A PJEPHODUOTIOK STUDY OF 1TGRSHER1T WHITE CEDAR By Thomas Charles Kelson A THESIS Subnitted to the School of Graduate Studiiie s o f Michigan State C ollege o f A gricu lture and Applied Science in p a r t ia l f u lf illm e n t o f the requirements fo r the degree o f DOCTOR OF PHILOSOPHY Department o f F orestry 1950 i ACKNOWLEDGEMENTS ERRATA The key i n f ig u r e s 1 , 2, and 3 should read "ten year record" in ste a d o f "nine y ea r record". Management S p e c ia lis t , were aeepxy Kppx-wcxetvou..-------Dr. L. E. G ysel o f Michigan S ta te C o lleg e la r g e ly d ir e c te d th e stu d ie s and gave in v a lu a b le su g g estio n s on th e w r itin g o f th is manu­ s c r ip t . Dr. P. A. H erbert, Dr. W. B. Drew and Mr. J. 0. Veatch o f Michigan S ta te C o lleg e are thanked f o r t h e ir p a r t ic ip a t io n upon th e Graduate Committee. Cusino V /ild life Experiment S ta tio n , S h in gleton , fu rn ish ed f i e l d h elp . E sp ecia l c o n sid e r a tio n i s due Mr. W. J. Koski f o r h i s many weeks o f f i e l d a id , h is c h e er fu l a t t it u d e and h is p erson al in t e r e s t in the p r o je c t. Mr. D. P. Sw itzenberg took the photographs w ith th e excep tion o f those in which mention o f th e photographer i s made in th e cap tion . E x c e lle n t co o p era tio n was r eceiv ed from o th er in te r e s te d a g en cies and o rg a n iza tio n s in c lu d in g Mr. W. M. Z i l l g i t t , in charge o f th e Upper P en in su la Experim ental F o r e st, Lake S ta te s F orest Experiment S ta tio n , Marquette, Mr. Gene A. H esterberg, Department o f F o r estry , Michigan C ollege o f Mining and Technology, and th e s t a f f o f th e F orestry D iv is io n , _____ A p o r tio n of the a id was made p o s s ib le through a Federal Aid in W ild life R esto ra tio n P r o je c t, 52-R. ACIQTOWLEDGEMENTS G rateful acknowledgement i s made to the Michigan Department o f Conservation f o r u ro v id in g th e p a rt-tim e fe llo w s h ip and oth er a id and 1/ fa c ilitie s which h elp ed to make t h i s study p o s s ib le . The encourage­ ment and a d v ice o f Mr. H. D. Ruhl, C hief o f th e Game D iv is io n , Mr. F. H. D a le, form erly Pittm an-R obertson Coordinator, Mr. L. A. Davenport, the p resen t Pittm an-R obertson C oordinator, and Mr. B. C. Jen k in s, Cover Management S p e c i a l i s t , were deeply a p p reciated . Dr. L. E. Gysel o f Michigan S ta te C o lleg e la r g e ly d ir e c te d the stu d ie s and gave in v a lu a b le su g g e stio n s on the w r itin g o f th is manius c r ip t . Dr. P. A. H erbert, Dr. W. B, Drew and Mr. J. 0. Veatch o f Michigan S ta te C o lleg e a re thanked f o r t h e ir p a r t ic ip a t io n upon th e Graduate Committee. Cusino W ild life Experiment S ta tio n , S h in gleton , fu rn ish ed f i e l d help. E sp ecia l c o n sid e ra tio n i s due Mr. W. J. Koski fo r h is many weeks o f f i e l d a id , h is ch eerfu l a t t it u d e and h i s p erson al in t e r e s t in th e p r o je c t. Mr. D. P. Sw itzenberg took the photographs w ith the exception o f th o se in which mention o f th e photographer i s made in th e caption. E x c e llen t coop eration v/as r e c e iv e d from o th er in te r e s t e d a g en cies and o r g a n iza tio n s in c lu d in g Mr. W. M. Z i l l g i t t , in charge o f th e Upper P enin su la Experim ental F o r e st, Lake S ta te s F orest Experiment S ta tio n , Marquette, Mr. Gene A. H esterberg, Department o f F o restry , Michigan C ollege o f Mining and Technology, and th e s t a f f o f th e F orestry D iv isio n , If A p o rtio n o f th e a id was made p o s s ib le through a Federal Aid in W ild life R esto ra tio n P r o je c t, 52-R. Michigan Department o f C onservation. F in a lly , and not in an attem pt to "be f a c e t io u s , s in c e r e thanks are due to th e Carbide and Carbon Chemicals Corporation, the makers o f ll6 l2 " , w ithout which most o f the f i e l d work would have been unbearable. With th e m osquitoes and b la c k f li e s p a r t ia l l y under c o n tr o l, the days spent wading knee-deep i n swamp-water, th e days w ith dew on th e w a ist-h ig h brush u n t i l noon, the warm days when the "swarap-gas11 crea ted a c o n sid e r a b le odor, and th e J u ly and August days in an a ld e r th ic k e t a r e now th e su b ject o f n o s t a lg ic memories rath er than b it t e r c u ssin g . TABLE OF CONTENTS Page THE PROBLIM............................................................................................................ 7 REVIEW OP LITERATURE..................................................................................... 8 DESCRIPTION OP AREA. P h y sio g r a p h y .............................................................................................. 11 C l i m a t e ....................................................................................................... 11 THE SPECIES AND ITS ENVIRONMENT The S e e d ....................................................................................................... 12 The T r e e ....................................................................................................... 16 Cover Types and A s s o c i a t i o n s .......................................................... 18 S o ils 20 . . ................................................................................................... INITIAL REPRODUCTION S eed lin g C h a r a c te r istic s and Development ........................................................................ Methods . . . . . R e s u l t s .............................................................................................. 24 24 S u rvival and M o rta lity M eth o d s.............................................................................................. R e s u l t s .............................................................................................. 30 33 Types o f Reproduction M eth o d s.............................................................................................. R e s u l t s .............................................................................................. 41 42 Environmental F actors A ffe c tin g I n i t i a l Reproduction Laboratory S tu d ies Seed V i a b i l i t y ................................................................... L i g h t ..................................................................................... A c i d i t y ................................................................................ Type o f Germinating M e d i a ........................................ T e m p e r a tu r e ....................................................................... F ie ld S tu d ies Season o f Seed Germination . ................................ E f f e c t iv e Seeding D istan ce Methods . . . . . ................................................. R e s u l t s .................. .................................................... 51 52 53 53 58 60 60 62 L igh t Methods ................................. . R e su lts ......................................... A c id ity Methods . .................................... R e su lts ......................................... Types o f Germinating Media Methods ......................................... R e s u l t s ......................................... Temperature M eth o d s........................................ R e su lts ......................................... S o il M oisture Methods .................. R e su lts ........................................ Drainage Methods ........................................ , R e s u l t s ........................................ ... Rodent and Snov/shoe Hare Browsing M eth o d s............................................, R esu lts ......................................... . S o il and Cover Types Methods S o il T y p e s ...................... ... Cover T y p e s .................. ... R esu lts S o il T y p e s .......................... Cover T y p e s .................. ... Age o f Stand M eth o d s............................................ , R e su lts ............................................ ADVANCED REPRODUCTION urvival and M ortality Methods ................. R esu lts ................. E ffect o f Deer and Snov/shoe Hare Browsing upon Advanced R«^production and Stand Composition Methods ........................................................................................ R esults ........................................................................................ E ffec t o f Deer Browsing upon Advanced Reproduction and S;and Composition Methods .................................................................................... R esults Page SILVICULTURAL PRACTICES E ffe c t o f C ertain S ilv ic u lt u r a l P r a c tic e s upon th e Reproduction o f Cedar and i t s A s so c ia te s Methods Boh* s Lake C u ttings . ........................................................... 150 Cusino C u t t i n g s ............................................................................. 156 R esu lts Bob»s Lake C u t t i n g s .................................................................... 159 Cusino C u t t i n g s ............................................................................. 165 P la n ta tio n s M eth o d s....................................................................... 169 R e s u l t s ............................................................................................................ 172 DISCUSSI01T R ep ro d u ctio n ............................................................................................................ 173 Management.......................... 178 SUMMARY ADD CONCLUSIONS...............................................................................................182 LITERATURE C IT E D ............................................................................................................ 188 APPENDIXES......................................................................................................................... 193 Appendix A. T ab les and M athematical C o n f u ta t io n s ............................ 194Appendix B. Common Names o f Trees and Shrubs and Their S c i e n t i f i c E q u iv a le n t s ........................................................... 199 Appendix C. Common Names o f B ird s and Mammals and Their S c i e n t i f i c E quivalents ...................................................... 200 Appendix D. Method o f Deer Browse A n a l y s i s ..........................................201 Appendix E. A s s o c ia te s o f Cedar from C o lle c tio n s and O bservations ............................................................................ 205 THE PROBLEM Northern w h ite cedar (Thuja o c c id e n t a lis L . ) stands occupy approxim ately 639*000 a c re s in M ichigan, Ahout 293*000 acres o f t h is t o t a l i s found in th e e a ste r n h a lf o f th e Upper P en in su la o f Michigan (3 5 ). Cedar i s th e s ta p le w in ter food o f th e w h it e -t a ile d deer ( O docoileus v ir g in !a n u s b o r e a lis M ille r ) , I t i s th e c h ie f source o f p o le s and p o s ts in th e Lake S ta te s as w e ll as being an important source o f o th er tim ber p rod u cts. Northern w h ite cedar has n e g a tiv e valu e in th e e x te n siv e stands managed fo r pulp p rod uction . Pewer e c o lo g ic a l and s i l v i c u l t u r a l s tu d ie s have been made on t h is s p e c ie s than on any o th er commercial t r e e s p e c ie s in th e reg io n . None have been made th at would serve to in d ic a te p o s s ib le management p r a c t ic e s . I t i s w ith th e se f a c t s in mind th at a study was i n i t i a t e d concerning th e f a c to r s a f f e c t in g i t s reproduction in swamp stan d s in the Upper P e n in su la . REVIEW OF LITERATURE L ite r a tu r e d ir e c t ly r e la t e d to northern w h ite cedar reproduct­ ion i s lim ite d . Papers d ir e c t ly concerned w ith any phase o f the management o f th e s p e c ie s a re not numerous. However, "bog s tu d ie s , reproduction s tu d ie s o f o th er s p e c ie s , and s tu d ie s in v o lv in g e c o lo g ic a l fa c to r s d e a lt w ith in t h is d is s e r t a t io n are m u ltitu d in o u s. S tu d ies on northern w hite cedar reprod uction are lim it e d to th e s t a t e s o f Maine and M innesota. C u rtis ( l 6) made a g en era l study o f th e s p e c ie s in clu d in g reproduction — p rim a r ily in swamp stan d s ch aracter­ iz e d "by h igh water ta b le s throughout th e y ea r. He a ls o conducted a sim ila r study on th e s p e c ie s in upland stand s (1 5 ). ca rried on fu r th er s tu d ie s o f rep rod u ction in Maine. Montgomery (i+2) Maki ( h i) stu d ied growth and reproduction in M innesota sta n d s. Stewart (5 0 ), in e f f e c t , c l a s s i f i e s northern w h ite cedar in a s in g le cover type in th e Lake S t a te s . Cedar i s lim it e d in th e type o f s i t e th a t i t o ccu p ies in th e southern p o r tio n s o f M ichigan, p re­ dominantly th o se s i t e s w ith a R i f le p eat s o i l typ e (5 5 )• ®ke s i t e p referen ce becomes l e s s marked as one p r o g r e sse s northward and westward. The su c c e ssio n o f cedar and i t s a s s o c ia t e s has been the su b ject o f co n sid era b le study. Gates (2h) c l a s s i f i e s th e Thuja a s s o c ia tio n as a clim ax type in th e hydrarch s e r ie s in bog la k e s in th e Douglas Lake region o f M ichigan. In wet s o i l s , he p la c e s Thuja as a temporary typ e, preceded by a bog a s s o c ia t io n or a low land f o r e s t . I t i s a s s o c ia te d w ith and rep laced by a P ic e a - A bies a s s o c ia tio n . Coouer ( l h ) , in d esc r ib in g an area e v o lv in g from Keweenawan la v a flo w s ( I s l e R o y a le), d if f e r e n t ia t e s between a bog su c c e ssio n and a d e lta swamp s u c c e ss io n . Thuja does not en ter in to th e hog su c ce ssio n except as a subdominant to l a r i x , a subclim ax s ta g e o f one l i n e o f development. On th e o th er hand, he p la c e s Thuja a s th e dominant in a s im ila r sta g e in th e d e lt a swamp su c c e ss io n . For the most p a r t, cedar seed and s e e d lin g s tu d ie s have "been aimed a t determ ining th e e f f e c t o f environmental fa c to r s on germ ination rath er than b ein g c a r r ie d out s p e c if i c a l l y a s s p e c ie s s tu d ie s . H eit and E lia so n (30) have lim ite d th e ir s tu d ie s to co n ife ro u s t r e e seed in c lu d in g a s p e c if i c study o f Thuja o c c id e n t a l^ regarding methods o f seed t e s t in g and fa c to r s a f f e c t in g germ ination and seed q u a lity . There i s no apparent marked d iffe r e n c e in the subterranean organs o f a g iven s p e c ie s growing in a bog and in comparable con d i­ tio n s in m ineral s o i l s (1 9 ). However, Rigg and Harrar (47) contend that e x c e s siv e root fu sio n o fte n occurs in sphagnum and e c c e n t r ic it y o f root growth and enlargement i s c h a r a c t e r is t ic o f t r e e r o o ts growing in sphagnum p ea t. Moore (43) m ain tains th at northern w h ite cedar i s th e o n ly n orth eastern c o n ife r th a t does not have m yccorhizal fu n g i in or on i t s r o o ts. The e f f e c t o f l i g h t and pH on cedar reproduction a r e m entioned in th e l i t e r a t u r e . Wiesner (60) and Burns ( 9 ) , u sin g photographic paper, c a lc u la te d th e average minimum li g h t in t e n s it y requirem ent fo r cedar, lu t z and Chandler (JjO) p o in t out th at northern w h ite cedar has been observed commonly on calcareou s s o i l s and i s sc a rce or absent on ad jacen t a c id s o i l s . Wherry (59) d isc u s se s th e re­ a c tio n ^ referen ce o f ea stern c o n ife r s in clu d in g cedar. I t was found n ec essa r y to turn to th e lit e r a t u r e fo r methods o f c la s s if y in g and determ ining th e e f f e c t s o f the va rio u s ty p es o f germ inating media found on th e swamp f lo o r (7 , 22, 26, ^ 5). McCullough (hij.) has developed a system o f c l a s s i f i c a t i o n fo r the v a rio u s sta g e s o f decay in lo g s on the w estern s p r u c e -fir f o r e s t s fo r u se in r e la t io n to t h e ir a c c e p t i b il i t y as ty p es o f germ inating media. H is c l a s s i f i ­ c a tio n was used in t h is stu d y. Son and A v er ell (6 2 ), stu d yin g growth in r e la t io n to swamp drainage, conclude th a t growth r a te d ecreases in p rop ortion to th e d ista n c e from the source o f d rain age. LeBarron and N e e tz e l (38) t a l l i e d sample quadrats over a lo n g p erio d o f tim e a t d esig n a ted d ista n c e s from a r t i f i c i a l arainageways in a cedar stand and found that e f f e c t i v e d rain in g would change a hog c o n ife r type to a hardwood c o n ife r type. Northern w hite cedar i s an important component o f many d ee r yards throughout the Lake S ta te s and i s su b ject to heavy browsing "by sev era l mammals, B a r t le t t ( 6 ) , S w ift (5 1 ), and Aldous and Smith (^-) d isc u ss the winter deer food problem in Michigan, W isconsin, and Minnesota r e s p e c tiv e ly . Davenport (17) p o in ts out th at northern w hite cedar proved to be th e o n ly s p e c ie s o f browse fe d e x c lu s iv e ly upon which deer were a b le to m aintain t h e ir stren g th . S w ift (52) rep orts th at in northern W isconsin, only th ree out o f every ten young cedar remain undamaged from d eer or snowshoe hare browsing. The harmful in flu e n c e o f snowshoe hare browsing on p la n ta tio n s in th e Lake S ta te s lias been d isc u sse d ( l ) . Cedar i s commonly damaged by snowshoe hare browsing, e s p e c ia lly when other food i s sca rce (3 6 ). Krauch (33, 3h) rep orts th at rod en ts are h ig h ly in im ic a l to n atu ral co n ifero u s regen eration in se v e r a l reg io n s. Cedar i s one o f the few c o n if e r s th a t reproduces v e g e t a t iv e ly . P otzger (k6) acknowledges th e f a c t th a t v e g e ta tiv e p ropagation occurs in c o n ife r s . Harlow (27) found o c c a sio n a l reju ven ation o f T, o c c id e n t a lis by n atu ral c u ttin g s in th e Adirondacks. Cooper (13) is r e s p o n sib le fo r r e c o g n itio n o f n a tu ra l la y e r in g among c o n if e r s , Aldous (3 ) rep o r ts th a t cedar food and cover p la n tin g s f o r d eer were u n su c ce ssfu l in Minnesota, DESCRIPTION OP AREA Physiography T his study was made in th e Great Lakes p la in s r e g io n , a g la c ia te d p a rt o f th e Great C entral Lowland o f the U n ited S t a t e s , and was co n fin ed to a p o rtio n o f t h i s p rovince in th e e a s te r n p a rt of the Upper P en in su la o f M ichigan. by p a le o z o ic sedim entary rock. The area as a whole i s u n d erlain The la r g e swamps, th e s p e c i f i c areas that th e cedar stu d ie s were made i n , have developed from l e v e l t i l l p la in s or o ld la k e beds. Upland r id g e s in th e v i c i n i t y prob ab ly are the r e s u lt o f g la c ia l a c t i v i t y . The beds dip g e n tly to th e south as the r e s u lt o f a g r ea te r amount o f r ecen t u p l i f t on the Lake Superior shore (5 5 ). Drainage i s poor and haphazard. In the Cusino swamp, the lo c a tio n o f t h is study, Stoner Creek p ro v id es the main drainageway, emptying in to Lake Michigan a t M anistique through th e M an istiaue R iver, Climate The Cusino swamp l i e s approxim ately a t 46°30' North la t it u d e . The c lim a te i s o f a marine type, governed by the fo r c e and d ir e c tio n of th e wind. C lim a tic co n d itio n s w ith in th e study area a re more sim ila r to th ose o f the weather s t a t io n n ear Seney, Michigan than to any o f the o th er nearby s ta tio n s a t which records have been k ept (5*4). A ten -year record shows an average January temperature o f 1*4.9° P. and a Ju ly average o f 6*1.8° P. Maximum temperature record ed a t th is s ta ­ tio n i s 103° E. and the minimum i s -* 4-7 . 0 ° P. The average date fo r th e la s t k i l l i n g f r o s t in the sp rin g i s June 15th and f o r the f i r s t f r o s t in f a l l i s August 27. The average growing season i s 73 days. A 17-year record in d ic a te s th a t average r a i n f a l l i s 2 6 .7 5 in c h e s w ith a growing season range o f 2.89 in ch es during May to 3 .33 in ch es in September. Light f r o s t s have been known to occur during June, J u ly , and August (55). During the 19*49 growing sea so n , f r o s t s were known to occur in the swamp on June 8 and J u ly 18. Climographs have been co n stru cted fo r 19*47. 19^48. and through the ].9*i9 growing season fo r the Seney sta tio n ( fig u r e s 1 , 2, 3 ). A climograph fo r the ten year p e r io d from 1930-39 in c l u s i v e lias been transposed on each y ea rly clim ograph fo r comparison. The 19*4? grow­ ing season s ta r te d out cool w ith th e l a t t e r p o rtio n o f th e season warmer than average. The season was dry throughout. The 19*48 season was dry but th e 19*49 season turned out to be warm and very wet. TEE SPECIES AND ITS EHVIHQHMEMD The Seed The seed s o f northern w h ite cedar are very sm all in s iz e and 1/ lig h t in w eight . They are approxim ately l / 8 in ch in le n g th with There are about 300,000 seed s p er pound (5&). CLIMOGRAPH OF CUSINO WEATHER 1947 1947 RECORD NINE YEAR RECORD V6 UJ t 40° INCHES OF PRECIPITATION F igu re 1 . Climograph o f Cusino w eather f o r 19^7 superimposed upon te n -y e a r com posite clim ograph. Graphs were con stru cted w ith Seney M igratory Waterfowl Refuge readings, th e n ea rest in lan d w eather s t a t io n . CLIMOGRAPH OF CUSINO WEATHER 1948 1948 RECORD NINE YEAR RECORD 70 ° 60 ' Ui 50 ' 40 “ | 30' UJ 2 3 4 5 6 7 INCHES OF PRECIPITATION Figure 2 . Climograph o f Cusino weather fo r 19*f8 superimposed upon te n -y e a r com posite clim ograph. Graphs were c o n stru cted w ith Seney M igratory Waterfowl Refuge read in gs, the n ea rest in la n d weather s ta tio n . CLIMOGRAPH OF CUSINO WEATHER 1949 1 949 RECORD NINE YEAR RECORD 70 ' 60 ' /- ■ /- UJ 111 S' 30' 320' 4 7 IN C HES OF PRECIPITATION Figure 3 . Climograph o f Cusino weather f o r th e f i r s t eig h t nonthB o f 19^+9 superimposed upon a ten -y ea r com posite clim ograph. Graphs were co n stru cted w ith Seney M igratory Waterfowl Befuge read in gs, the n earest in lan d w eather s t a t io n . a p a ir o f wide wings and brown in c o lo r . High q u a lity seed should con tain roughly 75 p ercen t f i l l e d seed and average p u r ity should a lso be about 75 percent (3 0 ). Seed has been sto r e d in sea led co n ta in ers fo r f iv e years a t 2° to ho C. w ithout any apparent l o s s in i t s o r ig in a l v i a b i l i t y (3 0 ). Seed produotion has been n oted on stems as young as 12 years in age in the Cusino area and cones were found on in d iv id u a ls as young a s 6 years of a g e i n Maine ( l 6 ) . Commencing a t 20 years o f age, seed i s produced q u ite abundantly w ith s u f f ic ie n t production fo r commercial seed g a th e rin g occu rrin g from 30 years o f age to m aturity. Seed occurs alm ost every year but heavy seed crops are borne a t approxim ately 5 year in t e r v a ls . I t i s not uncommon fo r an a v era g e -siz ed tr e e w ith a f a i r l y f u l l crown to produce o f con es. bushel A bushel o f cones averages one pound o f clean seed (h-1). I s o la te d t r e e s , and e s p e c ia lly th ose on upland s i t e s , produce seed more p r o l i f i c a l l y than th e t r e e s on swamp s i t e s in the study area. Seed i s shed throughout th e f a l l months and i s aid ed con sid erab ly in d is tr ib u tio n by c lip p in g o f the branches by s q u ir r e ls . The Tree Northern w hite cedar i s th e o n ly member o f i t s genus Thuja (Thuya S a rg .) th at i s found in the Lake S t a te s . I t commonly grows from hO to 60 f e e t in h eig h t and sev era l f e e t in diam eter. Growth form i s poor in swamp-grown cedar 1/ u; " p is to l-b u tts '1 For a hO fo o t tr e e — a common h eig h t in swamp stands — th at lias a diameter a t b reast h eig h t o u tsid e bark o f 8 .0 in ch es, Gevorkiantz and Duerr (25) c a lc u la te th e diam eter in s id e bark a t 1 fo o t above th e ground as 9 .2 in ch es and 3 .0 in ch es a t 30 f e e t above th e ground. and ''bowed-butts'* are common. C u rtis ( l6 ) p u ts fo r th sev era l hypotheses to exp lain th e se phenomena. V e g eta tiv e reproduction of th e la y e r in g typ e w i l l account fo r curved b u tts . Poor stem anchorage causes stems o f a l l ages to be fo rced from th e v e r t ic a l because o f snow, i c e , or wind a c tio n . S e ed lin g s e s t a b lis h them­ s e lv e s on ro tten lo g s , stumps, and moss hummoclcs th at gradu ally lea n as the su b str a te d is in t e g r a te s and t h e ir weight in c r e a s e s. Upland cedar seems to a t t a in b e tte r form than swamp cedar. Forked- heads and stag-h ead s a re a ls o common in Upper P en in su la swanro stan d s. Height and diam eter growth o f cedar are slow ; Gevorkiantz and Duerr (25) u se l6o y ears as a b ase in determ ining s i t e in d ex. Although o ld stems u su a lly show b u tt r o t , r e l i a b l e r in g counts have been obtained th a t in d ic a te t r e e s 300 y ea rs o ld are p resen t in stands in the Upper P en in su la . l a t e r a l root systems seem to develop on swamp-grown t r e e s , rep la cin g th e tap root o f th e young s e e d lin g s . The root system s are shallow , and th e branch ro o ts a t t a in a red c o lo r a tio n . Northern w hite cedar was one o f the s p e c ie s in clu d ed in th e (,p in e c u t11 in the Lake S t a te s . s h in g le s and p o le s . I t s u se then was p r in c ip a lly fo r Today th e wood i s u sed c h ie f ly f o r p o s ts , p o le s , t i e s , lumber in boat m anufacturing, cab in l o g s , r u s tic fe n c e s and fu r n itu r e , n o v e lty ite m s, and mine la g g in g . Northern w hite cedar f o li a g e i s th e most inroortant w inter browse fo r w h it e -t a ile d deer in th e Lake S ta te s region both from a p a l a t a b i l i t y and a n u tr itio n a l r a tin g ( 2 ) . Although th e amount o f f o lia g e produced by the t o ta l tr e e i s c o r r e la te d w ith diam eter at b rea st h eig h t, the browse a v a ila b le fo r deer d e c lin e s sharply beyond a d .b .h . o f 3 .0 in c h e s .