-15 June 16-30 J u l y 1-15 J u l y 16-31 August 1-15 Augus t 16-31 1957 40 © ? •p ft OS o t i o 20 o a-May June June J u l y J u l y August August 16-31 1-15 16-30 1-15 16-31 1-15 16-31 GRAPH 2: Winter Marten A c t i v i t y Related to Snow Depth i n Major Porest Types. GRAPHS 3 to 14: Seasonal V a r i a t i o n i n Major Pood types of Marten. 1955 4-July ftujuSt 1956 SO<f (terns - Sca+S ko Za • Matj J ci rye. J u l y August Small mammals Co f o Zo 4 1957 L H-H iie**s - 3*f9 SOTS n 5 u n e J u l » j A u g u s t 4-Ju \y flu ^ U5t fee u o o 4-4, Jo-n J u I u flugyst n Moij June Ju/u Robust toOn H-o So B i r d s 10 M o i j J u n e J u l w flujL<s + Insects l-o Zo-8 n M a i < J u n e J u l t j flujust 4o 11 o l n , n n ,11 n May June, J u l ^ A u g u s t 6o. 12 401 13 14 4-S +-4- Zoi. Z0--H-o. ZO • J u l i j fluju6t M c * y J u n e . Ju\y fluji;st B e r r i e s n I I M<*K J u n e Ju|i) ftoausf GRAPH 15: Small Mammals Snap-trapped and Small Mammals Eaten by Marten'. Sorex spp. and B l a r i n a Clethrionomys Microtus Synaptomys Peromyscus ITapaeozapus Zapus 20 40 % composition i n marten sea snap-trapped 60 MAP 1: Marten Study Area and Small Mammal T r a p - l i n e s • l . l o c a t i o n - o f l i v e - t r a p — arbitrary""bounda.ry of study area • l o c a t i o n of small mammal trap l i n e s , 1956. i MAP 2: Minimum Foraging Ranges, Male No. 55, o l d a d u l t P R O J E C T : MARTEN ECOLOGY [ S T U D Y A R E A ] WILDLIFE RESEARCH A R E A , A L G O N Q U I N PARK O N T . • captures, 1955 © capture, Feb. 8, 1956 range, May 18-June 28, 1956 range, July' 7-Aug. 10, 1956 range, Aug. 11-Aug. 23, 1956 MAP 3: Minimum Foraging Ranges, Male Ho. 58, adu l t • captures, 1955 * captures, 1957 range, May 18-June 27, 1956 - range, J u l y 5-Aug. 1, 1956 range, Aug. 4-Aug. 19, 1956 range, Aug. 20-Aug.30, 1956 4: Minimum Foraging Ranges, Female No. 37, a d u l t MAP 5: Minimum Foraging Range, Female No. 41, adult MAP 6: D i s t r i b u t i o n of Male Marten Showing Range Overlap - 1956 — No. 40 adul t • (i) No. 73, j u v e n i l e ---.Ho. 55 adul t • U) No. 69, adul t No. 58 .old a d u l t • (3) No. 70, adul t No. 67 old a d u l t MAP 7: D i s t r i b u t i o n of Male Marten Showing Range Overlap - 1957 No. 40 a d u l t No. 55 a d u l t No. 67 o l d a d u l t No. 76 a d u l t -. :— No. 86 j u v e n i l e No. 88 " y e a r l i n g " — No. 91 " y e a r l i n g " • d) Ho. 70 a d u l t • U) No. 77 o l d a d u l t • O) No. 82 adu l t • wNo. 87 j u v e n i l e • 00 No. 89 j u v e n i l e • WNo. 90 " y e a r l i n g " • (?)No. 93 " y e a r l i n g " M P 8: D i s t r i b u t i o n of Female Marten - J u l y , August 1955 and May 1956. Ho. 4 1 , a d u l t l i m i t of trapping area No. 64, adul t © No. 95, a d u l t No. 61, imm. • No. 63, immature MAP 9: D i s t r i b u t i o n of Female Marten - June to August 1956 P R O J E C T : MARTEN ECOLOGY [ S T U D Y AREA] WILDLIFE RESEARCH A R E A , A L G O N Q U I N PARK O N T . 0 a d u l t : Bos. 37, 65, 66, 68, 71. immature: Bos. 72, 74, 75. ! Ma) No. 65 -captures before and a f t e r appearance of No. 7] • U) No. 72 • (*)No. 75 <3) No. 74 MAP 10: D i s t r i b u t i o n of Female Marten - May to August 1957 P R O J E C T : MARTEN ECOLOGY [STUDY AREA] W I L D L I F E RESEARCH A R E A , A L G O N Q U I N PARK O N T . a d u l t : Nos. 37, 65, 66, 6.8, 78. .immature: Eos. 85, 92, 9-4. • U) Ho. 78 • U) No. 92 • U) No. 85 • (*) No. 94 P i g . 1. Winter a e r i a l photograph of a p o r t i o n of the study area. C o n t r a s t i n g c o n i f e r and hardwood f o r e s t s can be seen; two hemlock r i d g e s show prominently. P i g . 2. Spring a e r i a l photograph of a p o r t i o n of the study area. Fig. 3 . -Component Forest Types 1) Hemlock 2) White pine, red pine 3) Black spruce 4) Balsam, white spruce, "black spruce, white pine, hemlock j?) White spruce, white pine, balsam, white birch, aspen 6) Hemlock, sugar maple, yellow birch 7) White birch, aspen, white spruce, balsam, white pine 8) Sugar maple, yellow birch, hemlock 9) Sugar maple 10) White birch b) Decadent stands P i g . 4. View over a p o r t i o n of the study area i n the foreground i n d i c a t i n g the r o l l i n g topography of the r e g i o n . P i g . 6. Hemlock f o r e s t - summer. P i g . 8 . White b i r c h , white spruce and pine f o r e s t - summer. P i g . 9 . Sugar maple, yellow b i r c h , hemlock f o r e s t type - w i n t e r . P i g . 11. Sugar maple f o r e s t - winter P i g . 12. Marten l i v e - t r a p , set by l o g . Photo, by Mr. D. Johnston. P i g . 13. Marten i n a l i v e t r a p . Photo, by Mr. M. D a n i e l . P i g . 14. L o c a t i o n of a marten nest den. A p o r t i o n of the l o g was removed and the remainder s p l i t open. Photo, by Mr. D. Johnston. P i g . 15. L o c a t i o n of a winter day den among boulders. A f o o t r u l e i n the foreground i n d i c a t e s the entrance used by a marten. 

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