( 2 ) . Cover Types and. A s s o c ia tio n s Stev/art (50) c l a s s i f i e s northern swamps in to a s in g le major type co n ta in in g Thuja o c c id e n t a lis . L arix l a r l c l n a . and P ic e a mariana — th e s i t e c o n d itio n s governing th e stand com position. The "best stan d s o f cedar occur on th e edges o f h igh lan d and swamp where th e p ea t la y e r i s ju s t a few in ch es th ic k . Stewart fu r th e r observed th a t in deep p eat s o i l s , growth o f cedar i s slow and recommends a r o ta tio n on an average s i t e o f 70 to 90 years fo r p o sts and 200 years fo r p o le s . Northern w hite cedar i s in clu d ed in e ig h t o f th e cover types d esign ated hy the S o c ie ty o f American F o r e ste r s (4 9 ), namely; 1 8 . red spruce type 20. paper b irch -r ed spruce-balsam f i r type 21. w hite spruce-balsam fir -p a p e r b ir c h typ e 23. black spruce type type 22. balsam f i r type 2b. northern w h ite cedar typ e 26 . b lack ash-American elm -red maple ty p e. 25. tamarack Of the typ es l i s t e d , o n ly f i v e occur in th e Lake S ta te s , th e w hite spruce-balsam f i r paper b irch typ e, black spruce typ e, northern w h ite cedar ty p e, tamarack ty p e, and the black ash-American elm -red maple typ e. The w hite spruce-balsam fir -p a p e r b irch type does n ot commonly con tain northern w hite cedar and occurs more e x te n s iv e ly in northern Minnesota than in northern Michigan and northern W isconsin. The b lack spruce type i s the t y p ic a l bog cover type th at i s lo c a te d on a c id peat w ith l i t t l e or no drainage. component o f the stand. Cedar i s a minor The type i s u t i l i z e d m ainly fo r pulpwood. XM 1/ The n orthern w hite cedar type , a s e s ta b lish e d by the S o c ie ty o f American F o r e ste r s, i s predom inantly composed o f cedar and i s a s s o c ia te d w ith tamarack, balsam f i r , y e llo w b ir c h , paper b irch , b lack a sh , red maple, b la ck spruce, w hite p in e, hemlock, and a ld e r . The type occurs in th e northern h a lf of th e Lake S ta te s region a s w ell a s in Hew England. I t i s lo c a te d on s it e s w ith slow drainage and a h ig h water ta b le th a t are not stro n g ly a c id . on lim e sto n e uplands. I t i s a ls o p resen t A ccording to i t s type d e s c r ip tio n , i t w i l l m aintain i t s e l f a s lo n g a s the swamp remains wet i f i t i s n o t d i s ­ turbed. The tamarack type in c lu d e s con sid erab le northern w h ite cedar and i s found in M ichigan, W isconsin, and M innesota. I t i s t r u ly a bog ty p e, th e southern stands o f t h i s type occu rrin g as pure tamarack. I t i s succeeded by b lack spruce or ced ar, depending p rim a r ily on the drainage and a c id it y . The b lack ash-American elm -red maple type con tain s cedar o n ly in th e Lake S ta te s and then in small numbers. I t occu p ies low ­ land s i t e s and i s con sid ered a clim ax type. C u rtis (15) prop oses a sep arate cover typ e fo r th e upland northern w h ite cedar th a t d if f e r s from the S o c ie ty o f American F o r e ste r ’ s cedar ty p e, a c l a s s i f i c a t i o n which in c lu d e s both swamp and upland stan d s. The Lake S ta te s F orest Experiment S ta tio n (35) estim ated th a t cedar stands occupy 639,000 a cr e s in Michigan or 3$ o f th e a Synonyms mentioned by th e S o c ie ty o f American F o r e ste r s in c lu d e the fo llo w in g types; cedar-tam arack-spruce, cedar-tam arack-sprucebalsam, cedar-spruce-balsam -w hite p in e , balsam -cedar-taraarack-spruce, cedar-paper birch-balsam -red maple, and mixed swamp. t o t a l f o r e s t land in th e s t a t e . I t i s fu r th e r estim ated that the acreage "by se c tio n s o f th e s t a t e i s as fo llo w s ; S ectio n Acres Eastern h a lf o f the Upper P e n in su la Western h a lf o f the Upper P en in su la Northern h a lf o f the Lower P en in su la Southern h a lf o f the Lower P en in su la 293.000 153.000 180.000 13,000 A sso c ia te s o f cedar th a t \*ere observed and c o lle c t e d are l i s t e d in the Appendix. T h eir occurrence does not n e c e s s a r ily in ­ d ic a te common occurrence on cedar s i t e s , however. P o t e n t llla fru cto sa Por example, was found r a r e ly on cedar swamp s i t e s where­ as Cyprl-pedlum a c a u le , Bubus tr lflo r u m , and Galium asprellum were commonly found. Sarracenia purpurea and Ledum groenlandicum were most abundant on the ty p es w ith cedar th a t approach the tru e bog co n d itio n s. I r i s v e r s ic o lo r and Tyoha l a t l f o l i a were found in th e m oister pock ets w ith in th e swamp. S o ils Although Thu,1a o c c id e n t a ils i s con sid ered to be a c a lc ic o le (2 0 ), i t s occurrence in the Lake S ta te s i s by no means lim ite d to s o i l s o f calcareou s o r ig in . a c id swamps o f th e reg io n . I t i s found e x te n s iv e ly throughout th e Cheney (12) summarized the occurrence o f cedar in swamps by s t a t in g th at cedar comes in only on the old er bogs where the p eat i s f a i r l y w ell decomposed. The area stu d ied l i e s w ith in the Podzol s o i l region o f North America. S o ils are formed from m a teria l d ep osited by the l a t e Wis­ consin g la c ie r and d e p o sits are m ainly o f lo c a l o r ig in . The m ineral s o i l s o f th e area f a l l in to two major groups (55)» s o i l s which have developed under f r e e d rain age and a e r a tio n and con tain a normal amount o f m oisture fo r the r e g io n , and s o i l s which a re permanently or in te r m itte n tly sa tu ra ted w ith w ater. It i s with th e l a t t e r group that our in t e r e s t l i e s alth ou gh cedar does appear on s o i l s o f th e f i r s t group. A g en era lized s o i l p r o f ile w ith poor drain age or e x c e s s iv e m oistu re in clu d es: 1 . a dark-gray or "black su rface la y e r o f organ ic m atter. 2. a gray or dra"b la y e r of sand or sandy loam, s l i g h t l y or not a t a l l colored "by organic m atter. 3 . a la y e r co n ta in in g maximum co n cen tra tio n o f c la y and maximum coherence or p l a s t i c i t y , or one co n ta in ­ ing maximum y ello w or "brown c o lo r a tio n and maxi­ mum cem entation from iro n oxid es. k. the substratum or parent m a teria l, Newton sand i s a s o i l o f the w etter swamps, th e water ta b le being o n ly s l i g h t l y low er than that o f peat sv;amps. The p r o f i l e i s ch a ra cterized by a 2 to 8 inch la y e r o f peat over a sand w ith smokeco lo red or r u s t-y e llo w sp lo tch in g . I t i s s tr o n g ly a c id and covered w ith a v e g e ta tio n common to p ea ts and mucks. Saugatuck sand i s a wet sandy type, w ith a gray, sandy su rface above a ruB t-colored or brownish-black sandy hardpan. r e l i e f i s more or l e s s a pit-and-mound co n d itio n . I t s su rface I t i s a c id and supports a somewhat more mesophytic type of v e g e ta tio n than does Newton sand. Ogemaw f in e sandy loam i s a wet land s o i l ch a ra cterized by a f in e sand or sandy loam over an impervious c la y . The h eig h t o f water ta b le i s l e s s than w ith the Nev/ton and Saugatuck s o i l s . In th e Cusino a rea , red maple, balsam f i r , hemlock, and w hite spruce are among th e most abundant t r e e s p e c ie s found on t h is s o il type although cedar does occur. The organic s o i l s o f in t e r e s t to t h is problem e x h ib it a co n sid era b le range in chem ical and p h y sic a l p r o p e r tie s. Spalding p ea t i s a t r a n s it io n type from the open heath-bog typ e to the fo r e s te d typ e o f organ ic s o i l s . I t i s s tr o n g ly a c id and composed o f brown woody or spongy fib r o u s, sligh tly-d ecom p osed m a teria l. R i f le p ea t i s an a c id brown, c o a r s e ly granular or woody p e a t. The su rfa ce 4 to 8 in ch es a re more decomposed, darker c o lo red , and co n ta in more woody m a teria l than the un derlying la y e r s . The l e s s decomposed woody p ea t and raw fib r o u s peat are under the su r fa ce la y e r . I t i s covered w ith a f o r e s t typ e v e g e ta tio n and i s in term ed iate in development between a Spalding p eat and Carbondaie muck. Carbondale p eat or muck i s th e most p rod u ctive type o f organic s o i l in th e reg io n . I t i s a dark-brown, loamy or granular muck, l e s s a c id than R if le p e a t. The d ark -colored s o i l grades in to a more p e a ty , l e s s decomposed m a teria l a t a depth o f 1 to 2 f e e t . Kerston muck i s co n fin ed to th e areas im m ediately adjacen t to stream co u rses and i s ch a ra cterized by an admixture o f a lte r n a te la y e r s o f organic m a teria l and a llu v ia l m ineral d e p o sits, L ongrie loam overlieB lim esto n e bed rock and occupies upland r id g e s in the swamps in small a creages. typ e on Longrie loam. Cedar occurs as an o ld f i e l d Figure b i l l u s t r a t e s a p astu re in w estern A lger county on Longrie loam th a t i s r ev ertin g to a northern w hite cedar typ e. Figure U-. Cedar s e e d lin g reproduction, a p o r tio n o f which i s in d ica ted "by the arrow, on a Longrie loam s o i l type in w estern A lger county. T h is pasture i s r ev e rtin g to a pure stand o f ced ar. The presen ce o f p ea t s o i l s in the eastern h a lf o f th e Upper P en in su la i s common. There i s a t o t a l o f 666,112 a cres o f peat and muck s o i l s in th e co u n ties o f th a t s e c tio n o f th e s t a t e th a t have "been surveyed ( ta b le l ) . INITIAL REPRODUCTION S eed lin g C h a r a c te r istic s and Development Methods M orphological c h a r a c t e r is t ic s and s e e d lin g development were observed throughout th e course o f th e experim ents and in corp orated in to f i e l d n o te s. At the beginning o f the study, i t im m ediately became apparent th at c r i t e r i a fo r sep a ra tio n o f northern w hite cedar se e d lin g s from th e other co n ifero u s s e e d lin g s and herbaceous m aterial o f th e swamp f lo o r would need to be found. I d e n tify in g c h a r a c t e r is t ic s were a rriv ed a t through standard taxonomic procedures. Data on the change from ju v e n ile to ad u lt f o lia g e , root growth and development, r a te o f top growth, r o o t-to p r a t io s , and the in d ic a tio n s o f m ycorrhizae were a lso ob tain ed through general f i e l d o b serv a tio n s. R esu lts Northern w hite cedar s e e d lin g s , in th e ir e a r ly y ea rs, are d istin g u ish a b le from v e g e ta tiv e reproduction by the presen ce o f ju v e n ile le a v e s alon g th e main stem. Ju ven ile le a v e s are o p p o site ly arranged and g e n e r a lly tend to form four v e r t ic a l rows on the young Table 1. — Abundance and frequency o f muck lands In the ea stern h a lf o f th e Upper Peninsular-1 • • • Count;/- : * : S ectio n s : co n ta in in g : muck •• * • Percentage muck • • • • • • : Average muck : acreage per : se c tio n Zt • Acreage muck in county • Percent Percent Acres Acres A lger 19.1 8 5 .7 13M 111,936 Chippewa 18.7 6 3 .^ 184.0 1 8 8 ,6 0 8 Luce 33.^ 8 5.7 243.3 195.392 S ch oolcraft 2 2 .0 7 8 .6 176.5 170.176 T otal 6 6 6 ,1 1 2 l / Adapted from Harmer's c a lc u la tio n s ( 2 8 ) and based on S o i l Surveys in co u n ties that have been surveyed. 2 / Based o n ly on s e c tio n s co n ta in in g muck. stem as compared w ith an a lte r n a te or whorled arrangement o f a d u lt f o lia g e . Some s e e d lin g s e s ta b lis h a d u lt f o lia g e during th e f i r s t year o f growth; o th ers that a r e 2 -y e a r s-o ld have been found w ith out ad u lt f o lia g e formed as y e t . A 1 0 -y e a r -o ld s e e d lin g , twenty s i x in ch es high, was found w ith ju v e n ile f o lia g e s t i l l p resen t on two sm all bran­ ches near th e base o f the stem. However, in g e n e r a l, th e ju v e n ile f o lia g e i s no lo n g e r p resen t a t t h is age. R eversion o f f o lia g e to the ju v e n ile form in sa p lin g s and p o le s has been noted but i s rare. P revious to the form ation o f a d u lt f o lia g e , northern w h ite cedar s e e d lin g s are somewhat d i f f i c u l t to d is t in g u is h from o th er sv/amp c o n ife r s e e d lin g s . Figure 5 i l l u s t r a t e s cedar s e e d lin g s both with and w ithout a d u lt f o lia g e . F igure 6 p ro v id es a comparison between cedar, balsam f i r , and spruce s e e d lin g s . The fo llo w in g d is tin g u is h in g c h a r a c t e r is t ic s have been noted: H orthem w hite cedar: 2 co ty led o n s; ju v e n ile le a v e s o p p o site, th e h~row c h a r a c t e r is t ic bein g recog­ n iz a b le in th e m ajority o f s e e d lin g s ; e a r ly le a v e s approxim ately 3 / l 6 in ch in le n g th , sh arp ly p o in ted . Balsam f i r : k la r g e primary le a v e s w ith a whorled to a lte r n a te arrangement; whitened ■undersurface o f l e a f w ith prominent m idrib; ea r ly le a v e s approximate­ l y 6 / l 6 in ch in le n g th , b lu n t a t the t ip (o b tu se ly p o in te d ). Spruce: e a r ly le a v e s whorled, n eedle-shaped, sharply p o in ted . LeBarron (37) has developed a method o f d is tin g u is h in g black spruce s e e d lin g s from w h ite spruce s e e d lin g s . However, in view o f th e f a c t that h is method i s not a b so lu te but based on a p o p u la tio n estim a te, no d is t in c t io n v&b made in t h is study. Figure 5» Northern w h ite cedar s e e d lin g reprodu ction. Note th e se e d lin g w ithout ad u lt f o lia g e th a t is above and to the r ig h t o f th e Beedling w ith adult f o li a g e . Figure 6 . L e ft to r ig h t — spruce, northern w hite cedar, and halssm f i r s e e d lin g s . M ich.) (Photograph hy Mary Jayne H a llif a x , Munising, . . Exposed to adverse c o n d itio n s , s e e d lin g s have th e a b i l i t y to reproduce by branch la y e r in g when approxim ately f iv e y ears in age* A cedar s e e d lin g , ?•& in ch es in h e ig h t, was found w ith two s id e branches d evelo p in g as le a d e r s . They were p a r t ia l ly covered w ith sphagnum and e v id e n tly had r e c e n tly assumed an u p righ t p o s it io n , th u s reducing the p o s s i b i l i t y o f b ein g smothered by th e sphagnum. The le a d e r s th u s form­ ed were A.6 and 6 , 2 in ch es in h e ig h t. Development o f a tap root during the f i r s t s e v e r a l y ea rs seems to be in h eren t in the s p e c ie s . This tap root system i s soon rep la ced by a fib r o u s system in swamps, however. An example was found o f an e ig h t-y e a r -o ld s e e d lin g th a t had a fib r o u s root system w ith a group o f f iv e major r o o ts com prising th e main support o f th e stem; a d ven ti­ t io u s b race ro o ts had a lso formed on th e stem and served as an a id in support. The m a jority o f r o o t-to p r a t io s in 1 - to S -y e a r -o ld Beedlings f e l l between 1*25 and 3»00* The h ig h e st r o o t-to p r a t io s were found in a ro tter, vood medium. A bnorm alities in the root h a ir s o f cedar s e e d lin g s were noted although no m ycorrhizae were found and in d ic a tio n s o f the fr e q u e n tly branched e cto tro p h ic typ e (57) were not encountered. Eor th e most p a r t, th e a b n orm alities c o n s is te d o f very th ic k root h a ir s w ith a pronounced bulbous c o n d itio n a t the t i p . The la ck o f m ycorrhizae i s in agreement w ith th e li t e r a t u r e . Moore (A3 ) found th at northern w h ite cedar i s th e on ly n orth eastern c o n ife r th a t does not have .mycorrhizal fungi i n i t s r o o t s . He did n o te — to a very lim ite d ex ten t — th a t northern w hite cedar has a f in e b r i s t l y or filam en tou s growth u n lik e th e mycorrhizae th a t are c h a r a c t e r is t ic o f o th er c o n ife r s . O bservations o f cedar s e e d lin g s in Upper Michigan in d ic a te th a t th ere i s v ery l i t t l e d iffe r e n c e in t h e ir development in t h i s geogra­ p hic reg io n and in Maine as observed by C u rtis ( l 6 ) . However, C u rtis in d ic a te s th a t young s e e d lin g s may have 2 or 3 co ty le d o n s. No seed­ lin g s w ith 3 co ty led o n s were found in t h is stu d y. S u rv iv a l and M o rta lity Methods I t was noted during th e f a l l o f 19^6 th a t cedar had a very suc­ c e s s f u l seed year in th e Cusino a rea . In th e summer o f 19^7» co n sid er­ ab le cedar s e e d lin g reproduction was found alon g w ith th e u su a l v e g e ta tiv e rep ro d u ctio n . Ebur m ila cre p lo t s were l a id out e a r ly in th e growing season o f 19^7 on a B i f l e p eat s o i l type to determ ine th e amount o f cedar r e ­ p rod u ction , i t s subsequent s u r v iv a l, and th e f a c to r s cau sin g i t s mor­ ta lity . open. Three o f th e se were fen ced w ith chicken w ire and one was l e f t One o f th e e x c lo su r e s i s shown in fig u r e 7* 2he s i t e s fo r th e in d iv id u a l p lo t s were o r ig in a lly s e le c t e d on th e b a s is o f varyin g de­ grees o f l i g h t in t e n s it y and d e n sity o f ground co v er. The t o t a l rep rod u ction w ith in th e p lo t s was t a l l i e d ^ / . The cedar s e e d lin g and v e g e ta tiv e reproduction was marked w ith w ire p e g s. M o rta lity counts were made a t p e r io d ic in t e r v a ls . The amount o f mor­ t a l i t y as w e ll as i t s probable cause were a sc e r ta in e d . P lo t d e s c r ip ­ t io n s fo llo w . lZ.The reproduction p resen t a t th e tim e o f p lo t estab lish m en t has been d esig n a ted "19^7 reproduction". The amount p resen t previous to the summer o f 19^7 i s unknown. Figure 7 . I ll u s t r a t i o n o f the type ex clo su re e s ta b lis h e d to study amount o f cedar reproduction, i t s su r v iv a l, and causes o f m o r ta lity . P lo t 1 * An ex clo su r e lo c a te d In mature cedar w ith some balsam in th e canopy* Browse l i n e a t 10 fe e t* Ground cover composed o f a mos s-g r a s s- 8 ed g e-v io 1 e t-b edstraw complex* Type o f Reproduction Cedar (s e e d lin g ) Cedar (v e g e ta tiv e ) Spruce Balsam f i r Bed maple Number o f stems 191 0 k 13 k P lo t 2 * An unfenced p lo t lo c a te d in mature cedar* Canopy was composed o f cedar-spruce-balsam * Ground cover e s s e n t i a l l y moss* Type o f Beproductlon Cedar (se e d lin g ) Cedar (v e g e ta tiv e ) Spruce Balsam f i r Bed maple Black a sh Number o f stems 7k 0 3 21 6 1 P lo t 3m An e x c lo su r e lo c a te d in an a rea o f mature cedar* Im­ m ediate canopy was a l l cedar* Ground cover o f sed ge-grass-m oes tend­ ed to predominate* Type o f Beproductlon Cedar (s e e d lin g ) Cedar (v e g e ta tiv e ) Spruce Baleam f i r Bed maple Number o f stems 68 1 2 11 3 P lo t 4 * An ex clo su r e in a sm all n a tu ra l opening in a mature cedar stand* Heavy ground cover composed o f grass-oedge-m oss* The lig h t in t e n s it y in t h i s p lo t was 2 to 3 tim es g rea ter than th a t in P lo ts 1 , 2 , and 3* Type o f Beproductlon Cedar (s e e d lin g ) Cedar (v e g e ta tiv e ) Bed maple Black ash Number o f stems 29 5 5 1 In a d d itio n to the p lo t s Just d escrib ed , three o th er unfenced, m ilacre p lo t s were e steib lish ed , two in commercial c u ttin g s and one on a sm all lim esto n e rid g e in a swamp* P lo t d e s c r ip tio n s follow * P lo t 5* Located in a commercial c u t tin g made durin g th e w in ter o f 1946-47* Cover wae composed o f ced ar-b alsam -sp ru ce-low lan d hard­ woods* Ground cover l i g h t and e s s e n t i a lly moss w ith Labrador t e a , Joe-Pye weed, and Bubus s p e c ie s . True o f Benroduction Cedar (s e e d lin g ) Cedar ( v e g e ta tiv e ) Spruce Balsam f i r Bed maple Number o f stem s 5* 9 7 4-6 27 P lo t 6 . Located in a lim esto n e outcropping (L ongrie loam) in which th e su rfa ce s o i l was removed f o r road f i l l (1922-23)* Canopy none — e n t ir e ly open and unshaded. Groundcover sp arse and c o n s is t s of g r a s s , sed g e, h o r s e t a il, yarrow, straw berry, and w illow * Type o f Beproduction Cedar (s e e d lin g ) Cedar ( v e g e ta tiv e ) Spruce Balsam White b ir c h Number o f stem s 20 0 8 8 8 P lo t 7 * Located i n an area s e l e c t i v e l y cut during th e w in ter of 1946-47* Canopy composed o f ced ar-sp ru ce-b alsam -lovlan d hardwoods. Ground cover bedstraw -m oss-grass-sedge* Type o f Beproductlon Cedar (s e e d lin g ) Cedar (v e g e ta tiv e ) Spruce Balsam f i r Bed maple Black ash Number o f stem s 28 0 4 37 7 2 Be s u it s The l o s s o f s e e d lin g s during and between th ree growing season s fo llo w s a con stan t tren d (fig u r e 8 ) i f th e p lo t s are co n sid ered in to t o * D if f e r e n t ia l m o r ta lity - - both as to tim e o f occurrence and the amount o f m o r ta lity - - occurred between t h e variou s p lo t s ( ta b le 2) . Although exp ressed in sm aller numbers, counts o f the 1948 PERCENTAGE SURVIVAL OF SEEDLINGS PRESENT COMMENCING JULY 1947 IN SEVEN MILACRE PLOTS. 100 90 80 PERCENT 70 60 50 40 30 20 IP JUL OCT JU L SEP MAY ’47 '47 ‘48 '48 '49 JU L S EP '49 . '49 Figure 8 . —/ The September 19^8 c a lc u la tio n o f th e p ercen tage s u r v iv a l i s based on o n ly f iv e m ila cre p l o t s . Table 2. — S ee d lin g reproduction t a l l i e s f o r th ree growing seasons In e s ta b lish e d p lo t s ( in in d iv id u a l stem s) Date P lo t 1 2 3 4 5 6 7 T o ta l S ept. : May : 1948 : 1949 : J u ly : 1947 : O ct. : 1947 : J u ly : 1948 : J u ly 1949 : * 191 74 68 29 51 20 28 190 72 65 25 45 20 27 159 64 47 20 37 15 21 156 63 46 20 — 15 —* 148 59 41 20 13 15 7 142 53 33 18 10 14 3 138 52 30 17 7 13 3 461 444 363 — 303 273 260 Sept. 19^9 se e d lin g s showed e s s e n t i a ll y th e same tren d s in l o s s a s did th e counts on th e 19^7 s e e d lin g s (ta b le 3)* Tlle h e a v ie s t m o r ta lity was i n P lo ts 5 and 7 , th e p lo t s lo c a te d in com m ercially cut areas th a t were cut by a s e le c t iv e - c u t t in g method ( ta b le s 2 and 3 )* D i f f i c u l t y was encountered in determ ining a c c u r a te ly th e causes o f in d iv id u a l ca ses o f m o r ta lity o f northern w h ite cedar seedlingB * O bservations, coupled w ith a knowledge o f th e c lim a tic c o n d itio n s , were th e o n ly t o o ls a v a ila b le * One o f th e c h ie f problems encountered stemmed from th e f a c t th a t th e f o lia g e o f northern w h ite cedar d id not show s ig n s o f in ju r y or death u n t i l q u ite some tim e a f te r th e cau sa l agent had a c te d ^ * The e f f e c t o f Bnowshoe hare and rodent browsing are d isc u sse d elsew here in t h i s d is s e r t a t io n and are n o t in ­ cluded here in view o f th e f a c t th a t o n ly a p o r tio n o f th e p l o t s were exclosures* In view o f th e d i f f i c u l t i e s in v o lv ed , th e ca u sa l agent o f only 3^ p ercen t out o f th e 212 m o r ta lit ie s recorded was determined* By f a r th e g r e a te s t amount o f s e e d lin g m o r ta lity was caused by d esic c a tio n * In many in s ta n c e s , seed s germ inated during th e w et sp rin g season on the top o f stumps or lo g s w ith very l i t t l e d ecay, in the th in la y e r o f l i t t e r co v erin g b race r o o t s , in clumps o f moss that la t e r d ried up, or on th e swamp f lo o r in areas th a t l a t e r became desiccated * Late fro stB were found to k i l l seed lin g s* A f r o s t occurred on June 8 , 19^9» during which th e a ir tem perature im m ediately above th e su rfa ce o f the swamp f lo o r was recorded a t 2^*0° F* This fo llo w ed 3J A s e e d lin g removed from th e swamp f lo o r in i t s n a tu ra l media and p laced in s id e w ithout w aterin g was s t i l l green a f te r a 3-week period* I t was d e f i n i t e l y d e s ic c a te d , however* LeaveB were b r i t t l e , drooi^* in g , and cu rled but th er e was no ap p reciab le l o s s in color* Ta"ble 3 . — S eed lin g reproduction t a l l i e s fo r tv;o growing season s in e s ta b lish e d p lo t s ( in in d iv id u a l stem s) Date P lo t Aug. 19W 1 2 3 it 5 6 7 T otal Percent S u rvival : : S ep t. 19^8 : : May 19it9 : : J u ly 19^9 : : Sept. 19it9 16 3 2 1 it 2 7 16 3 2 1 it 2 7 16 3 2 1 2 2 3 16 2 1 1 1 2 3 lit 2 1 1 0 2 1 35 35 29 26 21 8 2 .8 7^.3 6 0 .0 1 0 0.0 1 0 0 .0 approxim ately a month o f f r o s t - f r e e weather* d ic a te d m o r ta lity caused by f r o s t . A p lo t t a l l y in July in ­ Examination o f th e p lo t s on November 1 4 , 1949, showed no in d ic a tio n s o f f a l l f r o s t as a fa c to r in m o r ta lity . Boot ro t was observed to be a l e t h a l fa c to r in s e e d lin g north­ ern w h ite ced ar. Although no is o la t io n s o f pathogenic organism s were made, appearances o f some s e e d lin g s in the swamp, e s p e c ia lly th o se on th e swamp f lo o r p roper, in d ic a te d th a t th e r o t was a post-em ergence damping o f f or a v ery e a r ly root r o t . A dark le s io n com p letely sur­ rounding th e s e e d lin g stem th a t appeared from varyin g d ista n c e above th e ground su rfa ce to the ends o f th e root system was noted . This fa c to r undoubtedly caused a c o n sid era b le number o f th e deaths c l a s s i ­ f ie d in th e "unknown" category but th e cause o f m o r ta lity was not d e f in it e enough to c l a s s i f y . The cause o f m o r ta lity d esig n a ted as " l it t e r and duff" i s in need o f ex p la n a tio n . Cedar s e e d lin g growth i s extrem ely slow . Coupled w ith t h is i s the f a c t th at la r g e cedar does n o t Bhed i t s in d i­ v id u a l le a v e s but th e a b s c is s io n la y e r forms between branch stems and sm all b re n c h lets ( c la d o p t o s is ) . Thus, i f a s in g le b ra n ch let or sever­ a l b ran ch iate happen to f a l l upon a sm all s e e d lin g , th e s e e d lin g covered w ith l i t t e r i s fo rced in to a p o s it io n p a r a lle l to th e swamp f lo o r and e v e n tu a lly k i l l e d . The d i f f i c u l t y a r is e s in fin d in g the remains o f the s e e d lin g fo llo w in g m ortality* I t i s su sp ected th a t t h is cau se o f m o r ta lity ( " l i t t e r and duff" ) i s la r g e ly lim ite d to swamp stands co n ta in in g cedar although i t may express i t s e l f on o th er co n ifero u s se e d lin g s a ls o . C lad op tosis i s p e c u lia r to cedar a lo n e o f th e Lake S ta te s swamp c o n ife r s w ith which we must contend* She hardwood l i t t e r was not observed to he as sig n i­ fic a n t a fa cto r as the cedar branches in m ortality o f t h is type* Seedlings were found w ith poorly developed root systemB which had f a ile d to branch ou t. In Borne c a se s , the development of the root systems was p h y sic a lly d i f f i c u l t due to the type o f medium upon which the seed germinated. This was e s p e c ia lly true when the germinating medium chanced to be a lo g or stump w ith l i t t l e decay. Com petition i s a ls o a cause o f s e e d lin g m o r ta lity . In the p lo t s under o b serv a tio n during th e 3 growing sea so n s, com p etition from gra ss was predominant. Grass growth would com p letely surround the s e e d lin g , thus ca u sin g i t s death towards the l a t t e r p o rtio n o f the growing sea so n . Another type o f co m p etitio n th a t was not encountered in th e se p a r tic u la r p lo t s due to th e la c k o f i t s presen ce w ith in th e study p lo t s i s com p etition from sphagnum. As p r e v io u sly s ta te d , cedar seed­ lin g s are e s p e c ia lly slow growing, slow er growing than th e accumula­ tio n o f sphagnum which may smother th e s e e d lin g s . O bviously, t h is fa c to r does not exp ress i t s e l f on th e whole o f the s e e d lin g popular tio n but o n ly in areas o f heavy sphagnum growth. Sphagnum grows much more r a p id ly in th e open than in the shade and th e com p etition be­ comes more n o tic e a b le under th e open c o n d itio n . The r e la t iv e amounts o f m o r ta lity th at th e variou s cau ses were r e sp o n sib le fo r in P lo t s 1 , 2 , 3* &nd 4 — e x c lu siv e o f snowshoe hare and rodent browsing — are as fo llo w s: Cause D e sic c a tio n Spring f r o s t s Boot r o t L it te r and d u ff P o orly developed ro o t system Com petition from g ra sses T otal Percentage m o r ta lity 38*5 19*2 15*^ 11*5 11*5 3*9 100*C Inclu d in g rodent and snowshoe hare brow sing as a m o r ta lity f a c to r , th e p ercen tage m o r ta lity caused by a l l f a c t o r s , known and un­ known, fo r P lo t s 1 , 2, 3 , and i s as fo llo w s i Cause Bodent and snowshoe hare browsing D e sic c a tio n Spring f r o s t Boot r o t L it t e r and d u ff P oorly developed ro o t system Com petition from g r a sse s Unknown T otal Percentage m o r ta lity 8*5 7*2 2«9 2*2 2*2 1*1 72*3 100.0 I t should be r e a liz e d th a t rodent and snowshoe hare brow sing as a cause o f m o r ta lity i s much e a s ie r to d e f in i t e l y a s c e r ta in than the other fa c to r s enumerated* I t i s undoubtedly a more r e lia b le estim a te o f m o r ta lity than are th e o th er v a lu e s obtained* In the p lo t s th a t are lo c a te d in th e com m ercially cu t areas (P lo ts 5 and 7)* th e m o r ta lity caused by d e s ic c a tio n was approxim ately 63*8 percent o f th e t o t a l m o r ta lity . The areas upon which th e se p lo ts were lo c a te d were found to be dry during th e l a t e summer and f a l l o f 19^8 * th e p eriod during which most o f the m o r ta lity occurred* I t i s p o s s ib le th a t some o f th e m o r ta lity c l a s s i f i e d as caused by d e s ic c a tio n might have been caused by su n scald . However, th ere seemed to toe enough canopy l e f t In th e g en eral v i c i n i t y o f th e p lo t s to p re­ clu d e t h is p o s s i b i l i t y . T a llie s made f o r sh o r te r p e r io d s, i . e . one and two growing sea so n s, on o th er p lo t s in d ic a te d th a t th e same cau ses o f m ortal­ i t y occur a s were p r e v io u s ly m entioned and in atoout th e same propor­ tio n s* The f a c to r s a f f e c t in g m o r ta lity o f cedar s e e d lin g s have re­ c e iv e d l i t t l e a t te n t io n in the Lake S t a te s . I t i s r e a liz e d th a t the r e s u lt s o f t h is study are in d ic a t iv e o n ly o f the stu dy area and th a t the importance o f each f a c to r w i l l vary w ith lo c a t io n . Montgomery (4-2) has t a l l i e d s e e d lin g rep rod u ction toy s in g le y ea rs in 1 - to 8- y e a r - c la s s e s i n a Maine stu d y. an in d ir e c t measure o f m o r ta lit y . He has thus provided His o b serv a tio n s in d ic a te th a t m o r ta lity o f s e e d lin g s approaches 100 p ercen t in f i v e y e a r s . D e sic c a tio n as a fa c to r in cedar s e e d lin g m o r ta lity has been reco g n ized toy C u rtis (1 6 ) alth ou gh he does n ot attem pt to estim a te i t s r e l a t i v e im portance. He a ls o concedes th a t nth ere i s stro n g e v i­ dence th a t 1- and 2-y e a r -o ld cedar s e e d lin g s may s u f f e r f a t a l l y from l a t e sp r in g f r o s t s " . Types o f Beproduction Methods I t has lo n g been known th a t northern w h ite cedar i s a b le to re­ produce v e g e t a t iv e ly as w e ll as toy seed . The nurseryman has found c u ttin g s to toe a s a t is f a c t o r y method o f propagating cedar commercial­ ly . However, l i t t l e i s known atoout th e s p e c if ic n a tu r el means o f v e g e ta tiv e reproduction and t h e ir r e la t iv e abundance in cedar stands* Hawley ( 29) l i s t s th ree ty p es o f v e g e t a tiv e rep rodu ction th at may be e f f e c t i v e in e s ta b lis h in g or renewing a f o r e s t : sp rou ts from th e stvunp or from th e r o o ts (root su c k e r s), c u t t in g s , and la y erin g * Pre­ lim in a ry ob serv a tio n s were lim ite d to c l a s s i f i c a t i o n in to th ese ca teg o r ie s* L ater, i t was p o s s ib le to fu r th e r sub d ivid e the c la s s e s according to th e apparent cause or method involved* A c r u is e o f twenty l/lO O th acre p lo t s was made to determ ine th e number o f stems f a l l i n g in to each ca teg o ry th a t was esta b lish ed * This n e c e s s ita te d a c r i t i c a l exam ination o f th e root system o f each stem* She p lo t s were lo c a te d in such a manner th a t c u t tin g had occur­ red in te n o f them* Some d i f f i c u l t y was exp erien ced in d eterm inin g th e o r ig in o f th e la r g e r stems* A lim it o f 8 f e e t in h e ig h t was used in an e f f o r t to o b ta in important d ata w ith in th e realm o f d e sir e d accuracy* lower lim it o f 2 f e e t was esta b lish ed * A A c l a s s i f i c a t i o n d esig n a ted "unknown v e g e ta tiv e type" was a lso e s ta b lis h e d to in clu d e stems which d id not d e f in i t e l y b elo n g in any o f th e s p e c if ic c la s s e s * B e su its Small s e e d lin g s can b e d istin g u ish e d from v e g e t a tiv e ty p e s, in most c a s e s, by the presence o f J u ven ile lea v es* Cases are found o f v e g e ta tiv e sh oots r ev e r tin g from a d u lt to Ju ven ile f o lia g e but are u s u a lly a s so c ia te d w ith rodent browsing* Several methods o f la y e r in g were noted* There i s a n a tu ra l tendency fo r the low er cedar branches to droop towards th e f o r e s t flo o r* The w eight o f the f o lia g e to g eth er w ith th e w eight o f snov and i c e fo r c e s some 'branches in to c o n ta ct w ith th e swamp flo o r * I t i s ap­ parent th a t r a p id ly growing sphagnum b eds b u ild upward in some c a se s and thus en g u lf th e branches* Duff and l i t t e r cover many o f th e branches in c o n ta c t w ith th e swamp flo o r * These p r o c e sse s a l l tend to provide a moi3t medium around th e branch th a t i s fa v o ra b le f o r th e development o f a d v e n titio u s r o o ts from th e tr a c h e a l rays o f the stem and thus th e form ation o f new in d iv id u a ls* has been d esig n a ted "branch la y e r in g 11* S his method o f la y e r in g Sigure 9 i l l u s t r a t e s t h is type o f v e g e ta tiv e reproduction* A B in gle branch may be covered a t se v e r a l p o in ts by m oist media and s ta r t form ation o f a d v e n titio u s r o o ts a t a l l o f th e s e p oin ts* Be­ cause o f th e u t i l i z a t i o n o f low branches in t h i s type o f reprodu ction, i t fo llo w s th a t t h is means o f rep rod u ction i s most p rev a len t i n th e v i c i n i t y o f stems o f th e low er age c la ss e s * Layering a ls o occu rs in co n ju n ction w ith broken or severed branches th a t are in co n ta ct w ith th e swamp flo o r * bo made as a r e s u lt o f wind-breakage or logging* (This con tact may In th e se c a s e s , the stems are not dependent upon th e root system o f th e parent tr e e but must form a root system o f t h e ir own by branch la y erin g * Two c a te ­ g o r ie s were e s ta b lish e d fo r th e se means o f v e g e t a tiv e rep rod u ction, "layerin g from windthrow" and " la y erin g from logging"* Vindthrows a lso produce v e r t ic a l steme in a manner somewhat d iffe r e n t than th a t p r e v io u sly d e s c r ib e d . o f windthrows e ith e r con tin u e In many c a s e s , th e branches to grow upward or assume a v e r t ic a l p o s it io n , o b ta in in g water and n u tr ie n ts from th e p o r tio n o f th e ro o t system o f th e parent stem th a t i s s t i l l in th e ground* An example o f F igure 9 . Branch la y e r in g . The stem h eld in th e r ig h t hend has become imbedded in sphagnum, thus p rovid in g a m oist medium fo r th e branch h eld in the l e f t hand. A d v en titio u s r o o ts have formed on th e branch and a new in d iv id u a l e s ta b lis h e d . t h is type o f rep rod u ction !/w as found in which th e parent cedar stem (3 in ch es d.b#h#) had f a l l e n and s i x o f i t s "branches had formed up­ r ig h t stems (fig u r e 1 0 )• stems# The o ld root system supported a l l th e se Three o f the stem s are now 1 in ch d .b .h ., one stem i s 2 in ch es d#b#h#, and two stem s are 3 in ch es d .b .h # The d e s ig n a tio n "windthrow" has been g iv en t h i s typ e o f growth# N atural c u t tin g s are d i f f i c u l t to tr a c e p o s i t i v e l y to th e ir o rig in # found# However, d e f in i t e examples o f t h is type o f reproduction were The f i r s t example o f t h is typ e was found in th e Upper Penin­ s u la Experim ental F o r e st, Dukes, Michigan# A stem o f v e g e t a tiv e re­ production was formed by ra b b it c u ttin g , th e c u t tin g f e l l i n g , b ein g p a r t ia l l y covered w ith d u ff , and rooting# branch r o o ts were formed# In t h i s in sta n c e , e ig h t S everal s im ila r examples have been found since# No ca se s were found o f s t o o l sp ro u tin g or o f root suckering# I s o la te d c a s e s o f form ation o f copp ice sh oots on stumps were found but th ey showed no in d ic a tio n o f development to a degree th a t might be c la s s e s as a stem o f reproduction# Cases were found in v o lv in g "topped"!/ stems in which the l a t e r a ls took over as a leader# How­ ev er, t h is probably should be c l a s s i f i e d as a transform ation rath er than a means o f reproduction in th a t no in c r e a se in th e number o f stems resu lted # !/jto r e o f a transform ation th a t tru e rep rod uction; but fo r purposes o f t h is study c la s s e d as a type o f v e g e ta tiv e reproduction# " T h e term "topped" r e f e r s to a t r e e th at has had i t s main stem cu t or broken a t a p o in t th a t i s above th e u su a l h eig h t o f cu ttin g # Eigure 10. from a windthrow. I l l u s t r a t i o n o f v e g e ta tiv e reproduction formed The sa p lin g stems have developed v e r t i c a l l y from branches on a windthrown t r e e . S ee d lin g s, 'branch la y e r s , and stem s o r ig in a tin g from vindthrow accounted fo r over 90 p ercen t o f th e reproduction (ta b le 4 ) . Approxi­ m ately 80*9 p ercen t o f th e v e g e t a tiv e reproduction was caused by la y e r in g o f one type or another* There waB no s ig n ific a n t d iffe r e n c e between the abundance o f the various types of v e g e ta tiv e reproduction in the p lo ts w ith c u ttin g and the p lo ts without c u ttin g (ta b le 5)<» The obvious exception i s in the category "layering from logging"* I t should be re-em phasieed th a t th e p lo t s sampled in determ in­ in g th e typ es o f reproduction were lo c a te d on organ ic s o ils * Obser­ v a tio n s have in d ic a te d th a t th e r e la tio n s h ip s expressed h ere between the v a rio u s typ es o f reproduction do not h o ld on th e s o - c a lle d "oldf ie ld " typ es in Upper Michigan* The reproduction i s predom inantly o f s e e d lin g o r ig in in th e s e stands* C urtis (1 6 ) m entions th ree typ es o f v e g e t a tiv e reprod uction by: 1* layerin g* 2* tops and branches o f thrown tr e e s th a t d evelop root systems* 3* thrown or p a r t ia l ly thrown stems when th e uppermost branches from them begin to grow v e r t i c a l l y in t h e ir new p o s it io n even though th e stem o f th e treo may n o t be in co n ta ct w ith th e f o r e s t flo o r * Moot o f the typ es e s ta b lis h e d in t h i s study cou ld b e c l a s s i f i e d tinder the same c a te g o r ie s; la y e r in g from windthrow and la y e r in g from lo g g in g end would thus f a l l in to th e c l a s s i f i c a t i o n "tops and branches o f thrown tr e e s develop root systems"* He does not mention n atu ral c u ttin g s as a means o f v e g e ta tiv e reproduction* (42) has evidence o f ro o t sproutin g and stump sprouting* Montgomery These types o f reproduction were not o f consequence in t h is study. Table 4 . — Types o f cedar rep rodu ction on a swamp s i t ©=' • • Type : Number o f stems : P ercentage • • S eed lin g Branch la y e r in g L ayering from windthrow L ayering from lo g g in g Windthrow N atu ral c u ttin g s Unknown v e g e ta tiv e typ es 108 326 14 9 58 1 24 2 0 .0 6 0 .4 2 .6 1 .7 1 0 .7 0 .2 4 .4 T o ta l 540 1 0 0 .0 l/ Based on t a l l i e s o f cedar 2 to 8 f e e t in h eig h t in twenty l/lO O acre p l o t s . Tahlo 5 . — R e la tiv e amounts o f v e g e ta tiv e reproduction In 10 •p lo ts w ith o u t c u ttin g and 10 u lo tB w ith c u ttin g « t Type Treatment ‘ • • Without C utting Percent Branch la y e r in g 7 4.2 Layering from windthrow 3 .0 0 .0 Layering from lo g g in g 1 6 .7 Windthrow 0 .4 Ne.tural c u ttin g s Unknown v e g e ta tiv e 5 .7 typ es • « T o ta l 1 0 0 .0 : With C u ttin g Percent 7 7 .4 3 .6 5 .3 8 .3 0 .0 5 .^ 1 0 0 .0 Kaki ( ^ l) , working in Minnesota, f e e l s th a t the proportions o f seed lin g and v e g e ta tiv e reproduction i s governed prim arily hy the type o f seed bed. He concludes that under dense, mature cedar stands, se ed lin g reproduction i s dominant due to the slow growth o f sphagnum and sparBeness o f herbaceous cover, thus providing l i t t l e in terferen ce w ith se e d lin g s. With a f a i r l y dense sphagnum mat, la y e r s are domi­ nant due to smothering o f seed lin g s by the sphagnum mats. In cut­ over areas, sphagnum u su a lly resumes growth w ith renewed v ig o r but both seed lin g s and la y er s are common. Observations in the Upper Peninsula o f Michigan do not concur w ith th ese r e s u lts o f Maki in th e ir e n tir e ty and the heavy emphasis on type o f seedbed as the determing fa c to r i s thought to be m isplaced. It i s agreed that dense, mature stands produce predominantly seed­ l in g reproduction but i t i s fo r the reason th at branch la y erin g i s the p rin cip a l type o f v e g eta tiv e reproduction and that the lower branches in a dense, mature stand have been self-p ru ned long sin ce; th is does not allow fo r the formation o f branch la y e r s . It i s agreed that sphagnum aicfc branch la y erin g but i t i s f e l t th at the age o f the stand and the se v e r ity of c u ttin g i f i t has occurred are the dominant in flu en ces in determining the type o f reproduction that w ill be p resen t. In other stu d ies concerned w ith the proportion of seed lin g reproduction to v e g eta tiv e reproduction, Curtis (l6 ) reports that re­ juvenation o f swamp cedar in Maine i s mainly through v e g e ta tiv e pro­ pagation and the number o f stems o r ig in a tin g through v e g e ta tiv e means i s probably la rg e in comparison w ith the number o r ig in a tin g from seed. T his, in e f f e c t , i s sim ilar to the r e s u lts in Upper Michigan. Ihvironm ental F actors A ffe c tin g I n i t i a l Beproduction Laboratory S tu d ies Seed V ia b ilit y Cones were c o lle c t e d during th e f a l l o f 19*4-8 from two owamp etande in th e Upper P en in su la o f Michigan and a ir-d r ie d * was ex tra cted by s ie v in g and p la ced in co ld storage* The seed Standard germi­ n a tio n t e s t s were run u sin g P e tr i p la t e s and f i l t e r paper* Further seed was obtained from a commercial seed source; i t had been c o lle c t e d in Door County, Wisconsin* Germination p ercen t­ age was ag a in c a lc u la te d fo r th e f i e l d run seed* S e le c tio n s o f w e l l - f i l l e d , h e a lth y appearing seeds w ere made and germ ination t e s t s conducted* The r e s u lt s o f a l l germ ination t e s t s were then computed on p ercen tage b ases to determ ine v ia b ilit y * Germination was determined from B a ld w in s standard (5 )* Hie d e fin e s germ ination as having occurred a t the tim e th a t th e t i p o f the r a d ic le has elongated fo r enough beyond th e seed coat to show a normal growing t i p and g iv e s in d ic a tio n s o f d ev elo p in g in to a h ea lth y seed lin g* Germination from 19*46 seed c o lle c t e d in swamp stand s in th e Upper P en in su la in d ic a te d approxim ately 3*0 p ercen t germ ination o f f i e l d run seed* Germination approximated 9*0 percent on seed ob­ ta in e d from Door County, W isconsin. By s e le c t in g o n ly w e l l - f i l l e d , h ea lth y appearing seed s from t h i s l o t , germ ination was in creased to 8 0 .0 percent under optimum co n d ition s* S everal methods o f approach in clu d in g B o r stia n 's procedure on southern w h ite cedar (32) were u t i l i z e d in an attem pt to determ ine the amount o f v ia b le seed per u n it o f f o r e s t f lo o r , hut none o f the methods proved sa tisfa c to ry * Germination counts on m ilacre p lo ts gave the o n ly in d ic a tio n th a t could he obtained o f the number o f viar* Die seeds* Throughout the course o f the study, the h ig h est germina­ tio n on an acre b a s i s i / was 4-27,000 se e d lin g s w ith counts above 150,000 seed lin g s per acre q u ite common* T a llie s and observations showed that a s a tis fa c to r y amount o f v ia b le seed was present in the areas studied* Light A study o f the e f f e c t o f lig h t on seed germination was i n i tiated * Prelim inary t e s t s under co n d ition s of f u l l su n ligh t and f u l l darkness were follow ed by a a e r ie s o f f u l l su n lig h t, f u l l darkness, end p a r tia l sunlight* Pour r e p lic a te s o f 100 seeds each were used in the l a t t e r tests* Three la y er s o f m oist f i l t e r paper were placed in P e tr i d ish e s, the f i l t e r paper serving as germ inating media* ed w ith Scotch tape to prevent m oisture lo ss* The p la te s were se a l­ Water was added as necessary* A sim ila r s e r ie s was wrapped w ith a heavy sh eet o f black paper, sea led , and then covered w ith a wrapping o f white paper* These wrappings were removed only long enough to obtain germination counts and to add water* The p a r tia l su n lig h t s e r ie s was constructed in the same manner as the f u l l su n ligh t se ries* However, i t was placed in \ l / Most o f the counts were on a m ilacre b a s is and expanded to an acre b aois fo r comparison w ith the r e s u lts o f other workers* p a r t ia l l i g h t co n d itio n s* The mean fo o t candle power fo r th e p a r t ia l li g h t r e p lic a t e s , o b tain ed from a a e r ie s o f s i x random read in gs during d a y lig h t h ou rs, was 76.3# Germination was checked sem i-w eekly. Germination under p a r t ia l l i g h t co n d itio n s appears to he b e t te r than germ ination in th e dark, from th e stan dp oin t o f v ig o r o f germina­ tio n and r a p id ity o f germ ination (ta b le 6 ) . There was no s ig n if ic a n t d iffe r e n c e in t o t a l germ ination between p a r t ia l l ig h t and darkness. F u ll su n lig h t in h ib ite d germ ination* A cid ity F iv e r e p lic a t e s , each c o n s is t in g o f IOC s e le c t e d se e d s, were ad ju sted to pH in t e r v a ls o f one u n it from 3*0 to 8*0 w ith i n i t i a l d e v ia tio n s not g re a ter than o n e-ten th pH u n it to determ ine th e e f ­ f e c t o f pH upon seed germ ination. D ifco standard agar a t a concen­ tr a tio n o f 20 grams p er l i t e r o f water was ad justed w ith n i t r i c a c id and sodium hydroxide to o b ta in th e d e sir e d co n cen tra tio n . d eterm in a tio n s. d egrees o f hydrogen-ion A Beckman e l e c t r i c pH meter was used to make th e pH Germination was a t room tem perature. The pH was measured again a t th e co n clu sio n o f th e experiment* A trend o f in crease in germination w ith a decrease in hydrogen- ion concentration i s apparent (ta b le 7)* I t becomes l e s s pronounced as the len gth of the germination period in crea ses. A s ta tis tic a l a n a ly sis o f the data in d ic a tes a s ig n ific a n t decrease in germination below pH Jf*0 under the conditions o f the experiment* Types o f Germinating Media A t e s t to determ ine the e f f e c t o f d iffe r e n t typeB o f Table 6 . — Accumulative. aeM. -gem lnatl on In d ifferen t, lig h t c o n d itio n s In lab oratory t e s t s (In number o f seeds germ inated) lig h t C ondition 7 F u ll su n lig h t--/ —/ P a r tia l lig h t Darkness 0 109 11+ 10 11+ 17 21 0 203 63 0 229 117 0 236 179 0 236 187 i f Four one-hundred seed r e p lic a t e s vrere i n i t i a t e d in each lig h t c o n d itio n . 2 / E sta b lish e d on J u ly 1, 19^9» Table 7. — Accumulative seed germination at various ~pH l e v e ls under lab oratory c o n d itio n s (In number o f seeds germ inated) » • T est P eriod in Days • • pH • • : 1/ “ '3 .0 4 .0 5 .0 6 .0 7 .0 8 .0 l/ 7 0 1 4 2 16 33 : 10 20 48 79 20 142 186 : 14 17 72 259 281 186 346 371 92 3^7 362 293 378 400 F ive one-hundred seed r e p lic a t e s were i n i t i a t e d at each pH le v e l. germ inating media under c o n tr o lle d temperature and m oistu re was i n i t i a t e d . Pour r e p lic a t e s , each c o n s is t in g o f 100 s e le c te d seed s, were germ inated in fou r ty p es o f n atu ral germ inating media; the organ ic swamp f lo o r , lo g s in va rio u s s ta g e s o f decay, hardwood l i t t e r , and m ineral s o il as a check. T^e swamp f lo o r sample was ob tain ed from th e top in ch o f an organic swamp s o i l c l a s s i f i e d as a R if le p e a t. The decaying lo g sample was obtained from th e same lo c a t io n and would f a l l in to the decay c la s s 5 according to McCullough's c l a s s i f i c a t i o n l / ( 4 4 ) . The hardwood l i t t e r sample was predom inantly composcAof alder le a v e s and was ob tain ed in a la rg e .ald er clump on th e edge of the swamp. A sm all percen tage o f red maple life te r was included in the sample. A v ery lim ite d amo^Jat of decay had occurred, but the le a v e s o f th e component s p e c ie s were reco g n iza b le. The m ineral s o i l sample th a t nerved as a check was ob tain ed from the Al h orizon of a Trenary f in e sandy loam. The samples th a t were kept in P e tr i p la te s a t room tenperatures w ith an adequate supply of m oisture.G erm ination was checked sem i-w eekly. An anal y s i s o f v a ria n ce showed no s ig n if ic a n t d iffe r e n c e in seed germ ination between any o f the media. R esu lts are shown i n ta b le 8 . 1 / McCullough d e fin e s decay c la s s 5 (^5) as f a ll e n t r e e s d e f in it e in o u tlin e but w ith decay w e ll advanced as in d ic a te d by the presence o f lo o s e fragm ents o f wood. Table 8. — Accum ulative seed germ ination in variou s typ es o f natural media under lab oratory conditions (In number of seeds germinated) Types o f Natural Media Swamp flo o r Decaying logs Hardwood l i t t e r Mineral s o i l l/ Test Period in Days 7 15 k lh 1 : lk> I 87 137 237 137 : 21 226 208 250 18*+ : 28 229 236 258 189 : 35 229 25^ 258 I 89 Pour one-hundred seed r e p lic a te s were in it ia t e d in each type o f natural media. Temperature P relim in ary t e s t s to determ ine the e f f e c t o f co n sta n t tem peratures upon the seed germ ination were e s ta b lis h e d in the la b o ra to ry a t 4°C entigrade in te r v a ls from 10° C. to 3^° C.lJ u sin g m oist f i l t e r paper i n P e tr i p la t e s as germ inating media. R efr ig e ra to r co n tro ls a t 1 0 ° ,1 4 ° , and 13° and in cu b a to rs at h igh er temperatures provided the tem perature chambers. Average d a ily v a r ia t io n interaperature approxLmateddt0»5 C. In a t h ir ty day p eriod fo llo w in g the estab lish m en t o f the t e s t s , germ ination occurred from 18° to 3^° w ith maximum germ ination to24-° to 26°. With th e se r e s u lt s in mind, a s e r ie s o f t e s t s were in s t ig a te d a t 10 °> 1 ^ ° , and 18° and a check a t room temperature to b e tte r e s ta b lis h th e minimum temperature a t which germ ination would occur. The t e s t s a t room temperature averaged 2 ^ .2 °0 . w ith an average d a ily v a r ia tio n o f 2 .6 5 ° C. as obtained from thermo­ graph rea d in g s. F iv e r e p lic a t e s of each treatm ent, each co n ta in !n g 100 s e le c te d se e d s'^ , were e s ta b lis h e d . Germination was checked sem i-w eekly. I t can be seen that lovr tem peratures have an ad verse e f f e c t upon seed g erm in a tio n (ta b le 9 ) • No germ ination occurred below 18° C. I t was a lso n o tice d th at i n i t i a l se e d lin g growth was l e s s vigorous a t the low tem peratures. l/M ech an ical d i f f i c u l t i e s n e c e s s ita te d 22° C. a 2i+° C. t e s t in stea d o f ^ S o u rce o f seed was Door County,W isconsin. Table 9. — Accum ulative seed germ ination a t va rio u s tem peratures under la b o ra to ry c o n d itio n s (In number o f seeds germ inated) • • T est P eriod in Days • • Temperature • » : 7 i / 10 ° c . 1ZjP 18° Check (2 4 .2 °) if 0 0 0 102 : 10 0 0 3 2^3 : 13 0 0 14 286 : 17 0 0 23 296 : 20 0 0 30 305 F iv e one-hundred seed r e p lic a t e s were in i t i a t e d at each temper­ ature l e v e l . F ie ld S tu d ies Season o f Seed Germination A s e r i e s o f ten m ila cre p lo t s were p ick ed c le a n o f cedar se e d lin g s reproduction p r io r to the 19^9 growing sea so n . Throughout th e growing sea so n , sem i-w eekly checks were made o f the p lo t s in an attem pt to fin d nev; s e e d lin g s . S eed lin g s found we re marked w ith numbered pegs to p revent d u p lic a tio n i n counting. The date and type o f medium upon which th e seed­ l i n g s were growing was a ls o recorded. I t i s acknowledged th a t the date a t which the s e e d lin g s were found was l a t e r in the growing season than the date o f emergence. In order to lim it t h is f a c t o r , as w e ll as the p o s s i b i l i t y of d isc o v e r in g se e d lin g s th a t might have been m issed in p reviou s in s p e c tio n s , the p resen ce o f th e two primary co ty led o n s in a h ea lth y s t a t e was deemed n ecessary fo r in c lu s io n in th e data. Emergence was f i r s t noted on th e 3rd of June and continued u n t i l the 2 8 th o f J u ly . The preponderance o f emergence occurred between th e 5th o f June and the 5th o f J u ly (fig u r e 1 1 ). The f i r s t emergence noted was th a t occurring on exposed stumps.Ho germ ination occurred on the swamp f lo o r u n t i l June 2 7 th, by that tim e, 77 p ercen t o f the germinatipn th at was noted on lo g s and stuirps in v a rio u s sta g e s o f decay hod been t a l l i e d . Germination on the swamp f lo o r was d e f in i t e l y la t e r than on lo g s and stumps. E ffe c tiv e Seeding D istan ce Methods O bservations were made throughout the course o f th e study to determ ine the e f f e c t iv e seed in g d ista n c e o f cedar. A concerted e f f o r t was made to lo c a te se e d lin g s th a t were s itu a te d i n r e la tio n SEASON OF S E E D THE 1349 G E R M IN A T IO N OURING GROW ING SEASON 10 9 9- • 1.3 S TE M S AMOUNT OF G E R M IN A T IO N 8 7 6 5 4 3 2 I 15 V--------------------j u P igu re 11. ne 30 ------------------------ 5 15 31 5 15 J U L Y ----------------------- ^ — AUGUST— ^ Germination p erio d as determined "by the number of new stems foiinc^ at each in s p e c tio n . *52 to a s i n g l e seed-b earing tr e e such t h a t the p a re n t could he d e f i n i t e l y determined. Observations were l i m i t e d to swamps or swamp borders to reduce the p o s s i b l e e f f e c t of slope. Distance and d i r e c t i o n of the s e e d l i n g from the p a re n t were noted. Frevalent winds in the area are from th e northwest. R e su lts numerous i n sta n c e s were found in which s e e d lin g s were l o c a te d from Ho to 53 f e e t from the n e a r e s t seed t r e e s . I t i s concluded, from the observations made, t h t adequate st ockin g w i l l occur HO to 50 f e e t from seed t r e e s , t h i s b ein g e s p e c i a l l y t r u e i n the d i r e c t i o n of the p r e v a i l i n g wind. The maximum distan ce of adequate se edling reproductio n from seed t r e e s was found in an area c l e a r cut according to the c l a s s i c a l d e f i n i t i o n (29) between 1957 and 19!'0. The area with s e ed lin g production i l l u s t r a t e d i n f ig u r e 12 contained 223 seedlings on a milacre b a sis. Ring counts showed a l l se edlings had developed a f t e r the c le a r c u t t i n g op e ra tion . The d ista n c e s to the n e a r e s t cedar t r e e s of se ed-bearing age were i n the c a r d i n a l d i r e c t i o n s and as fo llo w s^/; D irec tio n llorth \tfest South E ast J Distance 93* Of ec-t 15S. 0 f e e t SO.O f e e t IH3.O f e e t I t i s highly improbable t h a t animals or b i r d s could have been the d i s p e r s i n g agent i n t h i s case . All i n d ic a t i o n s suggest wind as the means of d i s p e r s a l . A H E A V IL Y SEEDED A R E A IN A C O M P L E T E .C L E A R S H O W IN G T H E NEAREST SEED TREES S c a le ; l " * 8 0 ’ 9 S e e d Trees O A re a With Heavy Seeding Browse Procurement Plot 13 Browse Procurement Plot 14 3Piguxe 12. C U T T IN G A word o f ex p la n a tio n i s due concerning th e l i g h t c o n d itio n s in t h is c le a r cu t area. The immediate area in which th e se seed­ li n g s were found was lo c a te d under cover o f p a r t ia l windthrow and h ig h h u tts o f le a n in g t r e e s . The lig h t in t e n s it y w ith in the area o f se e d lin g estab lish m en t was o n ly 10 p ercen t o f th a t throughout th e remainder o f the area. Because cedar seldom grows is o la t e d , and in view o f th e f a c t th a t few t e c h n ic a lly c le a r cu t areas con tain in g s a t is f a c t o r y l i g h t c o n d itio n s fo r seed germ ination are a v a ila b le , s a t is f a c t o r y work areas were d i f f i c u l t to f in d . B e su lts on t h is phase o f the study are thus o b serv a tio n a l o f n e c e s s it y . Bate o f f a l l i s an important fa c to r in d isp e r s a l by wind. Cedar seed i s l i g h t in w eight (approxim ately 325,000 seed s p er pound) and p o s s e s s e s a r e l a t i v e l y la rg e wing to aid in seed d is p e r s a l. On th e other hand, th e t o t a l h eig h t o f th e s p e c ie s i s sm all and a c ts towerd lim it in g the d isp e r s a l d ista n c e . Seed d isp e r s a l o f northern w hite cedar stands i s not dependent upon the wind a lo n e. Mention should be made o f the e x te n siv e work th a t i s undertaken by the red s q u ir r e l in t h i s r e s p e c t. Through­ out the course o f th e study, t h is c h a tterin g l i t t l e mammal was o fte n observed scu rrying about th e cedar t r e e s . In the f a l l , he c u ts cone c lu s t e r s and branches con tain in g th e c lu s t e r s . The per­ centage o f seed d isp ersed or ea ten by t h is animal i s not known but i t i s f e l t th a t i t has a profound e f f e c t on seed d is t r ib u tio n . Light Methods Throughout the course o f th e study, l i g h t read in gs were taken on a l l m ilacre p l o t s e s ta b lis h e d to determ ine th e e f f e c t o f l i g h t upon the i n i t i a l esta b lish m en t o f cedar s e e d lin g s and v e g e ta tiv e typ es o f rep rod u ction . Headings were taken w ith a DeJur Amsco Model 5B in c id e n t l i g h t meter and were taken as near to 10 AM and 3 PM as p o s s ib le . These tim es were a r b it r a r ily chosen because o f the sim ila r fo o t candle power in f u l l su n lig h t during th e two p e r io d s. I t was a ls o f e l t th a t th e two p erio d s were most r e p r e se n t­ a t iv e o f d a ily co n d itio n s in th at read in gs at th e se two hours would tend to b alan ce the read in gs in p lo t s th a t were shaded during the morning and open in the aftern oon and v ic e v e r sa . Headings were taken only during periodB o f a clea r sky. Except in a s in g le group o f p l o t s , read in gs were taken from oneh a lf hour b efo re th e d esign ated tim e to o n e -h a lf hour fo llo w in g the d esig n a ted tim e. In the s e r ie s o f p lo t s lo c a te d -in S ectio n 13 T44N H26W, tr a v e l d ista n c e s were o f such a magnitude th a t rea d in g s had to exten d to a maximum o f one hour p a st th e d e s ig ­ nated tim es. An extended e f f o r t towards u n iform ity in method Was made in tak in g a l l rea d in g s. That i s , the l i g h t meter was h e ld at w a ist l e v e l w ith the o p e ra to r 's bade to the sun and at a d ista n ce such th a t the o p er a to r 's shadow d id not f e l l a cro ss any p art of the m ilacre p lo t . A s in g le operator made a l l th e rea d in g s. The e n tir e p lo t was covered w ith the m eter. The reading s e le c t e d was the apparent modal read in g w ith the operator s t i l l b earin g in mind the minimum and maximum rea d in g s. This in te g r a tio n was thought to be most r e p r e se n ta tiv e o f the l i g h t co n d itio n s w ith­ in the u l o t . I t was a lso upon th e recommendation o f the DeJur Amsco Corporation en gin eers (39) th a t li g h t f lu x f a l l i n g upon the o b ject was used as a c r it e r io n fo r measurement rather than th e amount o f l i g h t r e f le c t e d from the o b je c t. C a lib ra tio n s from l i g h t meter read in gs to t o t a l l ig h t f l u x fo o t can d les were fu r n ish e d b y th e DeJur Amsco Corporation and are shown in fig u r e 1 3 . The in d iv id u a l p lo t l i g h t meter read in gs were converted in to fo o t candle power and a mean o b ta in ed f o r the p lo t . C la sses were then e s ta b lis h e d in f o o t candle power corresponding to l i g h t meter u n it s in so fa r as th e accuracy o f the instrum ent i s in b lock u n it s . The standard d e v ia tio n o f th e se ed lin g t a l l i e s was ob tain ed fo r each c la s s and p lo t r e j e c t io n s were made at a l e v e l o f g rea ter than two standard d e v ia tio n s from the a rith m etic mean. Several problems p resen ted th em selves in the d e sig n a tio n o f h eig h t c la s s e s to be included w ith in the t a l l i e s . I t i s acknowledged th a t some o f the v e g e ta tiv e rep rod u ction i s over 2 f e e t in h e ig h t at the time o f o r ig in , e s p e c ia lly th ose forming v e r t i c a l l y from windthrows. However, i t i s f e l t th a t the m ajority o f the stem s formed from branch la y e r in g are under 2 f e e t in h eig h t at the time a d v e n titio u s r o o ts form. I t i s a ls o acknowledged th a t se e d lin g reproduction and v e g e ta tiv e rep rod uction of th e same h eigh t would be q u ite w id ely separated in age. With th ese f a c t s in mind, the h eig h t c la s s from 0 to 2 f e e t in h eig h t was d esig n a ted as i n i t i a l rep rod u ction. Be s u i t s F igu res 14 and 15 i l l u s t r a t e g r a p h ica lly the abundance and C A L IB R A T IO N S FR O M L IG H T M E T E R R E A D IN G S TO THE 3 2 LOG OF LIGHT FLUX FOOT CANDLES L O G A R IT H M OF L IG H T F L U X FO O T C A N D LE S 0 2 4 6 8 10 L IG H T M ETER 12 R E A D IN G Figure 13. 14 16 18 20 fre q u en cy !/ o f s e e d lin g rep rod u ction under 2 f e e t in h e ig h t. The se e d lin g rep rod u ction abundance and frequency d a ta are in clu d ed in ta b le 35 (Appendix). The r e s u lt s in d ic a te th a t l i g h t in t e n s it y i s not a c r i t i c a l fa c to r in se ed lin g germ ination and i n i t i a l estab lish m en t throughout th e normal range o f l i g h t c o n d itio n s found in our swamp stan ds and swamp c u ttin g s . a t b oth extrem es. There i s a d ecrease in b oth abundance and frequency The c o n d itio n s at the lower end o f the t a b le , i . e . c o n d itio n s w ith an average f o o t can d le power o f 5 to 7 . were found in very dense stands and were u s u a lly lo c a te d in an area w ith a s u f f ic ie n t amount o f rep rod u ction in the somewhat la r g e r s iz e c la s s e s . The average minimum l i g h t in t e n s it y req uirem ents/ approximates 0 .0 6 7 . The c o n d itio n s at th e upper end o f the ta b le w ith a f o o t candle power o f 331 and upwards were found in 1 0 -y e a r -o ld c u ttin g s th at were h arvested under a system o f ab solu te c le a r c u ttin g . The r e s u lt s a lso in d ic a te th a t l i g h t in t e n s it y was not a c r i t i c a l fa c to r in v e g e ta tiv e rep rod u ction . I n i t i a l estab lish m en t was found throughout the range o f l i g h t c o n d itio n s th a t are p resen t in our 'swamp stands and swamp c u t tin g s ( ta b le 1 0 ). There was a decrease in both abundance and frequ en cy a t the low er extreme. The p lo t s at th a t end o f th e t a b le , i . e . c o n d itio n s w ith fo o t candle power o f 5 to 10, were found in very dense stands w ith a s u f f ic ie n t The term •'abundance11 i s used in t h is d is s e r t a t io n to d esign ate a count o f a g iv en sp e c io s p resen t on a u n it area b a s is . I t i s acknowledged th a t t h is i s a ra th er broad u se o f th e term but was used in view o f th e f a c t th a t d e n sity seemed d i f f i c u l t to v is u a l­ iz e in r e la t io n to a stand o f s e e d lin g s . Frequency i s d efin ed as a measure o f the degree o f d isp e r s io n measured on a p e r c e n tile b a s is . 2 / The r a t io between in t e n s it y f a l l i n g on the f o r e s t f lo o r and in t e n s it y in the open. ABUNDANCE OF N O R TH ER N UNDER W H ITE CEDAR VARIOUS LIG H T CONDITIONS IN SEEDLING REPRODUCTION 210 M IL A C R E P LO T S . 25 20 w is 15 z ow 10 111 O J 8 12 17 25 FO OT Bigure l b . L 34 J I I L 50 68 100 138 195 275 400 CANDLE POW ER Abundance o f northern w h ite cedar se e d lin g repro­ d u ction under v a rio u s l i g h t c o n d itio n s . Bach dot rep resen ts th e mean number o f s e e d lin g s in the p lo t s examined th a t were t a l l i e d in that s p e c if ic l i g h t c l a s s . The number o f m ilacre p lo t s in each lig h t c la s s rep resen ted in t h is fig u r e i s in d ic a te d in Table 35* 70 FR EQ U EN C Y UNDER OF NORTHERN V A R IO U S LIG H T W H IT E CEDAR C O N D ITIO N S IN S E E D L IN G 2 '0 R EP R O D U C TIO N M IL A C R E PLOTS. 0 9 8 7 6 .5 .4 3 2 6 8 12 17 25 34 50 68 100 138 195 275 400 F O O T C A N D LE F igu re 15. POWER Frequency o f northern w h ite cedar s e e d lin g repro­ d u ctio n under variou s l i g h t c o n d itio n s . Each dot rep resen ts th e p ercen tage o f m ilacre p l o t s in which cedar se e d lin g reproduction was found. Table 10. —Abundance and frequency o f v e g e t a tiv e reproduction under 2 f e e t In h eig h t In v a rio u s lig h t c o n d itio n s F oot-can d le power Abundance Frequency Number o f steraB Percent Number o f m il­ acre p lo t s in c la s s 5 -7 7 -1 0 10-14 14-20 20-28 0 .7 0 .6 1 .9 2 .3 1 .7 22. 4 3. 5 9. 68. 52. 9 14 17 22 27 28-40 40-58 58 -8 1 81-114 114-162 162 p lu s 2 .8 4 .9 0 .8 1 .4 0 .5 3 .9 69. 87. 55. *»4. 50. 64. 13 16 11 9 4 11 amount o f rep rod u ction in the la r g e r h e ig h t c la s s e s and extrem ely heavy se lf-p r u n in g . ITine o f th e rea d in g s in th e most in te n se c la s had f o o t candle power above 331 and were lo c a te d in an area th a t was c le a r cut ten y ea rs p rev io u s to th e p lo t exam ination. The co n clu sio n s correspond very c lo s e ly w ith th ose o f Malci (4 1 ). From lim ite d o b se r v a tio n s, he concludes th a t th ere i s no c o r r e la tio n between r a d ia tio n in t e n s it y and the number o f s e e d lin g s w ith o u t regard to age. Probably B allw in ( 5 ) b e s t summarizes th e e f f e c t o f l i g h t on seed s o f f o r e s t t r e e s by s ta t in g th a t l i g h t i s r a r e ly n ecessa ry but in n ea rly a l l ca ses germ ination i s improved and h asten ed by i t . T his d is c u s s io n i s not in anyway meant to in v a lid a te th e e f f e c t o f lig h t upon se e d lin g growth and development fo llo w in g seed germ ination and i n i t i a l esta b lish m en t. O bservations in d ic a te d that l i g h t i s b e n e f ic ia l, in c r e a se s th e growth o f rep rod uction, and that p o s s ib ly the s p e c ie s becomes more l i g h t to le r a n t w ith age. A c id ity Methods A study was i n i t i a t e d to determ ine the e f f e c t o f pH upon se e d lin g and v e g e ta tiv e reproduction under f i e l d co n d itio n s. In 156 o f th e m ilacre p lo t s e s ta b lis h e d throughout the course o f the p r o j e c t, pH read in gs were ob ta in ed . During the 19^8 growing season, d eterm in ation s were made w ith a " so ilte x " k i t . However, in view of A/ T his r e l a t i v e l y low fig u r e was chosen as an upper c la s s d esig n a tio n because o f the d i f f i c u l t y in o b tain in g an adequate number o f p lo t s in the more open a rea s. th e d i f f i c u l t i e s encountered due to th e organ ic matter in muck peat s o i l s clou d in g th e s in g le co lorom etric rea d in g , a LaMotteMorgen s o i l t e s t in g k i t w ith a s e r ie s o f in d ic a to r s was u sed during the 19^9 seaBon to a lle v ia t e t h i s d i f f i c u l t y . Samples were c o lle c t e d from w ith in the top 2 to 3 in ch es o f s o i l and s e e d lin g and v e g e t a tiv e reproduction was t a l l i e d . P lo t r e j e c t io n s were made a s p r e v io u s ly . R e s u lts S eed lin g reproduction in c r e a s e s w ith a decrease in hydrogenion con cen tra tio n above 6 .0 to a n e u tr a l and s li g h t l y a lk a lin e c o n d itio n (c irc u m -n eu tr a l). There seems to he no app reciab le d iffe r e n c e in reproduction in the range between pH ^-.0 and 6 .0 . In d ic a tio n s p o in t to a p o s s ib le red u ctio n below t h is l e v e l . Figure 16 i l l u s t r a t e s the se ed lin g abundance and fig u r e 17 shows the seed ­ lin g frequency data in a graphic p r e s e n t a t io n l/. The r e s u lt s in ta b le 11 in d ic a te a decrease in abundance and frequency o f v e g e ta tiv e reproduction at the h igh er pH l e v e l s (above pH 7 . 0 ) . However, t h i s i s above the organic s o i l range in most o f the study area. A ll the o b serv a tio n s in the c la s s 7»0 to 7*9 f a l l a t pH 7 .5 or below. General ob serv a tio n s a ls o bear out th ese fin d in g s concerning v e g e ta tiv e reproduction. In th e Longrie loam type^/ th a t i s p resen t in or near swamps, seed lin g rep rod uction seems to take over o ld f i e l d s th at have been u t i l i z e d as p a stu r e s. T his i s in co n tra st 2*1 The se e d lin g abundance and frequency data are p resen ted in ta b le 35 (Appendix). 2 / Longrie loam i s ch a ra cterized by a lim estone bed rock and i s n eu tra l to a lk a lin e in r e a c tio n . ABUNDANCE OF NORTHERN W H ITE CEDAR S EE D LIN G REPRODUCTION AT VARIOUS Ph LEVELS IN LOWLAND STANDS IN 156 MILACRE PLOTS. ea,t Newton sand T o ta l 1*6 . 5 ^5.5 •'46.2 15 57 13 1*6.5 1*5.1* 1*6.2 0 13 0 1 3 8 .2 85 138.1 13 Figure 19. An example o f th e typ e o f medium c l a s s i f i e d as " lo g s in varyin g sta g e s o f decay". Woody medium v a r ie s from s l i g h t l y decayed to advanced decay c la s s e s . Table 1 3 . — Percent age o f swamn f lo o r covered Tw lo g s In varyin g sta g e s o f decay on three d iff e r e n t s i t e s S it e P lo t : : Oarbondale neat Percent : : R if le neat Percent : : Newton sand Percent 1 2 3 39 11 20 lb 23 9 22 15 4 5 6 11 1 35 8 7 15 30 10 11 7 8 9 32 16 15 17 18 31 1 0 5 2 0 .0 15 .8 Mean 1 3 .1 3* the s e e d lin g s growing on ddcayed la g s had b e t te r developed r o o t system s than th o se growing on th e swamp f lo o r . IK th e p resen ce o f s u it a b le media in a swamp stand i s not adequate insurance in i t s e l f th a t cedar se e d lin g reproduction w i l l occur. Ao. absence o f se e d lin g s was noted in deer runways and other areas w ith an abundance o f deer and ra b b it s c a ts (fig u r e 2 0 ). T his might be p a r t ia l l y exp lain ed by th e tram pling e f f e c t o f deer which cau ses death o f young se e d lin g s and changes the p h y sic a l co n d itio n o f the organ ic s o i l . Several o f th e se areas su ggest a h y p o th e sis th a t th e co n ce n tr a tio n s o f s c a ts may exert a chem ical e f f e c t th a t i s in im ic a l to seed germ ination and se e d lin g esta b lish m en t. There seems to be a q u estio n in the minds o f the f o r e s t e r s and game managers throughout the reg io n as to whether the seed­ lin g s growing on p a r t ia l l y decayed lo g s w i l l be ab le to su r v iv e. The freq u en t occurrence o f co n crete examples le a d s to the h y p o th esis th a t th ere i s p o s s ib ly a fa r g rea ter number o f stems th at o r ig in ­ ated in t h is typ e o f medium p r esen t in mature stan d s than h e r e to fo r e susp ected . F ig u res 21 and 22 i l l u s t r a t e common examples o f seed­ lin g produced stem s. A num erical comparison o f adult stems th a t had o r ig in a te d on lo g s and on the swamp f lo o r was not attem pted. O bservations in d ic a te d th a t the form ation o f v e g e ta tiv e reproduction i s more dependent upon the m oisture c o n d itio n o f the medium than upon i t s conroosition. I t was found th a t a m oist co n ife ro u s l i t t e r or sphagnum moss was the b e st medium fo r v eg e t­ a tiv e form ation, probably due to the importance o f branch la y erin g as a means o f v e g e ta tiv e rep rod u ction . In a s e r ie s o f twenty a t Figure 20. I llu s tr a tio n o f h eavily used deer runway. the concentration of sc a ts in the run. Note Figure 21. An example o f a cedar p o le th a t developed from th e se ed lin g typ e o f rep rod u ction. Although i t s o r ig in i s d i f f i c u l t to prove, d ed u ction in d ic a te d th a t s e e d lin g type o r ig in would he th e o n ly p o s s i h i l i t y . Figure 22. An example o f an adult cedar th a t has developed from a seed which presumably germ inated in th e d u ff and moss on th e f a ll e n lo g . Again, d ed u ction p reclu d es th e p o s s i b i l i t y o f a v e g e ta tiv e type o r ig in . l/lO O th acre p l o t s , en average o f 8 .4 stems o f v e g e ta tiv e repro­ d u ction were found in f i v e p lo t s where sphagnum moss was sparse as compared w ith a mean o f 1 8 .9 stem s in 15 p l o t s where sphagnum moss was medium or heavy. Although lo g s in v a rio u s s ta g e s o f decay occupy a r e l a t i v e l y sm all p rop ortion o f the t o t a l a v a ila b le germ inating media ( 2 0 .0 $ , 1 5 .8 $ , and 1 3 *1$ r e s p e c tiv e ly on 3 s o i l ty p e s ), th ey are the fa v o red medium fo r seed germ ination in the f i e l d . The f a c t th a t th ere was no s ig n if ic a n t d iffe r e n c e in seed germ ination between ty p es o f germ inating media, in c lu d in g hardwood l i t t e r , in the la b o ra to r y under sim ila r c o n d itio n s o f temperature and m oisture in d ic a te s th at under s a t is f a c t o r y f i e l d c o n d itio n s o f th e se two fa c to r s among o th e r s , th ere should be no d iffe r e n c e between the typ es o f media as regards germ ination and i n i t i a l development. However, o b serv a tio n s su b sta n tia te d by m oisture s tu d ie s in d ic a te d th at th e hardwood l i t t e r in the swamps, predom inantly a medium o f ald er le a v e s , had a low er m oisture content than e ith e r the organic swamp f lo o r or lo g s in varying sta g e s o f decay. I t i s f e l t th at m oisture i s d e f in i t e l y the lim it in g value in th e estab lish m en t o f se e d lin g cedar reprodu ction in ald er clumps. The most exposed wood media are a lso su b ject to con sid erab le d e s ic c a tio n . S eed lin g s th a t might e s ta b lis h them selves on the upper p o rtio n o f a stump as in d ic a te d in fig u r e 23 would be su b ject to d e s ic c a tio n during the l a t t e r p o r tio n s o f the growing season. The r e s u lt s o f McCullough (44) in d ic a te th a t the e s t a b lis h - Figure 23. An exposed stump, a type o f germ inating medium found in swamps th a t was c l a s s i f i e d as Mlo g s in varyin g sta g es o f decayH. S ee d lin g s th a t became e s ta b lis h e d near the cut end of stumps were o fte n found to be su b ject to d e s ic c a tio n . ment o f Engelmann spruce and a lp in e f i r s e e d lin g s i s n o t dependent upon the herbaceous and shrub s u c c e ssio n or th e degree o f decay o f lo g s but th a t i t can occur on lo g s showing o n ly s lig h t s ig n s o f decay. With t h is in mind and th e f a c t th a t th e s p e c ie s in th e northern swamps th at c o n tr ib u te f a l l e n lo g s and strumas have very d ivergen t wood c h a r a c t e r is t ic s , no such c l a s s i f i c a t i o n was d esign ed . Logs w ith s lig h t decay, in the moss s ta g e s o f su c c e ssio n w ith a s c a tte r in g o f herbaceous v e g e ta tio n , were th e b e s t germ inat­ in g media. Although the e a r ly decay c la s s e s provide th e most s a tis fa c t o r y germ inating media, i t does n ot fo llo w th a t lo g s in the e a r ly sta g e s o f decay are th e b e s t typ es o f media fo r the u ltim a te e s t a b lis h ­ ment and su r v iv a l o f the rep rod u ction . O bservations in d ic a te th a t the la t e r sta g e s o f decay ore su p erior in t h is r e s p e c t. s ig n if ic a n t f a c t s are o ffe r e d in i t s support. Several There i s l e s s p h y s ic a l impediment to root p e n e tr a tio n in the la t e r c la s s e s . They a lso appear to have a higher c a p a city fo r m oisture absorption than the s l i g h t l y decayed lo g s . I t i s probable th at u n t il the p resen t century or the l a s t few y ea rs o f the p a st century — w ith the ex cep tio n o f swamps w ith heavy and com plete burns — a goodly number o f the swamp stands in the Upper P en in su la o f Michigan were v ir g in . With the very slow decay th a t occurs under th ese c o n d itio n s , the percentage o f the swamp stands occupied by a medium o f th e type most s u ita b le fo r se e d lin g development was probably at an optimum. With ev e r - in c r e a sin g u t i l i z a t i o n o f f o r e s t products and red u ctio n o f woods w a ste, the p ercen tage o f area occu p ied by t h is type o f medium w i l l no doubt d ecrea se. Such a c o n d itio n cannot h e lp but have an in ­ flu e n c e upon the fu tu r e amount o f medium most s a t is f a c t o r y fo r germ ination. Jrom a lon g time a sp e c t, there i s the p o s s i b i l i t y o f a c o r r e la tio n o f the amount o f s a t is f a c t o r y germ inating media w ith the means o f s la s h d is p o s a l. Hot o n ly was t h is type o f medium ( lo g s in va rio u s sta g e s o f decay) found very s u ita b le fo r cedar se e d lin g rep rod u ction but o b se rv a tio n s in d ic a te d th a t i t i s eq u a lly as advantageous fo r th e s e e d lin g rep rod u ction o f spruce and balsam f i r . I t i s r e g r e tte d th a t th e opp ortun ity to observe and study the e f f e c t o f f i r e upon germ inating media and cedar rep rodu ction in gen eral d id not p resen t i t s e l f . I t i s a fa c to r th at should r e c e iv e a tte n tio n . Temperature Methods The e f f e c t o f tenroerature upon seed lin g reprodu ction was undertaken in an attempt to o b ta in s p e c if ic inform ation concerning swamp ten p eratu res and to determ ine th e e f f e c t o f sev era l a sp e cts o f tenroerature upon reproduction: 1 . l a t e spring and e a r ly f a l l f r o s t s upon se e d lin g m o r ta lity . 2 . the low tenroerature o f th e f o r e s t flo o r in r e la t io n to the d elay or p r o h ib itio n o f seed germ ination. 3 . th e r o le o f s o i l temperature as a fa c to r determ ining th e p referen ce f o r the type o f germ inating medium in swamps. Two p a r t ia l weather s ta t io n s were con stru cted w ith in the swamp. M i n i m um and maximum temperatures were ob tain ed in th ree p o s it io n s at each s i t e , one set. at 2 to 3 in c h e s below th e su rfa ce o f th e swamp s o i l , another s e t from the in s id e o f decayed lo g s (th e most s a t is f a c t o r y germ inating media f o r cedar s e e d lin g s in the swamp), and a th ir d s e t from 2 to 3 in ch es above th e su rface o f the swamp f lo o r . F igu res 2k and 25 i l l u s t r a t e th e ty p es o f i n s t a l l a t i o n s . The s e t a t the Cusino s ta t io n was used as an upland check o f a ir tem peratures. Headings were ob tain ed sem i-w eekly. H e su it 8 F ig u res 2 6 , 27, and 28 in d ic a te th e mean, minimum, and maximum tem peratures throughout th e growing season at 2 to 3 in ch es above th e surface o f th e swamp f lo o r (A ir ), 2 to 3 in ch es below the su rfa ce of th e swamp f lo o r ( S o i l ) , in decaying lo g s , and a t an upland check s ta t io n (A ir ). I t can be seen that during th e periCd o f seed germ ination (June and J u ly ) , the temperatures w ith in lo g s in v a rio u s sta g e s o f decay were h igher than the temperature of th e f o r e s t f lo o r . The maximum tem peratures a tta in e d in the decaying lo g s was a lso h igh er; minimum tem peratures over­ lapped in range. This l a t t e r phenomenon appeared to occur at tim es when the w ater ta b le would drop beneath the gauges in the swamp f lo o r . The average sem i-weekly v a r ia tio n between minimum and max­ imum tem peratures was a ls o g rea ter w ith in the lo g s . I t averaged 6 . 8 ° F. f o r the swamp s o i l s ta tio n s as compared w ith 1 0 . 6° F. fo r the decaying lo g s t a t io n s . The average v a r ia tio n o f the a ir ju s t JTigure 2*4-. An i l l u s t r a t i o n o f th e lo c a t io n o f thermometers d esign ed to o b ta in minimum end maximum tem peratures above the su r fa ce o f th e swamp f lo o r . Figure 25. An i l l u s t r a t i o n o f th e lo c a t io n o f thermometers design ed to o b ta in minimum and maximum tem peratures in decaying lo g s . MEAN MONTHLY TEMPERATURES FOR SWAMP S O IL , LOG, AND AIR STATIONS AND AN UPLAND CHECK STATION. 80 70 p 60 40' 30 MAY JUN JUL AUG SEP ffigure 26 . i / 2J 3 f I f Temperature read in gs were in d egrees y a h re n h e it. ■*' Readings obtained during 19**9 season. 3 / The in la n d check s ta tio n measured a ir tem peratures. MONTHLY M INIM UM TEMPERATURES FOR SWAMP S O IL , LOG, AND AIR STATIONS AND AN UPLA N D CHECK STATION. 50 TEM PERATURE 40' 30 20 MAY JUN JUL AUG SEP Figure 27.l l Z l 3 / •*/ Temperature readings were in degrees F ah renheit. ■2/ Readings were obtained during 19^9 season . 3 / The upland check s ta tio n measured a ir tem peratures. MONTHLY MAXIMUM TEMPERATURES FOR SWAMP S O IL, LOG, AND AIR STATIONS AND AN UPLA N D CHECK STATION. 100 90 P 80 70 60 50 MAY JUN JUL AUG S EP F igu re 2 8 . 1 / 2 / 3 / —/ Temperature readings were in degrees F ah renh eit. 2 / Readings obtained during 19**9 growing season. 2 / The upland check s ta tio n measured a ir tem peratures. above the swamp flo o r was 30. 0° P. R e su lts at th e two s ta t io n s were in very c lo s e accord w ith each o th er. Table 14- in d ic a te s th e v a r ia tio n between the two s o i l and lo g s ta t io n s fo r the month o f August, 1949* I t i s a lso noteworthy th a t th e average a ir tem peratures w ith in the swamp are con sid erab ly low er than th ose o f a check s t a t io n lo c a te d on an adjacent upland r id g e . This i s e s p e c ia lly true o f the minimum tem peratures and the average monthly temp­ era tu res ( fig u r e s 26 and 27 ) . The e f f e c t o f l a t e spring and e a r ly f a l l f r o s t s on repro­ d u ction are d isc u sse d in d e t a il elsew here in t h is d is s e r t a t io n . In summation, l a t e spring f r o s t s were found to k i l l s e e d lin g s . Ho in d ic a tio n was found o f damage by ea r ly f a l l f r o s t s . (term ination o f seeds on the swamp f lo o r was l a t e . I t was d e f in i t e l y behind seed germ ination on other ty p es o f media. It i s f e l t th a t the low temperature o f the swamp f lo o r caused t h is d ela y . The f a c t th a t the mean temperature o f th e swamp f lo o r was found to be lower than th a t o f lo g s and stumps in verying sta g e s o f decay co rr e la te d w ith the amount o f se e d lin g reproduction. The fa c to r causing th e d iffe r e n c e in germ ination between th ese two media, in the c a s e s th a t were not eaqposed to such a degree th a t m oisture supply i s the lim it in g f a c to r , i s tem perature. The lim ite d work o f LeBarron and H eetzel ( 38 ) at Dukes, Michigan on swamp s o i l tem peratures p rovid es a fu rth er check on the r e s u lt s ob tain ed in t h is study. On June 11, 194-1, tw elve Table 14-. — Temperature r e la tio n s h ip s during August 1949 ( in d egrees F ahrenheit) S ta tio n 1 2 1 2 Minimum Maximum V a ria tio n ^ / L ocation : Mean : temperature temperature temperature S o il S o il Log Log 5 9 .0 5 6 .5 59.5 59.3 4 5 .0 4 6 .0 4 0 .0 4 5.5 6 8 .0 7 1 .0 7 2 .0 7 2 .0 1 0 .0 8 .8 13.6 1 1 .0 i f Based on the average sem i-weekly v a r ia tio n between minimum and maximum tem peratures. in ch es "below th e su rface o f th e swamp f lo o r , the tem perature was 52 ° P . w ith a range o f between elsew here in the swamp. and 49° in sim ila r p o s it io n s Readings f o r th a t p erio d during 19^*9 at 2 to 3 in ch es "below the swamp su rfa ce at Cusino averaged 49° 2*. fo r S ta tio n 1 and 4 8 .5 ° fo r S ta tio n 2. The la b o ra to r y r e s u lt s in d ic a te d th at the low er th resh o ld fo r germ ination o f cedar occurred somewhere between lU° and 18° 0 . (5 7 .2 ° and 6*f.4-° P . ) . I f exposure to co ld tem peratures during the w in te r p lu s a lte r n a tio n o f temperature has not markedly low ered the th re sh o ld o f germ ination, the temperature stu d ie s would in d ic a te th at the seed i s a b le to a t ta in the surface temperature o f the media in volved and commence germ ination w ith ­ out a warm la y e r o f s o i l . The h y p o th e sis th a t the exposure to co ld and a lte r n a tin g tem peratures have s l i g h t l y lowered the th resh old seems more te n a b le . T his co n clu sio n i s based on the f a c t th at swamp s o i l s are ch a ra c ter ize d b y a very h igh water ta b le during the p erio d o f germ in ation , thus p o s s ib ly reducing the temperature d if f e r e n t ia l between the surface and underlying la y e r s . S o il Moisture Methods S o il m oisture s tu d ie s w ith regards to i n i t i a l s e e d lin g reproduction were in s t ig a t e d w ith th e fo llo w in g o b je c tiv e s in mind* 1 . to o b ta in s p e c if i c inform ation on swamp m o istu res. 2 . to determ ine the e f f e c t o f low mid-summer m oisture on s e e d lin g m o r ta lity . 3 « to determ ine i f s o i l m oisture i s a fa c to r in flu e n c in g p referen ce f o r type o f germ inating media in th e swamp. Paired sanrolea o f approxim ately 50 grams each were ob tain ed semi w eekly a t each o f two s ta tio n s in each o f two typ es o f germ inating media — lo g s i n v a rio u s sta g es o f decoy and w ith in the top 3 ” o f the organ ic s o i l . These samples were taken from May 12 to September 26 during the 19*+9 growing season. Samples were oven d ried a t approxo im a tely 100 C. and reweighed when oven dry. The a ctu a l p ercen tage o f m oisture was then determ ined. Samples were obtained from media that was supporting cedar s e e d lin g reproduction. Weighings were made to a 2 mg. accuracy and m oisture p ercen tages c a lc u la te d . Two w ater l e v e l gaugeB were in s t a lle d during the summer o f 19^7* One was lo c a te d in S e c tio n 19 3y+7N B16W, c lo s e to the up­ land but w ith in a cedar-spruce ty p e. I t was s u b je c t to co n sid era b le drainage from the uplands and thus showed marked flu c t u a t io n s . The other gauge was lo c a te d in S e c tio n 30 T*+7N B l 6tf — w ith in the swaiip proper and not su b jec t to the flu c tu a tio n s d escrib ed fo r the previous gauge. Readings were taken ir r e g u la r ly during 19^7, w eekly during the summer o f 19 **8 , and weekly during the e n tir e growing season fo r 19^9 F igu res 29 and 30 show the w ater l e v e l during the 19^3 and 19^9 growing season s. R esu lts I t was noted th a t the m oisture content o f the swamp f l o o r was g rea te r than th e m oisture con ten t o f lo g s in v a r io u s sta g es o f decay upon which cedar was found germ inating (ta h le 15)» The m oisture p ercen tages o f "both typ es o f media throughout th e growing season i s w e ll above the w ilt in g percen tages arrived a t by F e u s te l and Byers (21) f o r p e a ts. The mean s o i l m oisture p ercen tage throughout the growing season was determined to be 5 6 0 .1 p ercen t fo r both s t a t io n s . The mean lo g m oisture percentage f o r that p er io d was found to be **73»9 p ercen t. The standard d e v ia tio n o f the in d iv id u a l sem i-w eekly determ inations from th e season al mean was found to be 120 p ercen t fo r th e s o i l medium and 135 p ercen t f o r the lo g medium. Water was stan d in g in the swamp on J u ly 2 3 , 19 ^9 . This c o n d itio n was not the r e s u lt o f recen t ra in s but rath er i s in d ic a tiv e o f the water ta b le a t that tim e. The data in d ic a t iv e o f the r o le o f d e s ic c a tio n in the s e c tio n o f the d is s e r t a t io n on se e d lin g m o rta lity were la r g e ly gathered fo llo w in g a sev e r a l month p eriod o f dryness in the 1948 growing season . Although determ inations were not c a r r ie d out on hardwood l i t t e r throughout the growing season , ob servation s in d ic a te d th at the p r in c ip le reason th a t v ia b le seed s o f cedar d id not tend to germ inate upon the l i t t e r was the la c k o f m oisture in the medium. Samples o f hardwood l i t t e r gathered on July 28, 19**9 fo llo w in g two days o f rainy weather w ith a p r e c ip ita tio n o f approxim ately 1 .2 4 < /> UJ ozX _) UJ > UJ _J < E UJ J<5 20 Lower Gauge Limit 25 JUL 5 JTigure 29. JU L 12 JU L 19 JU L 26 AUG 2 AUG 9 AUG 16 AUG 23 AUG 30 SEP 6 Water l e v e l readings during the 19^6 growing season in d ic a tin g w eekly flu c tu a tio n s in th e water t a b le . The S ectio n 19 gouge was lo ca te d c lo s e enough to th e upland so th at i t was su b ject to rapid d rainage. This phenomenon probably accounts fo r the g rea ter v a r ia tio n in i t s readings as compared w ith the S ectio n 30 o b serv a tio n s. 10,‘ Ground Level < /) UJ zo z z -J UJ > UJ -J ou: 20 25 Pigure 3 0 . 9 23 6 (9 5 Id 15 30 Water l e v e l readin gs during the 19*<9 growing season in d ic a tin g w eekly flu c tu a tio n s in th e water t a b le . The gauge in S ectio n 30 was inundated during most o f the ob servation p erio d . A comparison w ith th e 19^8 season i l l u s t r a t e s the extreme v a r ia tio n between growing seasons (fig u r e 29)• Table 15. — Average m oisture -percentage o f the two major typ es of germ inating media fo r c e d a i^ (In percentage) : S o il Bog Month : S ta tio n 1 : S ta tio n 2 S ta tio n 1 : S ta tio n 2 May June J u ly August September 591 579 566 5*i4 480 604 540 600 514 547 507 404 495 457 574 489 421 480 504 536 Mean^/ 5 6 0 .6 5 59.7 470.7 477.2 i f Data c o lle c t e d during 1949. 2 / Based on an average o f th e sem i-weekly readings in ch es had an average m oisture p ercen tage o f 1 4 8 .4 as compared w ith 666.2 p ercen t i n th e organic swamp s o i l and 4 4 7 .7 p ercen t f o r lo g s in varyin g sta g e s o f decay. During th e d rier p erio d s o f the growing season, t h is type o f medium was found so dry th a t crumpling w ith the hands would break th e le a f components o f the l i t t e r in to numerous p ie c e s . I t should be emphasized th a t hardwood l i t t e r im portant p o r tio n o f th e ground area o f swamps. covers an The areas which are in im ic a l to cedar se e d lin g reproduction are mainly ald er clumps or a ld er stands c o n s tit u tin g a s ig n if ic a n t acreage. The o b se rv a tio n s a ttr ib u te d to the "swamp flo o r" are composed o f m osses, co n ifero u s l i t t e r , and the remains o f shrubby and herbaceous a s s o c ia t e s among o th er th in g s. The n e e d le - t y p e ch aracter of the co n ife ro u s le a v e s i s o f such a nature th a t i t does not form a mat s im ila r to th a t formed by hardwood l i t t e r . ex cep tio n s occur i n northern w h ite cedar and w h ite p in e . The O bservations in d ic a te d that cedar b ra n ch lets on th e swamp f lo o r have a h igh m oisture r e ta in in g ca p a city and seem to be m oist enough to a llo w adequate rep rod u ction . White p in e n eed les, however, are in c lin e d to form a mat th at has a tendency to dry out on th e su rface la y e r . Drainage Methods A study was in s t ig a t e d to determ ine th e e f f e c t o f n a tu ra l d r a in a g e ^ upon n orthern w h ite cedar se e d lin g rep rod uction. Experim ental d e sig n fo llo w e d th a t o f LeBarron and N eetzel ( 3 8 ) in s o fa r as p o s s ib le . With th e ir r e s u lt s in mind, fo u r l i n e s were e s ta b lis h e d p erp en d icu lar to the C reighton creek (S e c tio n 19 T^TN, B16W) on meanders i n such a manner th a t the c l o s e s t d ista n c e from the p lo t s to a drainageway would be along th e e s ta b lish e d l i n e . The l i n e s were 125 f e e t in le n g th w ith p lo t s lo c a te d from th e bank o f th e stream to th e end o f the l i n e at 25 f e e t in t e r v a ls . A m ilacre was examined a t each p lo t lo c a t io n f o r cedar s e e d lin g rep rod u ction under 2 f e e t in h e ig h t. P lo t r e j e c t io n s were a t 2 standard d e v ia tio n s . In t h is d is c u s s io n none o f th e p lo t s has been in clu d ed th a t f a l l o u tsid e the lim i t s o f 2 standard d e v ia tio n s . Thus approxim ately 5 p ercen t o f the more e r r a tic p lo t s are e lim in a ted . Two standard d e v ia tio n s was chosen as the p o in t o f r e j e c t io n in accordance w ith the su g g e stio n o f Bruce and Schumacher ( 8 ). I n c lu s io n o f p lo ts w ith g rea ter than two standard d e v ia tio n s from th e a rith m etic mean would perm it s in g le p lo t s to ex ert undue in flu e n c y upon c a lc u la tio n s o f means. 1/ A r t i f i c i a l l y induced dralnagewoys are of l i t t l e consequence in swamp stands but th e presence o f streams and n a tu ra l drainageways e f f e c t co n sid era b le a rea s. T herefore, th e l a t t e r s itu a tio n s were chosen fo r s tu d ie s on the e f f e c t o f drainage. R esu lts from th e fou r l i n e s were t a l l i e d . These r e s u lt s prompted the esta b lish m en t o f fo u r more f i f t y f o o t lin e s w ith p lo t s at 25 f o o t in t e r v a ls . I n i t i a l rep rodu ction (0 to 2 f e e t in h e ig h t) was t a l l i e d from m ila cr e p lo t s in the same manner as p r e v io u sly d escrib ed . The area i s lo c a te d i n a R if le p e a t s o i l typ e. The cover was predom­ in a n tly cedar w ith an admixture o f spruce and balsam. Stream drainage i s in to the C reighton R iver, th e M anistique R iver, and th en ce in to Lake M ichigan. R esu lts The t a l l i e s o f the fou r l i n e s e s ta b lis h e d in d ic a te d a d e f in i t e d ecrea se in s e e d lin g reproduction from a 50 f o o t d ista n c e to th e stream 's edge. Four more l i n e s e s ta b lis h e d perp end icu lar to the drainageway and w ith p lo t s a t th e stream 's edge, 25 f e e t , and 50 f e e t supported the i n i t i a l r e s u lt s . The d ata f o r the eig h t l i n e s were grouped ( ta b le 1 6 ). I t was found th a t the p l o t s d ir e c t ly ad jacen t to the stream were in water during a p o rtio n o f the study p e r io d . I t was a lso observed th a t th e p ercen tage o f herbaceous ground cover d ecreased as one tr a v e le d away from the strea m 's edge. The v e g e ta tio n ad jacen t to the stream was m ainly composed o f sed ges. L ight read in gs in d ic a te d th a t the swamp su rface ad jacen t to the stream *3 edge r e c e iv e d more l i g h t than the p lo t s fa r th e r removed due to the opening made by the stream i t s e l f . However, li g h t measurements c a r r ie d on throughout the course o f the study in d ic a te d th at the v a lu es o b tain ed were not too high f o r s e e d lin g estab lish m en t. 110 Table 16. — Average se e d lin g reproduction a t variou s d is ta n c e s from a natural drainageway. • • D ista n ce from drainage : ! : Average number o f se e d lin g s Number o f o b serv a tio n s 3 .7 5 5 .3 8 8 .1 2 11.75 13 .50 10.75 8 8 8 b b b • • Stream1s edge 25 f e e t 50 « 75 " 100 » 125 " I I t i s f e l t th a t the r e la tio n s h ip s ob tain ed in t h is phase of study were c o r r e la te d w ith in u n d ation hut no experim ental evid en ce can he produced i n h e h a lf o f t h is h y p o th esis. Rodent and Snowshoe Hare Browsing Methods Rodent and snowshoe hare browsing was n oted very fr e q u e n tly in cedar reproduction i n the study area. With th e se o b servation s in mind, a s e r ie s o f p lo t s were in s t ig a t e d to check th e combined e f f e c t of th e ir browsing upon i n i t i a l s e e d lin g rep rod u ction(0 to 2 f e e t in h e ig h t). Two rodent and snowshoe hare proof p lo t s and two check p lo ts were co n stru cted and lo c a te d in such a manner th a t rodent and snowshoe browsing w ith in th e p lo t s could be the on ly s ig n if ic a n t d if f e r e n t ia t in g f a c t o r . Costs o f m aterial and la b o r lim ite d fu rth er r e p lic a tio n . There was n e c e s s a r ily some d iffe r e n c e between l i g h t co n d itio n s in the check and th e animal proof p l o t s but t h is was reduced in so fa r as p o s s ib le by p la c in g a p a r t ia l w ire along th e check p lo t s id e s . The p lo t s each covered one square meter o f the swamp su rfa ce. O ne-half inch p ip e was cu t and assembled in to a square meter frame w ith a bar down th e cen ter f o r support. A union was inclu ded in each h a lf o f the frame to enable i t to be tig h ten ed to the exact s iz e . This frame was attached to 6 -fo o t le n g th s o f th ree- fo u rth s inch p ip e and d riven th ree f e e t in to the ground. The tops o f the p lo t s were covered w ith k x 4 mesh hardware c lo t h . The s id e s of the animal p roof p l o t s were dug i n to a depth o f 6 in ch es below the swamp f lo o r l e v e l . The w ire s id e s of th e check p lo ts were p laced a t l*f in ch es from th e ground l i n e to allow ample room f o r rod en ts and snowshoes to en ter the p l o t . E igu res 31 (a) and (b) i l l u s t r a t e the rodent and snowshoe proof p l o t s and the check p lo t s . An in d ex o f th e rodent p o p u la tio n i n the study area was obtained during th e summer o f 19*1-3 ( ta b le 17)» fo llo w in g i n gen eral th e procedure o u tlin e d by C a lh o u n (ll). A t o t a l o f 605 trap n ig h ts in th ree days trapping on fou r l i n e s were used to o b ta in the in d ex . The trap lin e s were p la ced a t a d ista n c e from the p lo t lo c a t io n s th at should p reclu d e th e p o s s i b i l i t y o f an in flu e n c e on th e rodent p o p u la tio n but were s t i l l c lo s e enough to be w ith in the same cover type. The w in ter rodent p o p u la tio n was not o b tain ed . No in d ex of snowshoe hare p o p u la tio n was determ ined although o b servation s in d ic a te d that the snowshoe hare were abundant i n the study area. R esu lts Table 18 in d ic a te s the s e e d lin g m o r ta lity in the p lo t s between th e tim e o f estab lish m en t (J u ly , 19*J8) and the tim e th e snow was o f f the ground in the fo llo w in g spring (A p r il, 19**9)• Examinations showed th a t the m o r ta lity o f 8 .5 p ercen t o f the s e e d lin g p o p u la tio n in the check p lo t s was d e f in i t e l y due to rodent and snowshoe hare browsing. Snowshoe hare s c a ts were found on the f lo o r s o f th e se p lo t s . In P lo t 299, a check, three o th er s e e d lin g s had l o s t th e ir t ip s due (h) Check p lo t EXT Figure J l , (e ) Rodent end snowshoe hare p roof p l o t , Table I? . — R esu lts o f 605 trar> n ig h ts on four rodent trap pin g lln e s ^ / ( in in d iv id u a ls trapped) S p ecies Sorex cinereus(M asked shrew) Clethriononws gapneri(R ed-backed v o le ) M icrotus t(ennsylvanicus(Meadow v o le ) Eutamius minimus(Least chipmunk) : Male 6 1 2 1 l / The trapping p eriod was during August 19*4-8. : Female 6 2 0 0 Table 18. — S eed lin g counts In rodent and snowshoe hare p roof p lo t s and checks ( i n number o f stem s) • • P lo t Type o f P lo t 300 296 299 294 Rodent and snowshoe p ro o f Rodent and snowshoe p ro o f Check Check J u ly : A p r il : 1948 : 19.^9 : 33 35 38 33 33 3433 29 Loss Caused by Browsing , 0 ,0 I /2 4- l / The lo s s o f the oth er 3 se e d lin g s was caused by fa c to r s other than rodent and Bnowshoe hare browsing. to 'browsing. damage. F iv e se e d lin g s i n P lo t 29 4, a ls o a check, showed sim ila r Thus a t l e a s t 20 p ercen t o f the s e e d lin g s in th e check p lo t s showed browsing m o r ta lity or in ju ry during a s in g le w in te r . Browsing o f sm all s e e d lin g s was not co n fin ed to the s p e c if i c area in clu d ed in t h is phase o f stu d y. Counts made in the Escanaba E lver area — s p e c i f i c a l l y , S e c tio n 13 T44N R26W — in d ic a te d wide­ spread damag e by rodent and snowshoe hare browsing a ls o . On a m il- acre p lo t o f 103 s e e d lin g s , 20 o f them (1 9 .4 $ ) were s e v e r e ly browsed. Another p lo t c o n ta in in g *101 s e e d lin g s had approxim ately 40 p ercen t o f i t s se e d lin g s browsed. Although the s tu d ie s in c lu d e both rodent browsing and snowshoe hare browsing, the r e s u lt in g damage was grouped. mice and snowshoe damage were found. Both A m icroscopic check in d ic a te d equal damage by th e two groups in s o fa r as i d e n t if i c a t io n cou ld be made. I t was not p o s s ib le to determ ine the type o f damage to seed­ lin g s i n th e ir f i r s t few y ears o f growth. The cedar stem was more of a su ccu len t than woody nature and methods o f i d e n t if i c a t io n (48) did not adapt th em selves. The s e e d lin g s were too sm all to in d ic a te more than a s in g le to o th -c u t. The e f f e c t o f rodents on s e e d lin g rep rod u ction i s not unique to northern w h ite cedar. Krauch (3 3 , 34) has dem onstrated th a t s il v i c u l t u r a l methods designed f o r c o n tr o llin g o f the rodent p op u la tio n i s n ecessa ry i n D o u g la s-fir s il v i c u l t u r e in order to o b ta in adequate sto ck in g . I t was thought th a t the check p lo ts might a ttr a c t rod en ts and. snowshoes due to the f a c t th a t th e hardware c lo t h used as a coverin g d id c o l l e c t a la y e r o f snow and thus exposed t h is area b efore the surrounding swamp f lo o r in the sp rin g . However, the r e s u lt s o f checks in other a rea s tend to d is c r e d it the p o s s i b i l i t y of t h i s being a major source o f erro r. S o il and Cover (types Methods S o il (types Some o f th e conponent f a c t o r s c o n s tit u tin g th e d iffe r e n c e s between and among s o i l type3 were stu d ied in d e t a i l rath er than as a study o f the sura ex p ressio n o f the edaphic f a c t o r s as s o i l and are p resen ted in o th er s e c tio n s o f the d is s e r t a t io n . However, in ­ so fa r as many o f th e o b serv a tio n s were made by a s o i l type grouping, r e s u lt s o f th e se o b serv a tio n s are in clu d ed in th e fo llo w in g d is c u s s io n . Cover (types Stand c o n p o sitio n s and b a sa l areas were obtained by l/lO t h acre c r u is e s on 15 p lo t s which were e sta b lish e d i n con ju nction w ith the s o i l type o b se r v a tio n s. R e su lts S o il (types Although northern w h ite cedar occurs on K erston muck, inundated c o n d itio n s caused by beaver dams on th e Ker3ton mack t r a c t s w ith in th e g en era l study area precluded th e p o s s i b i l i t y o f stu d y in g t h is typ e. Spalding p e a t stands in c lu d in g cedar were found to be l i g h t l y stocked. Cedar was s c a tte r e d , o ld , and sm all; cedar repro­ d uction was lim ite d . Eeproduction on Ogemaw f i n e samdy loam was extrem ely lim it e d . Cedar i s not one o f the dominant s p e c ie s in the o r ig in a l sta n d s. Although t h is s o i l c la s s would f a l l in to a drain age caten a w ith Saugatuck and Newton sands more c l o s e ly than w ith th e upland s o i l ty p es, i t showed l i t t l e promise o f becoming a cedar-producing type. Newton and Saugatuck sands are more or l e s s in term ed ia te i n number o f se e d lin g stems between th e poorly d rain ed upland s o i l s w ith cedar occu rrin g, in the canopy and the organ ic swamp s o i l s . The h ig h e st numbers o f stems o f rep rodu ction were found i n th e more h ig h ly decomposed organic s o i l s of the swamps, B i f l e p ea t and Oarbondale muck. Although Longrie loam does not cover e x te n siv e a rea s, i t was found i n swamp border and rid g e s it u a t io n s . E x c e lle n t cedar reproduction occurs in a f o r e s t typ e that can j u s t i f i a b l y .be in clu d ed in the " old fie ld " cedar type ( 1 5 ) on lo n g r ie loam. The depth o f organic s o i l was measured to provide a c le a r e r p ic tu r e o f the c o n d itio n s e x is t in g throughout the swamp and adjacen t r id g e s. The depths v a ried markedly — from 2 to 3 in ch es on upland s i t e s and Newton and Saugatuck sands to as deep as 7 6 .5 in ch es in the deep E i f l e p e a t. Watson ( 5 6 ) in d ic a te s th a t the s i t e i s pro­ g r e s s iv e ly b e tte r as the organic s o i l la y e r d ecrea ses in depth. S it e in d ic e s by s o i l types were obtain ed fo llo w in g the su g g estio n s o f GevorkLantz and Duerr ( 2 5 ) . The v a lu es obtain ed p laced the s i t e s w ith in a narrow range of d iff e r e n c e s . With the excep tio n o f a Newton sand area and the Oarbondale muck typ e, the s i t e in d ex fo r cedar in the s o i l typ es stu d ied f e l l in the 45 c la s s . The Newton sand a rea was a somewhat poorer s i t e , the s i t e in d ex o f one o f the p lo t s f a l l i n g in to th e 35 c l a s s . Oarbondale muck p l o t s , a s i t e in d ex o f 55 On one o f th e o b tain ed . No stan ds of L ongrie loam o f s u f f i c i e n t h e ig h t to s a f e ly f i t the ta b le s were found. S it e in d ex as u s u a lly d e f in e d ^ d id n ot seem to ad eq u ately d if f e r e n t ia t e cedar s i t e s . Cover Types Northern w h ite cedar was found to occur most commonly in the Cusino area in th ree major f o r e s t ty p es; th e b la ck spruce ty p e, the northern w hite cedar type, and th e b lack ash-Araerican elm -red maple typ e. I t s occurrence in o th er ty p es was lim it e d . In c lo se d stan d s, i t was found th a t an in c r e a s in g amount of cedar reprod uction seemed to be roughly c o r r e la te d w ith an in c r e a s in g b a sa l area o f ced ar. Total reproduction under 2 f e e t i n h e ig h t in 6 m ilacre p lo ts lo c a te d in stan d s w ith l e s s than AO square f e e t b a sa l area o f cedar p er acre showed an average o f l e s s than one s e e d lin g p er p lo t as com­ pared w ith 6.^ s e e d lin g s per m ila cre in 9 p lo t s on which the b asal area o f cedar ranged from ho to 100 square f e e t . V eg eta tiv e repro­ d u ctio n averaged 1 .8 stems per m ila cr e f o r th e low er b a sa l a rea s and 9*9 stems per m ila cre f o r th e h igh er ones. There i s a d e f in i t e r e la tio n s h ip between s o i l and cover ty p es. The black spruce or tamarack types are c o r r e la te d w ith Sp ald ing p ea t. The b lack spruce typ e i s a ls o found to be p rev a len t ^ S i t e index i s u s u a lly d efin ed as the h e ig h t in f e e t o f th e asym ptote of the h e ig h t-d .b .h . cu rve. on Newton sand. Ogemaw f i n e sandy loam areas supported an a s s o c ia t io n that would prohahly he c l a s s i f i e d as a w h ite spruce-halsam fir -p a p e r "birch ty p e. Ihe L ongrie loam s o i l type i s a s s o c ia te d w ith th e northern w hite cedar cover typ e and Saugatuck sand w ith th e "black ash-American elm -red maple type. I li f l e p e a t, K erston muck, and Oarbondale muck support a ced arspruce-halsam f i r tamarack a s s o c ia t io n , a l l o f which would he c la s s e d as the northern w h ite cedar typ e. On the Oarbondale muck, th ere i s an in flu x o f lowland hardwoods and th e most h ig h ly decomposed a reas of th is type might he found supporting the "black ash-American elm -red maple typ e. Age o f Stand Methods O bservations throughout th e study in d ic a te d th a t th e age o f main stems a t the tim e o f c u ttin g when con sid ered on a stand b a s is might in flu en ce the la y e r in g a b i l i t y o f the low er branches. It i s a ls o known th at se lf-p r u n in g red u ces the number o f lower branches th a t would be capable o f la y e r in g at th e time o f c u ttin g . Two swamp cedar stands were chosen fo r the study. Their s e le c t io n was p rim a r ily based on the r e la t iv e amounts o f v e g e ta tiv e reproduction over 2 f e e t in h e ig h t. A com p letely c le a r cu t area at Cusino had e x c e lle n t v e g e ta tiv e rep rod u ction . The B ob 's Lake r e le a s e c u ttin g s (P lo t 36 ) had l i t t l e or no reproduction o f a v e g e ta tiv e n atu re. The stand adjacent to the c u ttin g at Cusino was p rim a r ily in th e 0 to 6 inch d .b .h . c la s s e s . The rem aining uncut stand at Bob' s Lake appeared mature w ith overmature stems in evid en ce. Five l i n e s ( l chain x 10 l in k s each) e s ta b lis h e d in each of the two lo c a tio n s were searched f o r cedar stumps o f the p rev io u s c u t, i . e . approxim ately 10 y ea rs ago. The stumps were measured w ith a diameter tap e, cut back to o b ta in a fr e sh c u t, and smoothed down to a surface such th a t the annual r in g s could be a sc erta in ed w ith the a id o f a hand le n s . Only 3 o f the stumps in the 10 p l o t s were decayed to the p o in t th a t a p a r t ia l estim ate o f age was n ecessa ry . T a llie s o f v e g e ta tiv e rep rod u ction o f cedar and the number o f stems o f other s p e c ie s were made in each o f the p lo t s . c la s s was d esig n a ted as from 2 to 8 f e e t . H eight The average age o f the cedar stumps in each lo c a t io n was c a lc u la te d and the mean diam eter o f the stems at each lo c a tio n was a scerta in ed . The number o f former stems by age c la s s e s was ob tain ed and summarized in an accum ulative t a b le . R e s u lts More cedar stems o f sm aller diam eter and younger age c la s s e s v/ere cut in the Cusino c u ttin g s than in the Bob 1 s Lake c u ttin g s ( ta b le 1 9 ) . The d iffe r e n c e in th e amount o f cedar v e g e ta tiv e rep rod u ction in favor o f the Cusino c u ttin g i s very apparent ( ta b le 2 0 ). In th e l i g h t o f o b se rv a tio n s o f th e rem aining p l o t s at B ob 's Lake, i t i s c le a r ly evid en t th a t the typ e o f c u ttin g ^ / d id not d if f e r enough from th e Cusino c u ttin g s to cause the d i f f e r ­ ence in r e s u lt s . R e lia b le e s tim a te s in d ic a te th a t the Cusino stand was cut a t a much e a r lie r age than the Bob' s Lake c u ttin g and sm aller stems were cut although we were not ab le to e s t a b lis h the ex a ct age o f th e stands at th e tim e o f c u ttin g . Table 21 p r e se n ts d ata showing th a t t h i s c o n d itio n e x i s t s — the f a c t th a t 64 p ercen t o f the stems cut at Cusino were in the 80-year age c la s s or under whereas only 36 p ercen t o f th e cedar stem s at Bob1s Lake f e l l w ith in t h is ca teg o ry . In a c tu a l numbers, th ere were tim es as many stems in the 80 -y ea r c l a s s or under at Cusino as at B ob 's Lake. Purther o b se rv a tio n s d is c lo s e d th a t the on ly v e g e ta tiv e reproduction found in the B ob 's Lake c u ttin g was lo c a te d in P lo t 40 — a p lo t cut to a 4 in ch es d .b .h . minimum diameter lim it but w ith young cedar stems from 2 f e e t in h e ig h t to 3 inches d .b .h . The Cusino c u ttin g was an a b so lu te c le a r c u t. The Bob's Lake c u ttin g (P lo t 3 6 ) was to a 4 in ch es d .b .h . minimum diameter l i m i t . ±23 Table 19. — Number o f cedar stumps, t h e ir mean ages and diam eters P lo t s Item : 1 : •• 2 : •• 3 •• 4 : : 5 : Weighted : mean Bob1s Lake C uttings (P lo t 2 6 ) Number o f stumps Mean age in years Mean diam eter in in ch es 11 130.1 7 .2 2 158.5 1 0 .6 6 152.3 5 .9 5 1 0 6 .2 4 .6 1 2 9 .4 6 .4 Cusino c u ttin g s (Browse Procurement P lo t) Number o f stumps Mean age in years Mean diam eter in in ch es 7 111.0 4 .6 6 4 8 .6 2 .1 15 7 7.0 3 .8 11 1 0 7 .0 ^ .3 15 74.6 3-3 83.7 3 .7 l / I n d ic a tio n s showed that the stumps sma.ller than 4 in ch es d .b .h . were in or along sk id road s. I 124 Table 20. - - Reproduc tio n from 2 t o 8 f e e t in h eigh t (In number o f steins) t • : Stems P lo t • : 1 : 2 : 3 : k : *5 : Per ecre Boh* s Lake C uttings (P lo t 36) Cedar (v eg eta .tiv e) Other stems per a c r e l / 0 ^3 0 53 0 66 o 50 0 6^ o 5520 Cu sin o C uttings (Browse Procurement P lo t ) Cedar (v e g e ta tiv e ) Other stems per a c r e ! / 52 35 26 7^ 20 68 22 56 38 5960 2820 1 / A l l a ld er and w illo w stems above 2 f e e t in h eig h t were t a l l i e d in view o f t h e ir rapid r a te o f growth. Other s p e c ie s t a l l i e d in clu d e spruce, "balsam f i r , tamarack, w h ite "birch, red maple, r e d -o sie r dogwood, "balm o f g i l i a d , "black cherry, serv iceb erry , and aspen. Table 21. — Accumulative t a l l y o f stumps by o.r:e c la s s e s (in number of stumps) Age n u i/ • 40 : 60 o CO :20 : : 100 : 120 : 140 : 160 : 180 j Over 180 Bob *s Lake c u ttin g s (P lo t 36) 1 2 3 4 5 0 0 0 0 0 0 0 0 1 1 2 0 1 1 1 4 1 2 1 1 7 1 3 2 1 7 1 3 2 1 7 1 3 5 1 7 1 4 5 1 9 1 4 5 1 11 2 6 5 1 Total 0 2 5 9 14 14 17 IS 20 25 Cusino c u ttin g (3rov;se Procurement P lo t ) 1 2 3 4 5 0 0 l 0 1 0 2 4 1 2 1 4 5 3 7 Total 2 9 20 2 6 11 4 11 3 6 13 6 12 4 6 13 7 14 7 6 13 9 15 7 6 15 10 15 7 6 15 10 15 7 6 15 11 15 4-0 44 50 53 53 54 in evid en ce. A com bination o f two f a c to r s appears to m an ifest i t s e l f in t h is ex p ressio n o f the age o f the stand upon v e g e ta tiv e cedar rep rod u ction. The stands co n ta in in g predom inantly mature and overmature stems w ith a h igh "basal area per acreXl are s e l f ­ pruned to a h eig h t th a t would p reclu d e th e p o s s i b i l i t y o f e x te n siv e branch la y e r in g , the p r in c ip le typ e o f cedar v e g e ta tiv e rep rod u ction. The younger stems th a t do have lower branches a lso seem to exp ress more v ig o r in the branch la y e r s th a t form than do old er t r e e s . ^ Gevorlciantz and Duerr (25) c o r r e la te d in c r e a sin g age w ith in crea sin g d e n sity o f cedar when expressed as b a sa l area. ADVANCED REPRODUCTION S u rv iv a l and M o rta lity Methods Twenty four l/lO O th acre p lo t s were e s ta b lis h e d i n a heavy winter deer yard on a R i f le p ea t s o i l type to determ ine the amount o f m o rta lity in the 2 to 8 f o o t h eig h t c la s s and the ca u sa l agen ts. This n e c e s s ita te d a c r i t i c a l exam ination o f each stem, in clu d in g the ro o t system , to determine the cause or causes o f m o r ta lity in so fa r as p o s s ib le . In the event th a t the e f f e c t o f two or more p o s s ib le causes was noted, e .g . both deer and snowshoe hare browsing, the fa c to r th at was most apparent was chosen. If the cause of m o r ta lity was not c le a r - c u t, the stem was c la s s e d in an "unknown11 category. Advanced reproduction was e s ta b lish e d as th at reprodu ction that was from 2 to 8 f e e t in h e ig h t. wa3 made fo r a combination o f rea son s. The h eig h t c l a s s i f i c a t i o n O bservations in d ic a te d that a d iffe r e n t s e t o f environm ental fa c to r s are able to exp ress them selves on th ese stems than on lower stem s. The h eig h t at establishm ent to the 2 fo o t l e v e l would be the most eq u ita b le s iz e c la s s fo r both i n i t i a l se e d lin g and v e g e ta tiv e reproduction. The d esig n a tio n o f 2 to S f e e t as advanced reproduction would a lso allow a more complete a n a ly sis o f the o r ig in a l p lo t t a l l i e s in so fa r as they were taken in 0 to 2, 2 to 5> 5 to 8 fo o t h eigh t c la s s e s . Comparable t a l l i e s were made in an unbrowsed p o rtio n o f the swamp t u t m o r ta lity numbers were i n s u f f i c i e n t to make a num erical comparison between the causes o f m o r ta lity . R e su lts A t o t a l of 72^- cedar stems were t a l l i e d in tw enty-four l/lO O th acre p l o t s . Of t h is t o t a l , U9 .3 p ercen t (357) stems were l iv in g and 5 0 .7 p ercen t ( 367 ) were dead. m o r ta lity r e p r e se n ts. I t i s not known how lon g a p erio d t h is Decay o f woody m a teria l i s known to be v ery slow in swamps and i t i s probable th at the m o r ta lity i s the accumulated m o r ta lity over a co n sid era b le p erio d o f y ea rs. Deer browsing can r e a d ily be d istin g u ish e d by i t s ragged c u ts and the " p u llin g o ff" o f browse. M o rta lity o f advanced reprod uction caused by deer browsing i s shown in fig u r e 3 2 . A comparison o f reproduction th at i s f r e e from browsing i s shown in fig u r e 3^* Figure 35 i l l u s t r a t e s stems th a t have developed to a h eig h t such th at th e ir top f o lia g e i s u n a v a ila b le to deer. Snowshoe hare browsing i s ch a ra cterized by sharp, smooth c u ts o f the lim b s. Although a com bination of deer browsing and snowshoe hare browsing was found to occur, l i t t l e tro u b le was experienced i n determ ining which cause was predominant. Oases were observed in which ro t had occurred on the branch from which the root system evolved in stems o f v e g e ta tiv e repro­ duction formed by branch la y e r in g . The r o t then seemed to p rogress­ iv e l y invade that p o rtio n o f the branch under th e su rface o f the swamp f lo o r . M o rta lity was l a i d to com p etition in heavy clumps o f repro­ d u ction th a t were under a heavy o v ersto ry . These cases were F igure 32. Cedar m o r ta lity caused "by d eer "browsing. th e presence o f o th er woody s p e c ie s th a t are uhbrowsed. Hote jL O U Pigure 3 3 . ■browsing. Deed, and d yin g young cedar as a r e s u lt o f deer P o ta g a n n issin g D eer Yard, Drummond Isla n d , 19^1. (Photograph hy S.C . W h itlo ck .) Figure 3^. Unbrowsed cedar rep rod u ction. w ith a h e a v ily “browsed c o n d itio n (fig u r e 3 2 ) . T his may 1)6 compared .JL O i< w Figure 35* Browse l i n e on cedar. Yard, Drurunond Isla n d , 193^» Long P oin t Base Lake Deer (Photograph hy P .E . G i l l e t t e . ) I thoroughly perused fo r deer and snowshoe hare "browsing b efo re the stems were c l a s s i f i e d under th e cause o f com p etition . In some stem s, formed by branch la y e r in g , th e a d v e n titio u s r o o ts f a i l e d to develop . number. Roots were found to be sh o r t and few in I t was rea l i z e d th a t many o f the sm aller r o o ts might have decayed and would not be in evidence i f m o r ta lity had occurred s e v e r a l years p reviou s to th e time o f exam ination. The lead er (main stem) o f some o f the rep rod uction was found broken. The probable cau ses were snow and i c e accum ulations with p o s s ib le wind a c tio n . Windthrow o f stems o f th e la r g e diam eter c la s s e s was found to d estro y reproduction during the p ro cess o f windthrowing. The wind­ throw wa3 not th a t o f the rep rod uction i t s e l f . The two most freq u en t causes o f m o r ta lity in twenty l/lO O th acre p lo ts lo c a te d in a p o r tio n o f th e swamp w ith no deer a c t iv it y were snowshoe hare browsing and root r o t . Deer browsing has lon g been known to be an important fa c to r in m o r ta lity o f cedar reproduction in the h eig h t c la s s e s between the snow le v e l and the upper lim it s o f a v a ila b ilit y to the deer. This p a r tic u la r m o r ta lity study on advanced reproduction a ttr ib u te d ^5 p ercen t o f the m o r ta lity to deer browsing (ta b le 2 2 ). Snowshoe hare browsing accounted fo r approxim ately 20 p ercen t o f the m o r ta lity encountered. Alaous and Aldous ( l ) concur in the importance o f snowshoe hare browsing in r e la t io n to f o r e s t repro­ d u ction by regarding i t as one o f th e lim it in g fa c to r s in p la n ta tio n estab lish m en t in th e Lake S ta te s . The r e s u lt s o f t h is study concerning snowshoes are in Table 22. — Advanced cedar re-production m o r ta lity in a w in ter deer yard on a R if le neat s o i l type • • Humber of stems Percentage lo s s Deer browsing Snowshoe hare browsing Hoot rot Competition F a ilu re o f root system to develop Broken le a d e r Y/indthrow Cause unknown 164 72 22 19 16 4 4 .7 1 9 .6 6 .0 5 .2 4 .4 7 2 65 1 .9 0 .5 17-7 Total 36? 1 0 0 .0 Cause : : « • agreement w ith Trippensee=/ although o b servation s in d ic a te th at snowshoe damage to northern w hite cedar i s a d e f in it e problem. Not o n ly have th e se s tu d ie s in d ic a te d th at ra b b it browsing i s an im portant ca u sa l agent o f m o r ta lity in advanced cedar rep rod u ction , but i t has a lso in d ic a te d th a t th e snowshoe markedly reduces the alread y lim ite d supply o f deer browse. A p r a c tic e su g g ested by Trippensee (53) f or reducing co n ifero u s p la n ta tio n damage by the varyin g hare might w e ll a id in reducing i t s damage to cedar, i . e . , r e ta in in g the enemies o f the v a ry in g h are, in c lu d in g timber w olves, c o y o te s, bobcats, lyn x, and the g rea t horned and barred o w ls. E ffe c t o f Deer and Snowshoe Hare Browsing Upon Advanced Reproduction and Stand Composition Methods A s e r ie s o f e x clo su r es were e s ta b lis h e d by C iv ilia n Conservation Corps crews during the summer o f 1937 under the d ir e c tio n o f Shaler E. Aldous at the Upper P en in su la Experim ental F o r est, Dukes, Michigan. Each exclosu re covers an area o f 600 square f e e t , liOO square f e e t o f which has been made deer proof and 200 square f e e t o f which has been made deer and snowshoe hare p ro o f. The e f f e c t o f tw elve years ex clo su re o f deer and the combined ex clo su re o f deer and snowshoes on reproduction i s thus a v a ila b le . i^T rippensee (53)» i n d isc u s sin g food p referen ce o f the snowshoe hare, s ta t e s th a t "the id e a 1 cover type f o r the snowshoe i s a m ixture o f hardwoods and c o n ife r s , hardwoods fo r fo o d and c o n ife r s fo r p r o te ctio n " . The four e x clo su r e s were lo c a te d in d if f e r e n t s i t e s . E xclosu re 1 was lo c a te d in S e c tio n 25, T 4-6 N R 23 W and had not been cut in recen t y ea rs. in 1937. E xclosu re 4 was i n an acre hloclc th a t was c le a r c u t i / E xclosure 5 was ad jacen t to i t in an uncut stand in S e c tio n 27 T 46 U R 23 W. S p e cies com p osition o f the stand composing the g en era l type in which E xclosu res 4 and 5 were lo c a te d was^/: S p ecies Stems per acre Oedar Balsam f i r Black spruce Red maple Black ash Y ellow b irch Other spp. 298 172 44 36 3^ 17 17 Range in d .b .h . in in ch es 2 2 2 2 2 3 2 -21 - 13 -13 -13 -12 - l6 -6 E xclosu re J (S e c tio n 23 T 46 H R 23 W) was lo c a te d in an overmature reserv e stand. Oedar stem s 18 in ch es d .b .h . and over are common. A ll ex c lo su r es f a l l w ith in the H ulbert lan d ty p e, predom inantly Carbondale muck. The snowshoe hare p o p u la tio n in the g en era l area i s very high. In the v i c i n i t y o f E xclosu re 1, snowshoes have been th e c h ie f cause o f p la n ta tio n f a il u r e s . an acu te problem in the area. The deer p o p u la tio n i s not Ho browse li n e was p r e se n t. The only fig u r e s a v a ila b le on deer p o p u la tio n in d ic a te th a t the "deer k i l l per square mile" i n the county i s below th e average f o r the Upper P en in su la o f M ichigan. In 1943, th e "deer k i l l per square mile" in Marquette county was 1 .9 5 as compared w ith an Upper ^ S p e c if ic a t io n s in d ic a te d a minimum d .b .h . o f 4 .0 in c h e s. However, »2 and 3 inch d e f e c tiv e stems were a lso cu t. • J This t a l l y was made p r io r to c le a r c u t tin g a p o r tio n o f th e area. P en in su la mean o f 2 .5 3 (2 3 ). A check (UOO square f e e t ) was e s ta b lis h e d adjacen t to each e x c lo su r e . Although steins in a l l the p lo t s were t a l l i e d accord in g to M ichigan C onservation Department Game D iv is io n Browse Study Form ITo. 1 (Appendix D), and r e p lic a t e s were e s ta b lis h e d \id th in each e x c lo su r e , s iz e c la s s e s and r e p lic a t e s were lumped fo r s t a t i s t i c a l treatm en t. I t i s r e a liz e d th a t snowshoe hare and deer browsing co u ld have l i t t l e e f f e c t on th e number o f stem s over 3 in ch es d .b .h . However, th e ir numbers per acre were not s ig n if i c a n t ly la r g e enough to in flu e n c e an a n a ly s is o f snowshoe and deer browsing on th e number o f stems per acre. R e su lts The g en era l com p osition o f stems over 2 f e e t in h e ig h t in each e x clo su r e area r e v e a ls th at cedar i s an important component o f the stand (ta b le 2 3 ) . E xclosure o f deer and snowshoes had no e f f e c t upon the number o f stems o f cedar (ta b le 2U ). Although no red u ctio n in number o f stems due to browsing i s in d ic a te d , damage from snowshoe browsing i s very apparent. For example, in th e v i c i n i t y o f E xclosure 1 and w ith in th e deer ex clo su re p o r tio n o f th a t p l o t , 20 to 25 p ercen t o f the cedar stem s from 2 to S f e e t in h eig h t were s e v e r e ly damaged due to snowshoe browsing. The a s s o c ia t e s o f cedar showed varyin g e f f e c t s o f browsing. Black spruce was not p resen t in la rg e enough numbers to provide an e stim a te . Ho s ig n if ic a n t d iffe r e n c e was noted between the v a rio u s treatm ents as regards the number o f balsam f i r stems per acre. Alder and red maple showed a h ig h ly s ig n if ic a n t l o s s in xu u Table 23. — Stand, ta b le in ex clo su re areas -I (in number o f steins per acre) « • E xclosure area • • S p ecies • • 1 4 Cedar Black spruce Balsam f i r .Alder Red Manle Black ash Other sp. 3594 73 1089 726 2977 109 145 2795 109 1416 5046 2251 3412 1597 980 109 109 9444610 3884 36 1016 73 290 182 2105 36 254 Total 8713 16626 10672 3956 - / Steins two f e e t in h eig h t and over. 5 7 Table 24. — Stems n er acre two f e e t in h eig h t and over • • E xclosure • • Treatment » • : 1 : 4 : 5 : 7 : Mean Cedar Check Deer ex clo su re Snowshoe hare and deer ex clo su re 2173 4900 3703 4792 1193 2396 980 4 36 1525 653 1525 871 2151 2015 2124 2178 3153 9S01 5^)4 5*j4 1742 0 109 436 871 930 3322 2722 2287 1307 1906 2478 4574 .Alder Check Deer ex clo su re Snowshoe hare and deer ex clo su r e 762 109 1307 Red maple Check Deer ex clo su re Snowshoe hare and deer ex clo su re 544 2287 6098 3158 1416 2178 1198 3920 6712 number o f stems per acre due to snowshoe brow sing ( t a b le 2^ ). There i s no s ig n if ic a n t lo s s i n numbers o f stems o f th e se s p e c ie s due to deer brow sing. B lack ash showed no s ig n if i c a n t d iff e r e n c e in numbers o f stems per acre due to browsing. I t i s in t e r e s t in g to note th at in th e c le a r cu t area (E xclosu re 4 ) , snowshoes p r eferred a ld er , red maple, b lack ash, and r e d - o s ie r dogwood to northern w h ite cedar and th e other c o n ife r s p re se n t fo r food . E ffe c t o f Deer Browsing upon Advanced Reproduction and Stand Composition Methods I t has been the op in ion o f numerous f o r e s t e r s and game managers th a t extrem ely heavy browsing o f northern w hite cedar by deer would change the com p etition w ith in a stand to such an ex ten t th a t th ere would be p a r t ia l e x c lu sio n o f northern w hite cedar. A group o f la rg e en clo su res were a v a ila b le f o r study at the Cusino W ild life Experiment S ta tio n . These e ig h t en clo su r e s, one acre each in s iz e , are lo c a te d in variou s typ es o f swamp and hardwood stands and have been su b jected to d if f e r e n t amounts o f deer browsing over the p a st ten y e a r s. C on stru ction i s o f la rg e p o le s w ith two h e ig h ts o f standard snow fe n c e . Figure 36 i l l u s ­ tr a te s a g en era l view o f the e n c lo su re s. The c o r r e la tio n o f cedar reproduction and young stand development w ith deer browsing i s la r g e ly dependent upon an accurate estim a te e f the browse co n d itio n s p r e se n t. A method Figure 36. A "browse p lo t viewed from th e o u tsid e showing th e type en clo su re co n stru c tio n used. The Andronogon fu rca tu s in th e foreground was noted o n ly alon g th e swamp roads. I t was not used "by deer to any exten t inasmuch as th e deer were ranging in th e uplands a t th e tim e th a t the g ra ss would "be o f v a lu e as fo ra g e. o f q u a n tita tiv e comparison o f browse co n d itio n s based on the in form ation acquired by u se o f the I-iichigan Department o f Conservat­ ion Game D iv isio n Browse Study Form Mo. 1 was d ev ised to o b ta in t h is e stim a te. A d isc u s sio n o f the browse a p p r a isa l method i s in ­ clud ed in Appendix D. Davenport e t a l (IS) p rovid e a rather d e t a ile d d e s c r ip tio n o f the browse p lo t s at Cusino th at were u sed in t h is study. However, th e ir d e sc r ip tio n was p rim a rily aimed at a d isc u s sio n of the p lo t s w ith carryin g ca p a c ity in mind and a fu rth er d e s c r ip tio n was f e l t n ecessary. Browse p lo t s 1, 5» 6 , 7« an& 8 were found to be s a tis fa c t o r y fo r the purpose o f th is s t u d y i/. These p lo t s were o r ig in a lly c r u ise d as cedar-spruce-balsam fir-swamp hardwoods w ith heavy sto c k in g , 0 to 6 in c h d .b .h . The stand d e n s it ie s and dates o f estab lish m en t are as follow s:P lo t 1 5 6 7 8 T o ta l Stems per Acre 8680 17360 11680 13980 13720 Date E sta b lish e d 1936-37 1938-39 1938-39 1938-39 193 S-3 9 The c r u is e d ata ob tain ed during the summer o f 19^9 was then analysed fo r "percentage o f a v a ila b le browse". R esu lts ^B row se p lo t s 2 and 3 were excluded from th e study because they were not pure stands o f c o n ife r s . P lo t 2 was a cutover stand of hardwood rep rod u ction and P lo t 3 n mixed swamp type o f both c o n ife r s and hardwoods. P lo t U was d isca rd ed because c r u is e records in d ic a te d a low sto ck in g in comparison w ith the other p lo t s at the tim e of estab lish m en t ( 31 U0 stems p er a c r e ). P lo t Percentage o f a v a ila b le "browse 1 5 6 7 S Thus a c l a s s i f i c a t i o n according to the p resen t "browse co n d itio n would in clu d e P lo t s 5 6 as com p letely "browsed o u t, P lo t S was very h e a v ily browsed, P lo t 1 as medium to heavy brow sing, and P lo t 7 w ith a high p ercen tage o f a v a ila b le browse. I t was d ecid ed th a t one p lo t in each group should be g iv en in te n s iv e study. P lo t 5 was s e le c t e d to rep resen t the co m p letely browsed out p lo t because P lo t 6 had a stream running through or adjacent to s e v e r a l o f the lin e s that might have p rovided a s u b s ta n tia l erro r. P lo t 6 a lso had an extrem ely heavy co n cen tra tio n o f deer p la ced i n i t fo r a s in g le yarding season (10 deer i n 193 ^ - 3 9 )• According to Davenport e t a l (IS ), at the tim e o f p lo t esta b lish m en t, 3& p ercen t o f th e t o t a l stems w ith in P lo t 1 were northern w hite cedar and h i . 3 p ercen t sp eck led ald er and h ig h bush cranberry. On the b a s is o f the commercial s p e c ie s i / p lu s a ld e r , cedar accounted fo r 39*5 p ercen t o f the stems above 2 f e e t in h e ig h t. Deer were browsed i n t h is p lo t throughout i t s e x is te n c e . The t o t a l known deer days^/ o f brow sing from the time o f esta b lish m en t to the summer o f 15^9 i s 2366 . ^ O ed ar, spruce (black and w h ite ), balsam f i r , b la ck ash, and .red maple. 2 / During World War I I , a l l p lo t s except P lo t 7> the c o n tr o l, were clean ed o f snow fen ce such th a t d if f e r e n t ia l browsing was by chance during t h is p e r io d . P lo t 7 remained in t a c t . P lo t 5 had the h ig h e st t o t a l number o f stem s r>er acre at the time o f esta b lish m en t. Oedar accounted fo r 17.S p ercen t o f th e stems o f commercial s p e c ie s and alder above 2 f e e t in h e ig h t. Alder was very heavy in the p lo t (5 8 .0 $ ) and a generous sp r in k lin g o f b la ck ash and red maple was p r e s e n t. P lo t 5 has been browsed s in c e estab lish m en t w ith 1106 days o f in d iv id u a l deer brow sing. P lo t 7 and a ld e r . 25 .8 p ercen t cedar stems o f th e commercial s p e c ie s This p lo t has been a com plete c o n tr o l s in c e 1938-39 w ith th e ex cep tio n o f a 132 pound a d u lt fem ale deer th a t escaped from P lo t S in to P lo t 7 fo r a p er io d not exceedin g 32 days during the 19^9 yarding sea so n . T herefore, in th is p lo t v i r t u a lly a l l the red u ctio n in a v a ila b le browse during the p a st ten years has been due to snowshoe hare browsing and n a tu ra l pruning. P lo t 8 o r ig in a lly was composed o f 30*2 p ercen t cedar stems as c a lc u la te d by th e same c t i t e r i o n u sed fo r th e oth er p lo t s . A t o t a l o f on ly 657 deer days o f browsing are on record . P ract­ i c a l l y th e o n ly food s w ith in th e p lo t a t the tim e o f e s t a b lis h ­ ment were cedar and a 3m all amount o f balsam. The "percentage o f a v a ila b le browse" on the cedar at the tim e o f p lo t esta b lish m en t fo r P lo t s 1, 5» 7» and 8 i s as fo llo w s: P lo t P ercentage o f a v a ila b le browse 1 56.3 5 7 8 ^9.9 5 6 .0 U1.9 R e su lts A ll the p lo t s showed a d ecrease in a v a ila b le browse during the eleven or more yard in g sea so n s. A comparison o f the ta b u la tio n s o f "percentage o f a v a ila b le browse" at th e tim e o f p lo t e s t a b lis h ­ ment and in 19^9 s u b s ta n tia te s t h is f a c t . (The decrease o f 1 0.8 p ercen t in P lo t 7 i s a ttr ib u te d to snowshoe browsing and n a tu ra l pruning. Pigure 37 i l l u s t r a t e s the ex ten t to which th e se fa c to r s can e f f e c t th e fo o d supply. I t should be noted th a t th e re i s an in c r e a se in cedar stems i n a l l the p lo t s during th e in te r v a l from th e time o f estab lish m en t to the summer o f 19^-9 ( fig u r e 3 8 ) . The in c r e a se in stems i s l e a s t in th e very h e a v ily browsed area and becomes in c r e a s in g ly la r g e r as the "percentage o f a v a ila b le browse" becomes la r g e r . Although th e re was no d iffe r e n c e in cedar stems over 2 f e e t in h eig h t a t the time o f p lo t esta b lish m en t, an a n a ly sis o f the 191+9 data shows a s ig n if ic a n t d iffe r e n c e in cedar stems between the p lo t s . There i s a c o r r e la tio n o f .978 between the number o f cedar stems per p lo t and th e "percentage o f a v a ila b le browse" (Appendix A). Table 25 enumerates the cedar reprodu ction from 2 to S f e e t in h eig h t on an acre b a s is as found i n the summer o f 19 ^9 * I* °an again be seen th a t th ere i s an in c r e a s e in cedar rep rod uction as th e "percentage o f a v a ila b le browse" in c r e a s e s . I t i s e s p e c ia lly evid en t in the check p l o t . Table 26 a lso b rin g s out an important c o n sid e r a tio n in swamp eco lo g y . In a l l c a s e s , the number o f ald er stems de­ creased over the elev en year p er io d , in s p it e o f heavy browsing o f the cedar and l e s s d e s ir a b le s p e c ie s in some o f the p lo t s . F igu re 37. Browse p lo t 7 in d ic a tin g a p o r tio n o f th e p lo t in which s e lf-p r u n in g and snowshoe hare "browsing have reduced th e "browse supply. I t i s not meant to imply th a t th e p resen t "browse c o n d itio n shown h ere i s th e r e s u lt o f o n ly th e se two f a c t o r s , however. Deer "browsing, p r io r to th e p lo t e sta b lish m en t, might have been a p a r t ia l cause, e s p e c ia lly in th e ca se o f th e o ld er stem s. N UM B ER OF C E D A R S TEM S IN HEIG H T PER A CRE E S T A B L IS H M E N T AT OVER TWO T IM E OF FEET PLO T AND AT PR ES E NT 6000 Time of Establishment 5000 1949 4000 in z UJ M3000 or o UJ 2000 1000 PLO T F igu re 38. P lo t 5 was com p letely browsed out in 1949, p lo t 8 was very h e a v ily browsed, p lo t 1 has been su b ject to medium to heavy browsing, and p lo t 7 was an unbrowsed check. Table 25. — Cedar re-production from 2 to 8 f e e t in h eig h t on an acre in 194-9 ( in number o f steins per acre) P lo t • • • • : Browse Condition : :Ko Browse • • 5 8 1 7 Yery h e a v ily browsed H eavily browsed Medium browsed No browsing % • 0 0 0 0 Browse Class : 0-33^ : 3h - 6 S i 100 40 180 160 560 600 400 20 • • : 67-1006 460 1720 1780 4800 : Total 1120 2360 2360 4980 Table 26. — Alder stems -per acre 2 f e e t in h eig h t and over at the time o f -plot estab lish m en t and summer 19^-9 (in number of stems per acre) P lo t Date o f t a lly Time of e s t a b lis h ment Summer 19^9 8 6580 6300 3100 5^00 ^060 5580 2580 2880 I t must "be remembered, th a t th e stand as a whole was com p aratively young a t th e tim e o f p lo t esta b lish m en t (0 to 6 in ch c l a s s e s ) , the heavy c u t probably having been 30 to ^10 y ea rs p r e v io u s. The number o f ald er stems has decreased as much as U6 .7 p ercen t in an e le v en year p e rio d . Although a good share o f t h is was probably due to the m o r ta lity o f a ld er stems w ith in th e clumps, c o n sid e r a b le evid en ce was p resen t o f c o n ife r s enroaching upon th e a ld e r clumps. There was no c o r r e la tio n between the "percentage o f a v a ila b le browse" and the in c r e a se or d ecrease i n number o f stem s o f th e o th er s p e c ie s . Upon com pletion o f the stud y, th e data from P lo t 6 was checked f o r comparison. E s s e n t ia lly i t fo llo w e d th e same trends as P lo t 5 — another h e a v ily browsed p lo t — in th a t th ere was an in c r e a s e i n cedar stems' and a decrease in alder stems over th e y e a r s . SILVICULTURAL PRACTICES E ffe c t o f C ertain S ilv ic u lt u r a l P r a c tic e s upon the Reproduction o f Cedar and i t s A sso c ia te s Methods Bob's Lake C u ttings A s e r ie s o f experim ental swamp c u ttin g s were i n i t i a t e d in th e w inter o f 1937 by th e Game D iv is io n , M ichigan Department o f C onservation w ith the co o p era tion o f the M ichigan Emergency C onservation Yfork program Camp Escanaba R iver ( C iv ilia n C onservation Corp). L.A. Davenport was r e sp o n sib le fo r th e p lann ing p h ases w ith variou s fo r e s t e r s and game managers r e sp o n sib le fo r th e f i e l d work in v o lv ed . I t was p o s s ib le to correspond and converse w ith se v e r a l o f th e te c h n ic ia n s r e sp o n sib le fo r th e p r o je c t as to th e conform ity o f f i e l d work w ith the s p e c if ic a t io n s . A checkerboard s e r ie s o f p lo t s each f iv e ch ain s square were la i d out with p ic k e t l i n e s sep a ra tin g the in d iv id u a l p l o t s . A t o t a l o f fo r ty -fo u r p lo t s were e s ta b lish e d in t h is manner. Figure 39 i l l u s t r a t e s th e p lo t s th a t were e s ta b lis h e d and cu t p r io r to the tim e o f photographing (October 1}-, 1939)* F igu re ^40 i l l u s t r a t e s th e p lo t d esig n and in d ic a te s the depth o f the organic s o i l la y e r throughout th e area stu d ied . The p lo t s d esig n a ted as th e "Bob's Lake R elea se Cutting" are lo c a te d in the Sg- o f th e S e c tio n 13 R26w. and the o f the 1TE^ o f the SW^ Odd numbered p lo t s v/ere u sed as check p lo t s w ith no s il v i c u l t u r a l treatm ent. There was no d if f e r e n t ia t io n between s p e c ie s regarding the diameter lim it on the p o r tio n o f the cu t p lo t s fo r which a lim it was d esign ated . A ll tr e e s on the p lo t s below th e s p e c if ie d lim it s \*ere not cu t or pruned in any manner except where n ecessa ry fo r roads or p ic k e t l i n e s . On the p o r tio n o f the cut p lo t s d esig n a tin g a lea v e stand in t r e e s p er acre, th e stems l e f t were good stem s, even ly spaced, and not n e c e s s a r ily the la r g e stem s. Wherever p o s s ib le the merchant­ ab le m a teria l was removed and the le a v e stems s e le c te d from the rem aining stand. The le a v e stems were s e le c te d in the fo llo w in g order o f p r io r it y by sp e c ie s; w hite p in e, northern white cedar, hemlock, balsam f i r , spruce, b ir c h , black ash, red maple, p oplar, JL V^V§. ^ B ><**B f c W ® ? 6 ’.%.vS V• Figure 39. : /.’V - • - C 21* <* \ An a e r ia l photograph o f th e checkerboard d esign in S ectio n 13 I 4^11 R Z6\I. th e c u ttin g s were made. This photograph was taken b efo re a l l CONTOUR MAP BOB'S LAKE CUTTINGS (DEPTH OF ORG. LATER) PT71 plots u t il iz e d in t h is STUDY. Figure *10. Contour map of the Bob's Lake cutting s showing the depth of the organic s o i l la y e r s. The experimental design was a checkerboard pattern o f 5-chain square p l o t s with the various cuttings occurring in the even-numbered p l o t s . and balm of g i l i a d . Stems containing available browse were given preference over stems that had no available browse, either from animal browsing or self-pruning. For in terp retatio n of the sla sh disposal s p e c i f i c a t i o n s , the term "browse species" was defined as including northern white cedar, hemlock, and red maple. Oruises at the time of p l o t establishment were made i n an e a s t west d ir e c t io n through the centers of the p l o t s , thuB covering a s t r i p f i v e chains by ten l in k s . with Browse Study Form ITo. 1. Oruises were made i n conformance The a v a i l a b i l i t y region was considered to be from the ground up to 6 .5 f e e t i n height. The 19^9 cru ise was conducted in a s l i g h t l y d i f f e r e n t manner in order to conform with procedure used elsewhere in t h i s d i s s e r t a t ­ ion. Five temporary l i n e s , each 1 chain in length, were esta b lish ed on each p lo t that was s e l e c t e d for study. eit h er end of the l i n e . Posts were placed at Location of p lo t s was according to the plan i l l u s t r a t e d i n Figure 4 l . Oruises were taken i n 10 l i n k widths. All stems above 2 f e e t i n height were t a l l i e d by browse c l a s s e s . A height of S. 5 f e e t was used as an upper l i m i t o f browse a v a i l a b i l i t y . At the time of establishment, four to s i x p l o t s were cut according to each of the cutting methods i n us e. then r ep lica te d as to slash dis posal. The p l o t s were The cutting methods used ore as follows: 1. 2. 3. 4. 5. Out a l l material over 2" d.b.h. Out a l l m a te ria l over 4" d .b .h . S e l e c t iv e cut leaving approximately 500 stems per acre. S e l e c t iv e cut leaving approximately 750 stems per acre. Uncut check p l o t . IL L U S T R A T IO N O F C RU IS E ON B O B 'S L IN E LAYOUT L A K E C U T T IN G S i o ' HOI " o HO X S X o X o ■ IC H > 1.5 CH. , , 5 : h 1 5 CH . Figure tyl. , j 1CH i S lash d isp o s a l fo llo w e d two s p e c if i c methods: 1 . P i l e and burn th e s la s h from a l l non-browse s p e c ie s as i t i s c u t. S ca tte r th e s la s h from a l l browse s p e c ie s u n t i l th e deer have had an op p ortun ity to fe e d on i t as much as th ey w i l l , then p i l e and burn a l l the rem aining s l a s h i / . 2. P ile and burn the s la s h from a l l th e non-browse s p e c ie s as i t i s c u t. Lop and s c a tte r th e s la s h from a l l browse s p e c ie s so th a t no p a rt o f th e s la s h la y s more than 2M- in ch es above the ground or snow. P lo t s e le c t io n s vrere made on a b a s is o f s i t e and cover ty p e. O ruises were made o n ly on p lo t s f a l l i n g w ith in a d e f in it e swamp cedar-balsam type. With t h is in mind, p lo t s s e le c t e d fo r co n sid e r a t­ io n were th o se enumerated i n ta b le 2 7 In cid en t to the c r u is in g , s o i l borings were made a t the south end o f each l i n e and th e data obtain ed was in corp orated in to a contour map o f the depth o f organic s o i l in the experim ental area. P lo t 32 was found to be a Newton sand type w ith a v ery sh allow organic la y e r and was d iscard ed from the study. The stand com position on an acre b a s is at th e tim e o f p lo t estab lish m en t p o in ts out th e dominance o f cedar (ta b le 2 8 ). A m ilacre p lo t was a lso e s ta b lis h e d at the south end o f each l i n e . Oedar s e e d lin g and v e g e ta tiv e reproduction from 0 to 2 f e e t in h eig h t was t a l l i e d w ith in th e se p l o t s . P lo t r e j e c t io n was a t 2 standard d e v ia tio n s from th e mean or g r e a te r . Ousino C uttings Twelve on e-acre r e le a s e c u ttin g s (2 ch ain s x 5 ch a in s) were l a i d out during the w inter o f 1935-36 and were c le a r c u t. A ll l / These types h e r e a fte r w i l l be d esign ated by th e treatm ent o f th e browse s p e c ie s . Table 27. — P lo ts s e le c t e d fo r comparison o f c u ttin g and s la s h disT>osal methods (P lo t numbers) C utting method :___________ SLflsh d isp o s a l method___________ ____________________________________ ; P ile and burn: No sla s h : Lop and s c a t t e r Cut a l l m a teria l Cut a l l m a teria l S e le c tiv e cut to S e le c tiv e cut to Uncut checlc over 2 in ch es over 4 in ch es 500 steins per 750 stems per d .b .h . d .b .h . acre acre 34, 40 12, 14 — , 16 24, 36, 20, 18, 11.27 26 38 28 32 Table 28. — Stand conroosition in the Bob's la k e c u ttin g s a t the time of e s t ablishment - / (in number of stems p e r a c r e ) • • Size C la sses : Stems 2 to 8 f e e t : Stems 6 in ch es : Total stems over 2 : in heifcht : d .b .h . and over: f e e t in h eigh t • • Species Cedar Snruce Balsam f i r Alder Red Maple Tamarack Aspen 189 2 200 31 51 7 4 158 58 33 0 0 0 0 100? 109 400 138 53 9 16 Total 484 249 1732 1 / Obtained from l / 2 0 t h acre c r u is e s on 9 p l o t s . are located in the W|- of Section 30 Thjll Rl6w. The s o i l type throughout the area i s R i f l e peat; swamp types c o n s t it u t e d the cover. The Land Economic Survey, Michigan Department o f Conservat­ ion, typed the cover i n 1932 as a swamp cedar-spruce-tamarack type with medium stocking, 0 to 3 6 inches d.b.h. No record has "been found as to the amount of material taken o f f each p l o t . Figure 1+2 shows the lo c a t i o n of the p l o t s . Browse procurement p l o t s were cut i n the same manner as the r e le a s e cutti ngs, i . e . a complete c le a r cut. Five l i n e s were e sta blish ed i n the Browse Procurement P lo ts, each 1 chain by 10 l i n k s . A ll reproduction above 2 f e e t i n height was cruised. Results Bob's Lake Cuttings Cedar se edlin g reproduction under 2 f e e t 10 to l 4 years a f t e r cutting i s shown i n table 29. I t can be seen that seecU- l i n g reproduction i s in c re a sin g ly heavier as the i n t e n s i t y of cutting decreased. In a l l cases, there i s a greater average number of se ed lin gs i n the p l o t s in which the browse sp ecies were lopped and scatt ered than i n the p l o t s i n which cedar has been p i l e d and burned a f t er browsing was completed. A good portion of the se ed lings t a l l i e d i n the Bob's Lake p l o t s was i n the 1- and 2-yea r-o ld c l a s s . the reproduction was of t h i s s i z e , however. Not a l l of Seedlings from 10 inches to 2 f e e t in height were frequently noted. Cedar v e g etative reproduction under 2 f e e t i n height was P igu re H-2. c u ttin g s . An a e r ia l photograph i l l u s t r a t i n g the Cusino X tll Table 29* — Average number o f cedar se e d lin g s per i.iilacre p lo t in summer 19^-9 C utting method Slash d is p o s a l method P ile and bum : Ho s la s h : Loft and s c a t t e r Cut a l l m aterial over 2 in ch es d .b .h . Cut a l l m aterial over ^ in ch es d .b .h . S e le c t iv e cut to 5 00 stems per acre S e le c t iv e cut to 750 stems per acre Uncut checlc 1 0.0 2 o.9 2 9 .8 3*0 1 6 .9 J1.6 3 6 .0 55*2 n e g lig ib le i n a l l c a s e s . Out o f 71 p lo ts c r i t i c a l l y examined, on ly S were found to have v e g e t a tiv e rep rodu ction p re se n t. The amount o f cedar reproduction per acre in the 2 to 8 f o o t h eig h t c la s s e s a t the tim e th e p lo ts were e s ta b lis h e d i s as fo llo w s : P lo t Number o f 11 12 0 VO 120 26 27 28 36 3s lK> 100 20 280 220 2140 Mean 135 There i s inadequate cedar rep rodu ction in th e 2 to 8 f o o t h e ig h t c la s s e s under a l l the c u t tin g methods and under both s la s h treatm ents ( ta b le 30)* There i s too l i t t l e reproduction i n th e se s iz e c la s s e s to c o r r e la te w ith any s il v i c u l t u r a l treatm ent. I t appears th a t a heavy c le a r cut fa v o rs ald er in v a sio n . Lopping and s c a tte r in g o f s la s h seems to have an in h ib it in g e f f e c t upon ald er in v a sio n . W ithin the lig h t e r c u ttin g , i . e . lig h t e r than a cut o f a l l m a te r ia l over 2 in ch es d .b .h ., i t would appear th at some fa c to r other than degree o f c u ttin g might in flu e n c e th e in v a sio n o f a ld e r . In a l l typ es o f c u ttin g , the amount o f ald er has in c re a se d over th at o f the uncut check p lo t s (ta b le 3 1 )* Balsam f i r reproduction i s favored by the h eavier cu ts Table 30. — Cedar re-production 2 to 8 f e e t in h eig h t in summer 19^9 (in steins per acre) Cuttins nothod :___________ Slash disposa l method________ _________________________________{Pile and burn? I7o slash; Lot> and sca tter Cut a l l material Cut a l l material Selective cut to Selectiv e cut to Uncut check over 2 inches over k inches 500 stems per 750 steins per d.b.h. d.b.h. acre acre 10 0 0 20 10 0 0 0 JLU*J Table y i . — Alder stems -per acre 2 f e e t in h eig h t and over in summer 19^-9 Slash d isp o s a l method P ile and burn: No s la sh ; Lop and s c a t t er Cut t in s method Cut a l l m ateria l Cut a l l m ateria l S e le c tiv e cut to S e le c tiv e cu t to Uncut check over 2 in ch es over k in ch es 500 stems per 750 stems per d .b .h . d .b .h . acre acre — 3080 3510 5620 7320 1970 2200 5360 250 (table 32). Spruce reproduction seems to fo llow the same trend as balsam f i r (table 33)* The order u sed in th e c l a s s i f i c a t i o n o f c u ttin g s from heavy to l i g h t was v e r i f i e d by ten th -a cre p lo t s w ith in the va rio u s c u ttin g ty p e s. An example o f the conformance to c u ttin g s p e c if ic a t io n s — and probably th e most accurate one — i s shown in P lo t 2S, s e l e c t i v e l y cut to 500 stems per acre. The 19^9 c r u ise produced a t a l l y o f 530 stems per acre 3 in ch es d .b .h . and over. Stems o f sp e c ie s oth er than th ose already d isc u sse d were s c a tte r e d and v a r ie d from p lo t to p lo t rath er than c o r r e la tin g w ith th e d iff e r e n t treatm en ts. On a p lo t w ith a cu t o f a l l m a teria l over 2 in ch es d .b .h . th a t was adjacent to upland aspen (P lo t 2U), a heavy i n f i l t r a t i o n o f aspen from the r e s id u a l stand was n oted (5^0 stems per a c r e ). Willow and r e d -o s ie r dogwood a lso invaded s e v e r a l of th e p lo ts . Very l i t t l e red maple and b la ck ash were t a l l i e d . Gusino C uttings Table 3 ^ summarises the r e s u lt s o f t h is phase o f study. I t can be seen th a t cedar d e f in it e ly i s a heavy component o f the u n d erstory. o r ig in . The reproduction i s m ainly o f a v e g e ta tiv e Counts o f in d iv id u a l stems were not made in th e se p lo ts . Alder i s not a dominant component o f the stand in any o f the c u ttin g s in which i t was not noted in the cover at the time o f esta b lish m en t. Table 32. — 3alssj.n f i r reproduction 2 to 3 f e e t in h e ig h t in summer 1949 (In number of stems p er a cre) C utting method Cut a l l m a teria l Cut a l l m a te r ie l S e le c tiv e cu t to S e le c tiv e cut to Uncut check over 2 in ch es over 4 in ch es 500 stem s per 750 stem s per : Slash d is o o s a l method : P ile and burn: Ho s la s h ' Lop and s c a tte r d .b .h . d .b .h . acre acre 1270 960 1150 460 960 4000 140 290 Table 33- — Snruce reproduction 2 to 8 f e e t in h eig h t on an acre b a s is in summer 1949 (In nvunber o f stems) Slash d isn o sa l method P il e and burn; ITo sla sh : lo p and s c a tte r C utting method Cut a l l m a teria l Cut a l l m a teria l S e le c tiv e cut to S e le c tiv e cut to Uncut check over 2 in ch es over 4 in ch es 500 stems per 750 stem s per d .b .h . d .b .h . acre aci'e 100 240 0 220 260 110 0 KEY 'TO SYMBOLS III TABLE A A lder 3 Balsam f i r 0 northern S Spruce T Tamarack w h ite cedar Good s to c k in g Modiiu:i r.to ckinp rO'T.1 Table 3^- *— Ou3ino c u ttin g s summary • • • » R elease c u ttin g p lo t 1 « • : Cover : Cover : 1935 : 19^9 • • • • ! : Cover ; ad jacen t : J to p lo t •• • • • • • • 1/AS' * 2 /ART C' • • • • Deer use ; Snowshoe hare : use • • • • • • SC" Medium Heavy 2 CSA«« BTS C" CSB Heavy Heavy 3 ASTC ASBT C" CST Heavy Heavy k A3S 3TA C" CST Heavy Heavy 5 - ST C ''' CST Heavy Heavy 6 - ACB C' CS Heavy Heavy 7 ASCT'' ABT C' SCA Medium L igh t 8 AC c a (ts ) CS" Medium Heavy c' 1 ' ACS' » ASB O' CS Heavy Heavy 10 C" CTB C ''» CST L ight L ight ll CAS'' SBT C" ' CS Heavy Heavy 12 - V - Heavy - 9 3»/ Symbol key i s included on opposite page. 2 / West & o f P l o t 12 AS i e a s t £ TSB . 3 / Overstory cover i s above the l i n e ; u n d e rsto ry type below the l i n e . S la sh p i l e s d e f i n i t e l y lim it rep rod u ction . cedar rep rod u ction w ith in th e ir b ou nd aries. out t h i s o b se rv a tio n . There was no P igure b rin gs Deer u se was noted in a l l th e areas and runways extend from p lo t to p l o t . Deer browsing has elim in a ted w h ite b ir c h and b la ck ash rep rodu ction in s e v e r a l o f th e a rea s. The reproduction on an area th a t was c le a r cut in the same manner as the oth er r e le a s e c u ttin g s i s as fo llo w s : S n ecies Stems per acre Cedar Spruce Balsam f i r Alder Tamarack White b ir c h Red maple B lack cherry R. 0 . dogwood W illow Amalanchier Aspen 5960 520 200 15*10 160 20 60 20 200 30 20 20 I t can be seen th a t an adequate amount o f cedar reprod uction i s p r e s e n t. Alder has not invaded th e area very s tr o n g ly . O bservations in d ic a te d th a t th e cedar rep rod u ction was predom inantly v e g e t a tiv e . P la n ta tio n s Methods P la n ta tio n s o f northern w hite cedar were made during the f a l l o f 19^1 and the sp rin g o f 19*+2 in th e study area. The 19^1 p la n tin g s were made p rim a r ily fo r in c r e a s in g deer food and cover. The 19^2 p la n tin g s were an attem pt to convert a spruce-balsam type to cedar Pigure 43. An area c le a r-c u t in 1939-*JO. r e la t iv e ly young at th e time of c u ttin g . was v e g e ta tiv e in o r ig in . The stand was The cedar rep rodu ction F igu re *A. A s la s h p i l e in a 1935-36 c u t tin g . R ed -osier dogwood i s the o n ly woody s p e c ie s th a t has been a b le to reproduce w ith in th e s la s h p i l e . Photograph taken in J u ly , 19*19. "by p la n tin g r e c e n tly cu t-o v er area s. The 19^1 p la n tin g s (SB£ and UBjf o f S e c tio n 19 and SB^ o f S e c tio n 8 T^7U R l 6 w) were made w ith 3 -2 p la n tin g sto c k , 9 in ch es i n h e ig h t, produced a t the Hardwood S ta te ITursery, Boyne B a lls , M ichigan. Borty thousand stems were p la n te d "by C iv ilia n C onservation Corps p erso n n el w ith Michigan p la n tin g "bars a t a 6 fo o t x 6 fo o t sp a cin g . No s o i l p rep aration was made. The s o i l typ es p la n te d were Saugatuck sand and R i f l e p e a t. The 19^2 p la n tin g s were made from the same age sto ck u sin g s im ila r p la n tin g methods. th e p la n tin g s . However, l o c a l lab or was u sed to make The s o i l types (S ectio n s 16 and 17 T^7N R l 6 w) were R i f le p ea t and Oarbondale muck. R esu lts The 19^1 p la n tin g s proved f a ilu r e s . Although i t i s d i f f i c u l t to d is t in g u is h n a tu ra l reproduction from p la n te d sto c k in p o r tio n s o f the area, v ery l i t t l e in d ic a tio n o f p la n ted sto ck was l e f t . N atural rep rod u ction i s p resen t in p o rtio n s o f the area not covered w ith an ald er typ e. The attem pt to con vert from a spruce-halsam type to a cedar type was a lso a com plete f a i l u r e . Adequate reproduction o f balsam f i r and b la ck spruce was p r e s e n t. The cedar tra n sp la n ts th a t have su r v iv ed show heavy deer and rab b it browsing. The f a ilu r e o f cedar p la n tin g s i s not co n fin ed to the Upper P e n in su la o f M ichigan. Aldous ( 3 ) rep o rts th e f a ilu r e o f cedar p la n ta tio n s in M innesota and concludes th at i t i s not p r a c t ic a l to p la n t w hite cedar as i t i s too s e v e r e ly browsed by d eer. The The o n l y s u c c e s s f u l p la n t in g s w ere th o s e tuxa d e e r - p r o o f e x c l o s u r e s . cecta r w i t h i n t h e e x c l o s u r e s p r o te c te d oy n a r e - E ig h t y - t h r e e p e r c e n t o f th e w h ite s u r v iv e d a g r o w in g s e a s o n s i n com ­ p a r is o n w ith 18 p e r c e n t s u r v iv a l o u t s i d e . DISCUSSION R e p r o d u c t io n The d i v e r s e e n v ir o n m e n t a l c h a r a c t e r i s t i c s swamps s h o u ld p r o v i d e an o p p o r t u n i t y f o r o f v a r ia tio n m o r ta lity . in th e r e la t iv e The v a r i e d sa n d and f i n d o ff a ffe c te d n a tu r e o f th e had 0 t o 80 p e r c e n t o f t h e to be r e l a t i v e l y m ig n t a l s o r e a c t to swamp f l o o r to (3 1 ) . r e p r o d u c tio n . in v o lv e d , sp ru ce, on c o a r s e e p ip h y to tic s in A lt h o u g h i t d am ping o f f , in e s p e c ia lly ced ar b e tw e e n wne T h ese w ere n ot th e in t a l l y i n g v e g e ta tiv e The p r e d o m in a n t t y p e o f v e g e t a t i v e r e p r o d u c t i o n , uranch la y e r in g , is t h e p a r e n t stem t o in e ffe c t a t r a n s f o r m a t i o n from a b ra n ch o f a stem o f r e p r o d u c t i o n . d e g r e e o f ju d g m e n t t h a t c o u ld b e q u e s t i o n e d . T h is i n v o l v e d a H o w ev er, t h e d e b a t a b l e c a s e s w e r e fe w i n c o m p a r is o n w it h t h e t o t a l number of is a s i m i l a r m a n n e r. d iffe r e n tia tin g t y p e s o f r e p r o d u c t i o n h a v e b e e n m e n t io n e d . o n ly d i f f i c u l t i e s b la c k w h e r e a s on a muck m e d ia dam pin g r e s is ta n t in v o lv e d o f f e r s u s an 2 p e r c e n t d am p in g o f f s e e d lin g s d am pin g o f f The d i f f i c u l t i e s f a c t o r s c a u s in g For i l l u s t r a t i o n , sa n d r e s p e c t i v e l y c o n s id e r e d a c o n s i d e r a b l e am ount im p o r t a n c e o f t h e e x a m p le o f t h e p o s s i b i l i t i e s . a n o t h e r swamp s p e c i e s , o f th e n o rth ern s te m s t a l l i e d . C u rtis (15) g iv e s f u r t h e r su p p o rt to the th e o r y t h a t a d u lt s t e m s o f s e e d l i n g o r i g i n may p o s s e s s v a r y i n g d e g r e e s o f sweep in h is d is c u s s io n s o f old f i e l d esta b lish ed from s e e d . varyin g d e g r e e s, stem s, a l s o adds t h a t cedar i s wood and grow r i g i d l y “Old f i e l d i s l a r g e l y due t o snow, or r o o t a n c h o r a g e 11. . . crowding, Watson ( 5 6 ) a p p a r e n t l y q u i c k t o fo rm c o m p r e s s i o n i n t o a curved stem . p h o t o t r o p i s m s a s an e x p l a n a t i o n f o r He a l s o m e n t i o n s some o f t h e stem c u r v a t u r e s . The a b o v e c i t a t i o n s a r e p r e s e n t e d i n an e f f o r t w idespread f a l l a c y stands . . . many i n d i v i d u a l s h a v e sw ee ps o f . "but t h a t t h i s occurrence o f m u ltip le cedar. to c o r r e c t a co n c er n in g cedar r ep ro d u c tio n ; a curved stem a b s o l u t e l y c la ssifie s namely, that a t r e e as b e in g o f v e g e t a t i v e o rig in . The e f f e c t o f l i g h t upon g e r m i n a t i o n i n t h e l a b o r a t o r y su b stan tiates t h e o b s e r v a t i o n s made i n t h e f i e l d . or near a b s e n c e o f l i g h t red uced g e r m in a t io n v i g o r . extrem e, i.e . fu ll lab oratory r e s u l t s . su n lig h t, field An a b s e n c e At t h e o t h e r s t u d i e s a g a in confirm ed S e e d l i n g g e r m i n a t i o n and i n i t i a l d e v e l o p m e n t w e re m a r k e d l y r e d u c e d i n b o t h s e r i e s . u p p e r end o f t h e t a b l e w i t h f o o t The c o n d i t i o n s a t t h e c a n d l e p o w e r s a b o v e 33^ were found i n th e 1 0 - y e a r - o l d c u t t i n g s t h a t were removed under a system o f com plete c l e a r cu ttin g. The e f f e c t somewhat t h e o f a c i d i t y under l a b o r a t o r y c o n d i t i o n s f o l l o w s sa n e t r e n d a s i n t h e swamps. The d e c r e a s e i n g e r m i n a t i o n and e s t a b l i s h m e n t a t t h e l o w e r e x t r e m e s f o l l o w s i n both c a s e s . The d e f i n i t e tren d o f i n c r e a s i n g g e r m in a tio n through­ o u t t h e a c i d i t y r a n g e m a n i f e s t s i t s e l f a bov e a p p r o x i m a t e l y pH 6 . 0 u nder f i e l d th e x ie ld c o n d itio n s . r e s u lts in I t w as n o t e d pH r e p l i c a t e s (a g a r) th is e x h ib ite d th e la b o r a to r y s u b s ta n tia te s r e sp e c t. in th e la b o r a to r y one u p p e r v a l u e s . m ed ia Tne t r e n d i n e x p e r im e n t s t h a t t h e lo w e r d if f e r e n t d eg rees o f g e la tio n A d iffe r e n c e in h a rd n ess c o u ld c a u se c h a n g e s in T h is p h en o m en o n m ig h t e x p l a i n th a n d id o f t h e g e r m in a t io n th e a v a i l a b i l i t y th e la b o r a to r y r e s u l t s o f m o is t u r e . or a t l e a s t i n d i c a t e t h a t pH c a n be o f c o n s i d e r a b l e c o m p l e x i t y — e v e n u n d er c o n tr o lle d c o n d itio n s . The e x p e r i m e n t a l d e s i g n u s in g c o n s ta n t tem p era tu re te m p era tu res. A lt h o u g h i t in t h e l a b o r a t o r y w as i n i t i a t e d a s a b a s e r a t h e r th a n a l t e r n a t i n g is a c k n o w le d g e d t h a t an a l t e r n a t i n g t e m p e r a t u r e • e x p e r im e n t w o u ld h a v e m ore c l o s e l y c o n d itio n s of seed uuced v a r i a b l e s It is r e a liz e d g e r m in a tio n th a t r e q u ir e in nave th e e f f e c t o f t e m p e r a t u r e a r e more f a v o r a b l e o f lo w e r in g in v a r y in g th e th r e s h o ld o f cedar sta g e s o A p x a m e d on a m o i s t u r e h y p o t h e s i s . is le s s flu c t u a t io n are u s u a lly th e lo c a te d swamp f l o o r w o u ld h a v e i n t r o - o f s e e d t h a n c o n s t a n t t e m p e r a t u r e s and m ig h t The m arked p r e f e r e n c e a m edium o f l o g s t h e swamp, i t th e u se o f h ig h - c o s t f a c i l i t i e s . th a t a lte r n a tio n s f o r th e g e r m in a tio n a p p r o x im a te d o f th e w a ter at a s lig h t s e e d lin g s Ib i s r e a liz e d fo r th a t th e r e decayed lo g s . c o n s id e r a b le and t h u s a r e r e l a t i v e l y t h a t o c c u r d u r in g r a i n y p e r i o d s . in th e f i e l d o f d e c a y c a n h a r d ly be s u p p ly i n to o f g e r m in a tio n . free They e le v a tio n above fro m t h e i n u n d a t i o n s The m o is t u r e s tu d ie s d u r in g t h e 1 9 4 9 g r o w in g season a c t u a lly show ed a g r e a t e r a v e r a g e w e e k ly v a r i a t i o n fro m t h e m ean f o r l o g s d u r in g t h e g r o w in g s e m i­ s e a s o n t h a n d id t h e swamp f l o o r , how ever. C u r tis i ' u i t e n w ood a s a medium i n w h ic h m o is t u r e i s (1 6 ) d e s c r ib e s m ore or l e s s c o n sta n t b en ea th th e f o r e s t ca n o p y . T h e r e w as a d e f i n i t e tw o t y p e s d iffe r e n c e te m p era tu re. th r e s h h o ld s e a s o n and a g r e a t e r a l t e r n a t i o n The h i g h e r mean t e m p e r a t u r e i s d u r in g J u n e an d J u l y , fo r th e p e r io d ( 5 7 .2 ° in g e r m in a tiv e c a p a c it y tem p era tu re o f th e o f s e e d g e r m in a tio n . to 6 4 .4 ° F .) at th e se swamp f l o o r The swamp f l o o r lo g s in v a r io u s d id n o t r i s e is sta g e s of decay, c a n an d d o e s a c c o u n t f o r how ever d e n s e , p o s s ib ility r e la tiv e ly in e le v a tio n in a p o r tio n of it . T h is v a r i a t i o n an a r e a a s I t w as a l s o d r y d u r in g p e r i o d s in s m a ll a s a m i l a c r e . r e g a r d in g sphagnum b e d s . s till if and m ounds o f It is th e lim it e d fe lt th a t th e fo r m s a r a t h e r o p e n t y p e medium is g rea t. th a t th e a p p a ren t r e d u c tio n not as lik e ly s lig h tly . and p i t s s p a c e o c c u p i e d by a i r t o ■chis f a c t o r . o n ly s tu m p s , S e v e r a l h y p o t h e s e s may b e o f f e r r e d o f s e e d lin g s d ecrease The m o n th ly mean a r e m a r k a b le d e g r e e o f v a r i a t i o n t e m p e r a t u r e and m o is t u r e w i t h i n i n w h ic h t h e Tne ab ove th e u pper l i m i t s e x c e e d in g ly v a r ia b le o r g a n ic m a t e r ia l a r e c o n s id e r e d sph agnu m , n o te w o r th y w ith a d e f i n i t e te m p e r a tu r e s. t h r e s h h o l d u n t i l A u g u s t and t h e n g e r m in a tio n e s p e c ia lly of s e e d g e r m in a tio n a t a c o n s t a n t te m p e r a tu r e f a l l s b e tw e e n 1 4 ° an d 1 8 ° C. is t e m p e r a t u r e b e tw e e n t h e o f m e d ia , t h e l o g m edium s h o w in g a h ig h e r mean te m p e r ­ a t u r e t h r o u g h o u t t h e g r o w in g o f th is in o b serv ed T here i s th e o f g e r m in a t io n i s due t h a t sphagnum m a ts becam e of lit t le r a in fa ll. H o w ev er, t h i s t o b e t h e c a u s e o f r e d u c t i o n i n g e r m in a t io n and e s t a b l i s h m e n t a s i t w as fo u n d t o occur la t e r in t h e g r o w in g s e a s o n x.nan t h e t im e a t w h ic h g e r m i n a t i o n and e s t a b l i s h m e n t n o r m a lly occur. J A l t h o u g h damage b y r o d e n t s and snow shoe h a r e i s a s e r i o u s problem i n n o r th e r n w h it e cedar r e p r o d u c t i o n , it i s not f e l t t h a t p o i s o n i n g i s n e c e s s a r y a s a c o n t r o l m ea s u r e u n l e s s t h e e f f e c t o f sn o w s h o e s up on a d v a n c e d r e p r o d u c t i o n m ig ht a l s o reach a c r i t i c a l tio n s in these stage. H o w ever, categories r e d u c t i o n o f anim al p o p u la ­ s h o u l d be g i v e n due w e i g h t i n c u t t i n g and s l a s h d i s p o s a l m e t h o d s . The masked s h r e w , a c c o r d i n g t o B u r t ( 1 0 ) , i n s e c t i v o r o u s and c a r n i v o r o u s a n i m a l . g r a s s e s and s e d g e s * and i n s e c t s . runways. c h i e f l y an The meadow v o l e e a t s chipmunk s e e d s , The f o o d o f t h e r e d - b a c k e d v o l e alth ou gh C u rtis (16) resp on sib le the l e a s t is concludes th a t t h i s nuts, b erries, i s n o t known sp ecies is p a r tia lly f o r s e e d l i n g l o s s e s by c u t t i n g s e e d l i n g s i n t h e The s h o r t - t a i l e d w e a s e l (M ustella c ic o g n a n ii c i c o g n a n i i B onaparte) has been o b ser v ed i n th e v i c i n i t y t h e p l o t s a l s o bu t i t s of c a r n i v o r o u s h a b i t s w o u ld l e a d t o t h e c o n c l u s i o n t h a t i t w o u ld r e d u c e t h e mouse p o p u l a t i o n r a t h e r than th e cedar s e e d l i n g p o p u l a t i o n . B o t h f r e q u e n c y and abundance i n d i c a t e that n eith er l i g h t , pH nor t y p e o f g e r m i n a t i n g media a r e c r i t i c a l f a c t o r s in the e s t a b lis h m e n t o f v e g e t a t i v e r e p r o d u c tio n throughout the range o f c o n d i t i o n s found i n t h e swamp s t a n d s i n v e s t i g a t e d . Indica­ t i o n s show t h a t a m o i s t g e r m i n a t i n g medium, e s p e c i a l l y sphagnum moss, d e f i n i t e l y a i d s v e g e t a t i v e r e p r o d u c tio n . few commercial s p e c i e s o f f o r e s t In so fa r as t r e e s reproduce v e g e t a t i v e l y , no m o r t a l i t y s t u d i e s hav e b e e n f o u n d i n t h e l i t e r a t u r e . Management It is con clu d ed t h a t s e e d l i n g r e p r o d u c t i o n under k f e e t i n h e i g h t i s u n a v a i l a b l e to deer a s w in te r fo o d i n the r e g io n under s tu d y . T his l e a d s to the d e c i s i o n t h a t l i t t l e i f any "browse can he p r o d u c e d by c e d a r s e e d l i n g r e p r o d u c t i o n i n l e s s t h a n 20 y e a r s u n d e r swamp c o n d i t i o n s t h a t c o v e r a good p o r t i o n o f the d eer yards in th a t area . It is r ela tio n to v e g e t a t i v e r e p r o d u c tio n because i t br o w s e i n o rig in . e v i d e n t t h a t more r e s e a r c h s h o u l d be c o n d u c t e d i n can p r o d u c e swamp s t a n d s more r a p i d l y t h a n t h a t o f s e e d l i n g V e g e t a t i v e r e p r o d u c t i o n up t o 5*0 f e e t in h e ig h t p r o b a b l y w i l l be w e i g h t e d down by i c e and snow and t h u s c o v e r e d during the w in te r . in d ica te A l t h o u g h t h e a g e c l a s s d e s i g n a t i o n may slow growth, t h i s i s not n e c e s s a r i l y the case because a g o od number o f t h e y e a r s r e p r e s e n t e d may h a v e b e e n a s a b r a n c h and n o t a s a b r a n c h l a y e r . The i n c r e a s e o f t h e number o f c e d a r s t e m s p e r a c r e i n t h e C u sin o b r o w s e p l o t s th a t accom panies the v a r y in g d eg r ees o f b r o w s i n g d o e s n o t h o l d t r u e i n t h e Dukes e x p e r i m e n t s . A l t h o u g h t h i s a p p e a r s t o be a c o n t r a d i c t i o n o f d a t a , s a l i e n t p o i n t s are e v id e n t . T here i s a d i f f e r e n c e i n t h e a g e s o f t h e s t a n d s a t Dukes and a t C u s i n o . a ll The C u s i n o p l o t s a r e compose d o f a y o u n g s t a n d w h e r e a s t h e r e from v e r y y o u n g r e p r o d u c t i o n ( B x c l o s u r e k ) s t a n d ( E x c l o s u r e 7 ) a t Du kes. several It i s a gradation t o an o v e r m a t u r e i s a ls o e v id en t that deer o v e r - p o p u l a t i o n h a s n o t b e e n s e r i o u s a t Dukes b u t h a s b e e n a t C usino. The c o n c l u s i o n s regard in g the and d e c r e a s e o f a i d e r , con d ition s that in overbrowsed, a la st w ill w ill resort. cedar stem s e v e n u n d e r medium and h e a v y "browsing do n o t a g r e e w i t h t h e He c o n t e n d s in crease o f con clu sion s tim e a good d e e r y a r d , if o f Sw ift (51). con stan tly g r o w up t o p l a n t s w h i c h d e e r e a t o n l y a s "Thus - — w h i t e cedar, m o u n t a i n m ap le . . . be r e p l a c e d by t a g - a i d e r , aspen, and r a s p b e r r y " . In sum m ation, of s ilv ic u ltu r a l p ra ctices the r e s u l t s the stu d ied in d ic a te : 1. Cedar s e e d l i n g r e p r o d u c t i o n i s g r e a t e r w i t h l o p p i n g and s c a t t e r i n g o f s l a s h t h a n w i t h p i l i n g and b u r n i n g . 2. I n i t i a l s e e d l i n g r e p r o d u c t i o n i n c r e a s e s w i t h an in c r e a s in g r e s id u a l stand. 3 . The a g e o f t h e s t a n d w i l l d i c t a t e t h e amount o f v e g e t a t i v e r e p r o d u c t io n to a l a r g e d e g r e e . H e a v i e r c u t s f a v o r b o t h a i d e r i n v a s i o n and b alsam r e p r o d u c t io n . The r e s u l t s d ifferen t of th is approaches in p u r p o s e s o f management. a seed lin g the p o s s i b i l i t y s i l v i c u l t u r a l methods It is felt that of fo r variou s co n d itio n s favorin g t y p e o f r e p r o d u c t i o n s h o u l d be s o u g h t f o r i n m anage men t f o r com m ercial p o l e , whereas a v e g e t a t iv e fa v o ra b le study i n d i c a t e post, and c a b i n l o g p r o d u c t i o n t y p e o f r e p r o d u c t i o n w o u l d be t h e m os t f o r game m an ag em en t p u r p o s e s . m ust be c o n s i d e r e d w i t h t h e same p o i n t s operator’s in ter est in creased lies game m a n a g e r ’ s i n t e r e s t 1. in In term ed iate i n m ind. stem g r o w th . cuts The c o m m e r c i a l The c e n te r s about; in c r e a s e d f o l i a g e growth w it h in th e h e i g h t c l a s s e s t h a t a r e a v a i l a b l e t o game a n i m a l s a s b r o w s e . 2 . red uction of se lf-p r u n in g . 3 . an a i d t o rep rod uction . I f the c o n d itio n ex p r essed in ta b le b h o ld s tru e in the m a j o r i t y o f swamp s t a n d s , d escrib e t h e d e s i g n a t i o n " c l e a r out" u s e d t o t h e v a r i o u s t y p e s o f c u t t i n g m eth o ds can n o t he u s e d s y n o n o m o u s l y a s e x p r e s s i n g a m ethod o f r e p r o d u c t i o n ( 2 9 ) . fact The t h a t a p p r o x i m a t e l y 65 p e r c e n t o f t h e r e p r o d u c t i o n t a l l i e d o r i g i n a t e d from l a y e r i n g would c h a n g e th e c l a s s i f i c a t i o n meth od t h a t i s u n d e s c r i b e d i n t h e l i t e r a t u r e . to a I t i s proposed t h a t two new m ethods o f r e p r o d u c t i o n be added t o f o r e s t t e r m in o lo g y to d e s c r i b e t h e r e m o v a l o f th e o l d s t a n d and t h e e s t a b l i s h m e n t o f t h e new crop by t h i s The " la y e r in g m ethod” i s m ain ly o f l a y e r s . p rin cip a lly of s e e d l i n g s and o t h e r stand i s The cut a m e t h o d c o m p o s e d w h o l l y or The r e n e w a l o f t h e by l a y e r s . type o f r e p r o d u c tio n . A lthough there forest is types o f v e g e ta tiv e rep rod u ction . The clear. " l a y e r i n g - w i t h - s t a n d a r d s method " e n t a i l s m a in l y from l a y e r s , the accom p lish ed u s u a l l y a m ixture in g o f sta n d a rd s above a l a y e r i n g f o r e s t . or — is but i n o t h e r w o rd s — the a rea i s a resid u al never the m a in ta in R eproduction i s cu tclea r. stand i s left Standards standing at end o f e a c h r o t a t i o n . It is felt th a t th e d e s ig n a t io n o f the r e p r o d u c tio n m e th o d a s a " l a y e r i n g method" o r " l a y e r i n g - w i t h - s t a n d a r d s m ethod " i s in use. more t h a n m e r e l y c h a n g i n g t h e name o f t h e m e th o d A la y erin g t y p e o f r e p r o d u c t i o n i n v o l v e s a new p h ilosop h y o f c o n i f e r o u s management i n t h e La ke S t a t e s a n d em phasizes con cep ts th a t are q u ite cu ttin g, seed -tree, sh elterw ood , d ifferen t from th e clear and s e l e c t i o n m e t h o d s . M o n t g o m e r y ( ^ 2 ) h a s made s t r i d e s m a n agem en t p r a c t i c e s feet hy s u g g e s t i n g i n t h e d i r e c t i o n o f new that t o p p i n g o f c e d a r k t o 10 in h e ig h t approxim ately at b r e a s t h e ig h t w i l l greater t o t a l number o f b r a n c h e s , branches, greater a n d more b r a n c h l a y e r i n g I t wag d i f f i c u l t se e d lin g s per acre Hawley ( ^ 9 ) a d v ises to that total in a len gth of t h a n an u n t o p p e d c h e c k . o b t a in an e s t i m a t e co n stitu te resu lt o f t h e number o f an a d e q u a te the e s ta b lish m e n t stock in g. of at le a s t 2 ,0 0 0 to 5*000 w e l l - d i s t r i b u t e d young p la n t s per a cre as adequate in d ir e c t seed in g . of are A ssum ing t h e s e cedar s e e d l i n g s to co n stitu e i n a m ixed c o n i f e r o u s th a t a cut o f a l l adequate swamp, m a teria l over 2 in ch es w ould be u n s a t i s f a c t o r y sele ctiv e fig u re s stock in g in d ication s or 4 in c h e s d .b . h . a s a r e p r o d u c t i o n method w h ereas a cut w ith a r e s id u a l s ta n d o f 500 or 750 stem s p e r a c r e w o u l d b e more s a t i s f a c t o r y . E vidence a ls o p o in t s to the p o s s i b i l i t y young sta n d s w ith the r e p r o d u c tio n o f means. I n m a t u r e an d o v e r m a t u r e of h eavier c e d a r by v e g e t a t i v e stan d s, a lig h t w i t h a t h i n n i n g from b e l o w m igh t be a d v i s a b l e is cu ts in if sele ctiv e cut t h e management f o r b r o w s e p r o d u c t i o n and t h e y o u n g s t e m s a r e d e v o i d o f browse. W indthrow, b o t h o f c ed a r and i t s a sso cia tes, e n c o u n t e r e d i n h e a v y c u t t i n g B an d a d j a c e n t t o be c o n s i d e r e d i n d e t e r m i n i n g a c u t t i n g m ethod. them. was I t must SUMMARY AND CONCLUSIONS N o r t h e r n t v h i t e c e d a r i s a s s u m i n g more i m p o r t a n c e i n t h e La^e S t a t e s e a c h y e a r . organic Although i t o c c u r s on b o t h m i n e r a l and s o i l t y p e s i n t h e e a s t e r n h a l f o f t h e Upper P e n i n s u l a or M i c n i g a n , th is study i s m ainly concerned v i t h swamp s t a n d s . L a b o r a t o r y s t u d i e s w e re c o n d u c t e d on v a r i o u s f a c t o r s a f f e c t ­ ing g e rm in a tio n . seed c o l l e c t e d con d ition s. I n d i c a t i o n s are that la b o r a to r y germ ination of i n swamp s t a n d s i s low un de r good g e r m i n a t i n g T e m p e r a t u r e s 1 8 ° C. and b e l o w had an a d v e r s e e f f e c t upon s e e d g e r m i n a t i o n and s e e d l i n g v i g o r . A trend of in c r e a s e in germ ination w ith a d e crea se in hydrogen-ion c o n c e n tr a tio n ttDOve pH 6 . 0 was a p p a r e n t , b u t became l e s s germ ination p eriod in c r e a s e d . p r o n o u n c ed as t h e No s i g n i f i c a n t d i f f e r e n c e was i o u n d i n t h e amount o f g e r m i n a t i o n on f o u r n a t u r a l t y p e s o f m edia: litte r , swamp f l o o r , l o g s i n v a r y i n g s t a g e s o f d e c a y , hardwood and m i n e r a l s o i l a s a c h e c k . F u ll su n ligh t in h ib ite d g e r m i n a t i o n and p a r t i a l l i g h t was more s a t i s f a c t o r y t h a n d a r k n e s s . F i e l d o b s e r v a t i o n s and e x p e r i m e n t s c o n f i r m e d t h e l a b o r a t o r y resu lts e x ce p t as n o ted . seed i s p r e s e n t i n swamp s t a n d s f o r r e p r o d u c t i o n a l t h o u g h v ia b ilib y i s low . F ield stu d ies in d ic a te that su fficien t The p r e p o n d e r a n c e o f s e e d l i n g e m e r g e n c e o c c u r r e d betw e en t h e 5 t h o f June and t h e 5 t h o f J u l y ; e a r l i e r germ ination o c c u r r e d on p a r t i a l l y d e c a y e d l o g s and stumps tha n on t h e swamp flo o r. factor R e s u l t s p o i n t o u t t h a t l i g h t i n t e n s i t y was n o t a c r i t i c a l i n s e e d l i n g g e r m i n a t i o n and i n i t i a l developm ent th ro u g h ­ o u t t h e n orm al r a n g e o f l i g h t c o n d i t i o n s f o u n d i n our swamp s t a n d s and c u t t i n g s a l t h o u g h t h e r e i s a d e c r e a s e in both a b u n d a n c e and f r e q u e n c y a t t h e u p p e r and l o w e r e x t r e m e s . Tne a b u n d a n c e and f r e q u e n c y o f s e e d l i n g r e p r o d u c t i o n i n c r e a s e s w i t h & d e c r e a s e in h y d ro g e n -io n c o n c e n t r a t i o n above approxim ately pH 6 . 0 . Hoy/ever, adequate g erm in a tio n occurred throughout the normal r a n g e o f pH v a l u e s f o u n d i n c e d a r s t a n d s . d istan ce Maximum s e e d i n g o b s e r v e d was 1 5 8 . 0 f e e t w i t h 4 0 t o 50 f e e t a p p e a r i n g t o p rovide adequate in su ra n ce of a seed supply. Unuer f i e l d co n d ition s, lo g s in v ariou s t n e p r e f e r r e d t y p e s o f g e r m i n a t i n g m ed ia f o r xound t o s t a g e s o f decay are s e e d l i n g s and were o c c u p y a b o u t 16 p e r c e n t o f t h e swamp gr ound a r e a . The s e e d l i n g s g e r m i n a t i n g upon them d e v e l o p e d b e t t e r r o o t s y s t e m s t h a n t h o s e g e r m i n a t i n g upon t h e swamp f l o o r . t h r o u g h o u t t h e g rov/ing s e a s o n i n t h i s w ith the sv/amp f l o o r i s Harawood l i t t e r media, t y p e o f medium as compared t h o u g h t t o be t h e r e s p o n s i b l e f a c t o r . and sphagnum b e d s w e re n o t s a t i s f a c t o r y g e r m i n a t i n g t h e hardvfood l i t t e r coo d r y . A h igh er tem peratrue — predom inantly a ld e r le a v e s — b e in g A n o t i c e a b l e a b s e n c e o f s e e d l i n g s was f o u n d i n d e e r runways. A pproxim ately 8 .5 p e r c e n t o f th e l o u n d t o be k i l l e d and 11*5 p e r c e n t damaged by r o d e n t and snov/shoe hare brow sing in a s i n g l e y e a r . root r o t, c o v e r i n g by l i t t e r develop m en t, moss s e e d l i n g p o p u l a t i o n was D esiccation , and d u f f , la te spring f r o s t s , inadequate r o o t system and c o m p e t i t i o n from h e r b a c e o u s c o v e r and sphagnum a r e o t h e r f a c t o r s t h a t c a u s e m o r t a l i t y o f you ng c e d a r seed lin g s. N a t u r a l d r a i n a g e w a y s had an i n h i b i t i n g e f f e c t upon s e e d l i n g r e p r o d u c t i o n from t h e s t r e a m ' s e d g e t o a d i s t a n c e o f 75 f e e t . of th is No e x p e r i m e n t a l e v i d e n c e i s phenomenon. o ff e r r e d in e x p la n a tio n Growth s t u d i e s i n d i c a t e t h a t a p p r o x i m a t e l y t e n y e a r s a r e r e q u i r e d f o r a s e e d l i n g t o r e a c h 2 f e e t i n h e i g h t and t w e n t y y e a r s t o r e a c h ^ f e e t i n h e i g h t on a B i f l e p e a t is site. B eproduction r o u g h l y c o r r e l a t e d w i t h t h e b a s a l a r e a o f c e d a r and t h e h i g h ­ e s t numbers o f s e e d l i n g s a r e f o u n d i n a r e a s w i t h t h e g r e a t e s t b a s a l a r e a o f c e d a r and on t h e f a v o r e d o r g a n i c s o i l types, B ifle p e a t and C a r b o n d a l e muck. T e m p e r a t u r e and m o i s t u r e o b s e r v a t i o n s were c a r r i e d o u t a s a phase o f th e stu d y . sta g e s o f decay i s The t e m p e r a t u r e s w i t h i n l o g s i n v a r i o u s s i g n i f i c a n t l y h i g h e r and showed a g r e a t e r v a r i a t i o n t h a n t h e t e m p e r a t u r e o f t h e swamp f l o o r . a i r t e m p e r a t u r e s w i t h i n t h e swamp a r e The a v e r a g e c o n s id e r a b ly low er than t h o s e o f a c h e c k s t a t i o n l o c a t e d on an a d j a c e n t u p l a n d r i d g e . M oisture p e r c e n t a g e s , o b t a i n e d t h r o u g h o u t t h e g r o w i n g s e a s o n from d e c a y i n g l o g s and t h e swamp f l o o r * c o n t e n t s t h a n hardwood l i t t e r . o f the i n d i c a t e d h i g h e r m o iB tur e The w a t e r l e v e l o f t h e p o r t i o n swamp t h a t i s a f f e c t e d by d r a i n a g e from t h e a d j a c e n t upland s l o p e s shows g r e a t e r f l u c t u a t i o n s t h a n t h e p o r t i o n s t h a t a r e l o c a t e d d e e p e r i n t h e swamp. A l t h o u g h s t u d i e s on t h e f a c t o r s a f f e c t i n g v e g e t a t i v e r e p r o ­ d u c t i o n were c a r r i e d on i n p a r a l l e l w i t h s t u d i e s on i n i t i a l se ed lin g reproduction, t h e numbers o f ste m s o f v e g e t a t i v e o rig in were l e s s and t h e r e f o r e n o t o f a m a g n i t u d e t o a l l o w s t a t i s t i c a l i n t e r p r e t a t i o n i n many c a s e s . B o t h f r e q u e n c y and abu nd ance o f i n i t i a l v e g e t a t iv e reproduction in d ic a te that n e ith e r l i g h t , pH, n o r t y p e o f g e r m i n a t i n g medium a r e c r i t i c a l f a c t o r s i n i t s estab lish m en t. v egetative The p r e s e n c e o f sphagnum moss d e f i n i t e l y a i d s reprod uction . The ag e o f t h e s t a n d i s felt t o be an M im portant f a c t o r i n d e t e r m i n i n g t h e amount o f v e g e t a t i v e rep rod u ction that w ill deer b row sing, resu lt. root r o t, R o d e n t and s n o w s h o e h a r e b r o w s i n g an d f a i l u r e of a d v en titio u s roots d e v e l o p w e r e n o t e d ag c a u s e s o f m o r t a l i t y regen eration . Growth, in r e la tio n d eterm ine i n view o f i t s parent p lan t However, it d eer browse that in to age, th is is i n many o f t h e type o f d ifficu lt d ep en d en ce upon th e r o o t to to system o f the forms o f v e g e t a t i v e p r o p a g a t io n . wag f o u n d t o b e more r a p i d i n p r o d u c i n g a v a i l a b l e than the of branches, s e e d l i n g m ethod. an d t h e la y erin g , form ation o f v e r t i c a l p rin cip a lly e t e m s fro m w i n d t h r o w n t r e e s w e r e t h e m o s t common means o f v e g e t a t i v e p ro p a g a tio n alth ou gh o th er s t a n d on R i f l e feet peat, i n h e i g h t was 0 f percent of vegetative t y p e s were found. 20 p e r c e n t In a r e p r e s e n t a t i v e o f t h e r e p r o d u c t i o n from 2 t o 8 s e e d l i n g o r i g i n and t h e r e m a i n i n g 80 o rig in . F o r t y - f i v e p e r c e n t o f advanced r e p r o d u c tio n m o r ta lity t h a t wag t a l l i e d wag due t o d e e r b r o w s i n g a n d 2 0 p e r c e n t c a u s e d by s n o w s h o e h a r e b r o w s i n g . m ortality, the c a u s e o f 17 p e r c e n t wa s u n d e t e r m i n e d . c a u s e s t h a t were n o t e d i n c l u d e r o o t o f the r o o t Of t h e r e m a i n i n g 35 p e r c e n t system to d e v e lo p , rot, co m p etition , broken le a d e r s , Other fa ilu re an d w i n d t h r o w . A l t h o u g h s n o w s h o e h a r e b r o w s i n g and a c o m b i n a t i o n o f s n o w s h o e an d d e e r b r o w s i n g d i d n o t r e d u c e t h e number o f c e d a r stem s p er a cre severely in a tw elve year p e r io d , the snowshoe b r o w s in g i n j u r e d a s much a s 2 5 p e r c e n t o f t h e d u c t i o n from 2 to 8 f e e t in h eigh t Among t h e a s s o c i a t e s o f c e d a r , in the area cedar repro­ stu d ied . sn ow shoe b r o w s i n g m arked ly r e d u c e d t h e number o f s t e m s p e r a c r e o f a i d e r and r e d maple JLO O b u t bad no s i g n i f i c a n t e f f e c t on b l a c k a s h or b a l s a m f i r . I n a p e r i o d o f e l e v e n y a r d i n g s e a s o n s , t h e number o f c e d a r s t e m s o v e r 2 f e e t i n h e i g h t showed an i n c r e a s e and a l d e r i n t h e same h e i g h t c l a s s e s Drowsing. a d e c r e a s e u n d e r know amounts o f d e e r T h i s t r e n d d i d n o t e x p r e s s i t s e l f a s r a p i d l y under he avy and medium d e e r b r o w s i n g a s i t d i d w i t h l i t t l e or no b ro w sin g , however. S t u d i e s on t h e e f f e c t o f s i l v i c u l t u r a l p r a c t i c e s upon cedar r ep ro d u c tio n i n d i c a t e t h a t v / it h l o p p i n g and s c a t t e r i n g w ith p ilin g and b u r n i n g . seed lin g reproduction i s o f t h e s l u s h o f br o w se s p e c i e s t h a n L e s s a l d e r i n v a s i o n was a l s o f o u n d w i t h l o p p i n g and s c a t t e r i n g t h a n v / i t h p i l i n g d u c t i o n 7/as f o u n d i n slash p i l e s . and b u r n i n g . In itia l No r e p r o ­ s e e d lin g reproduction i n c r e a s e d w i t h an i n c r e a s i n g r e s i d u a l s t a n d . leavin g a resid u a l greater A selectiv e cut s t a n d o f 500 ste m s p e r a c r e or 750 s t e m s p e r a c r e was s a t i s f a c t o r y . Tne a g e o f t h e s t a n d a t t h e t i m e o f c u t t i n g d e t e r m i n e s t h e amount o f v e g e t a t i v e r e p r o d u c t i o n , younger sta n d s fa v o r in g i t . the Heavy c u t s a p p r o a c h i n g and i n c l u d i n g a c l e a r c u t f a v o r b o t h a l d e r i n v a s i o n and b a ls a m f i r r e p r o d u c t i o n . Management o f c e d a r on a c o m m e r c i a l b a s i s s h o u l d r e l y on a system t h a t fa v o rs s e e d lin g r e p r o d u c tio n in view o f the i m p o r t a n c e o f form . Management f o r mass f o l i a g e p r o d u c t i o n s h o u l d f a v o r v e g e t a t i v e r e p r o d u c t i o n a s i t r e a c h e s an a v a i l a b l e h e i g h t f o r browsing s o o n e r . R e p r o d u c t i o n methods a r e s u g g e s t e d i n s o f o r a s t h e d a t a and o b s e r v a t i o n s o b t a i n e d i n t h i s It is s t u d y would a l l o w . p r o p o s e d t h a t t h e t e r m s " l a y e r i n g method" and " l a y e r i n g - v / i t h - s t a n d a r d s method" be u sed i n f o r e s t t e r m i n o l o g y t o d e s i g n a t e t h e t y p e s o f r e p r o d u c t i o n methods t h a t d e s c r i b e uedar r e p r o d u c tio n . A t t e m p t s t o i n c r e a s e d e e r f o o d and c o v e r and t o c h a n g e cut-over s p r u c e - b a l s a m t y p e t o c e d a r t y p e by c e d a r p l a n t a t i o n s nave been u n s u c c e s s f u l . LITERATURE CITED A l d o u s , C. M. and A l d o u s , S . E. 1944 The sn ow sho e h a r e — a s e r i o u s enemy o f f o r e s t p la n ta tio n s. Jour. F orestry 42:88-94. A .ld o u s, S. E. 1941 D e er management s u g g e s t i o n s f o r n o r t h e r n w h i t e cedar t y p e s . J o u r . W i l d . Management 5 : 9 0 - 9 4 . 194 9 E x p e r i m e n t a l p l a n t i n g o f f o o d and c o v e r f o r d e e r . U. S . D. I . , F i s h and W i l d l i f e S e r v i c e W ild life L ea flet 320:x-9. A l d o u s , S , E. and C. F . S m i t h F a l l and w i n t e r f o o d h a b i t s o f d e e r i n n o r t h e a s t e r n 19 59 M in nesota. U. S . D. Af , B i o l . S u r v e y , W i l d l i f e R e s . and Mgt. L e a f l e t B. S . 1 3 7 . B aldw in, i. I . Forest tr e e seed. 1942 C h r o n i c a B o t a n i c a C o . , Waltham, M a ss. B a r t l e t t , I . H. W h i t e t a i l s , p r e s e n t i n g M ichigan*s deer problem. 19S8 M ic h i g a n D e p t . C o n s . , Game D i v . , B u i . C. and ’il. K. Goddard Bramble, E f f e c t o f a n i m a l c o a c t i o n and s e e d b e d c o n d i t i o n 1942 on r e g e n e r a t i o n o f p i t c h p i n e i n ' t h e b a r r e n s o f c e n tr a l P en nsylvan ia. Ecology 2 3 :3 3 0 -3 3 5 . o r u c e , D. and F . X. Schumacher F o rest m ensuration. 194.2 M cG raw -H ill Book C o . , I n c . , New Y ork, B u r n s , G. P. S t u d i e s i n t o l e r a n c e o f New En g la n d f o r e s t t r e e s . 1916 I I I . D iscontinuous l i g h t in f o r e s t . V t . A gr . Exp. S t a . B u i . 193: B u r t , W. i . The mammals o f M i c h i g a n . 1946 The U n i v e r s i t y o f M i c h i g a n P r e s s , A.nn A r b o r . C a l h o u n , I . D. Announcement o f pr ogram. 1948 N or th A m erica n Census o f S m a l l Mammals R e l e a s e No. 1 , Roden t E c o l o g y P r o j e c t , John Hopk ins U n i v . C n e n e y , E . G. American S i l v i c s and S i l v i c u l t u r e . 1 94 2 Tne U n i v e r s i t y o f M i n n e s o t a P r e s s , M i n n e a p o l i s . Co o n e r , W. S. R e p r o d u c t i o n by l a y e r i n g among c o n i f e r s . 1 31 1 Bot. Gaz. 52: 56 9 - 5 7 9 . 1915 I he c l i m a x f o r e s t o f _ s l e x i o y a l e , Lake S u p e r i o r , and i t s d e v e l o p m e n t . b o t . Gaz. 5 5 :1 -44,115-140,189-255. JLOC7 15. C u rtis, 1 94 4 J . D. N o r t h e r n w h i t e c e d a r on u p l a n d s o i l s Jour. Forestry 42:756-759. i n M ain e. 16 . 1946 17. 18. 19. 20. 21. 22. 25. 24. 25 . 26 . 27. 28. 29. P r e l i m i n a r y o b s e r v a t i o n s on n o r t h e r n w h i t e c e d a r i n M a in e . E cology 27:23-36. D a v e n p o r t , L. A. Deer f e e d i n g e x p e r i m e n t s . M i c h i g a n D e p t . C o n s . , Game D i v . , Unpub. R e p t . 318. D a v e n p o r t , L. A . , Yf • S n a p t o n , and W. C. Gower 1 94 4 A study o f the c a r r y in g c a p a c it y o f deer yards a s d e t e r m i n e d by b r o w s e p l o t s . N i n t h N o r t h Amer. W i l a l . C o n f . T r a n s . :144-149. Em erson , F . V/. 1921 S u b t e r r a n e a n o r g a n s o f bog p l a n t s . B o t . G az. 72:359-374. F e r n a l d , M. L . 191 9 L i t h o l o g i c a l f a c t o r s l i m i t i n g th e ranges o f P inus bank s i ana and Thu.ja o c c i d e n t a l i s . Rh odor a . 2 1 : 4 1 - 6 7 . F e u s t e l , I . C. and H. G. B y e r s 1936 The c o m p a r a t i v e m o i s t u r e - a b s o r b i n g and m o i s t u r e r e t a i n i n g c a p a c i t i e s o f p e a t and s o i l m i x t u r e s . U. S . D. A . , T e c h . B u i . 532. F i s h e r , G. M. 1935 C o m p a r a t i v e g e r m i n a t i o n o f t r e e s p e c i e s on v a r i o u s k in d s o f s u r f a c e - s o i l m a te r ia l in th e w estern w nite pine ty p e. E cology 16:606-611. Game D i v i s i o n , M i c h i g a n D e p a r t m e n t o f C o n s e r v a t i o n 1 94 9 1 9 4 8 deez' h u n t i n g i n f o r m a t i o n w i t h 1949 r e g u l a t i o n s G a t e s , F . C. 1926 P l a n t s u c c e s s i o n s a b o u t D o u g l a s L a k e , Cheboygan County, M ichigan. B o t . G a s. 82:170-182. G e v o r k i a n t z , S . R. and V/. A. Duerr 1959 Volume and y i e l d o f n o r t h e r n w h i t e c e d a r i n t h e Lake S t a t e s . Lake Sx-ates F o r e s t E xp . S t a . , m u l t i l i t h e d , 55 p p . Grano, C. X. 19 49 I s l i t t e r a b a rrier to the i n i t i a l e sta b lish m en t o f s h o r t l e a f and l o b l o l l y p i n e e s t a b l i s h m e n t ? Jour. Forestry 47:b44-548. H arlow, W. M. 1 92 8 R e p r o d u c t i o n o f A d i r o n d a c k w h i t e c e d a r by natural c u ttin g s. Jour. F orestry 26:244. Harmer, P . M. 1941 The muck s o i l s o f M ic h i g a n M ic h . A g r . Exp. S t a . S p e c . B u i . 314. H aw ley, R. 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I n f l u e n c e o f r o d e n t s on n a t u r a l r e g e n e r a t i o n o f D o u g la s -fir in th e Southw est. Jour. F orestry 43:585-589. Lakes S t a t e s F o r e s t Experim ent S t a t i o n 1936 F o r e s t a r e a s and t i m b e r v o l u m e s i n M i c h i g a n . Econom ic N o t e No. 5 , m u l t i l i t h e d , 4 0 pp. 36 . 1936 57. 38. 39. 40. 41. /. *: k * 45. 44. Woody f o o d p r e f e r e n c e s o f t h e sn owshoe r a b b i t i n t h e Lake S t a t e s . T e c h n i c a l N o t e No. 109. L e B a r r o n , R. K. 1948 S i l v i c u l t u r a l management o f b l a c k s p r u c e i n M in nesota. U. S . D. A . , C i r c . 791. L e B a r r o n , R. K. and J . R. N e e t z e l 1942 D r a i n a g e o f f o r e s t e d swamps. E cology 23:457-465. L e o n a r d , A. A . , C h i e f E n g i n e e r P e r s o n a l C o r r e s p o n d e n c e , DeJur Amsco C o r p o r a t i o n , Long I s l a n d C i t y 1 , New York. L u t z , H. J . and R. F. C n a n d l e r , J r . 1946 Forest s o i l s . Jobn W i l e y & S o n s , I n c . , New York. M&jci , T. E. 1951 Some f a c t o r s a f f e c t i n g g r ow th and r e p r o d u c t i o n o f Tnu .j a o c c i d e n t a l i s i n swamps i n M i n n e s o t a . Unpub. t h e s i s , U n i v e r s i t y o f M i n n e s o t a , S t . P a u l . Montgomery, R. D. Some f a c t o r s a f f e c t i n g t h e r e p r o d u c t i o n o f .n o r t h e r n w h i t e c e d a r ( Thu.ja o c c i d e n t a l i s L . ) i n Maine. Unpub. m a n u s c r i p t . Moore, B. 192:' Humus and r o o t s y s t e m s i n c e r t a i n n o r t h e a s t e r n f o r e s t s i n r e l a t i o n t o r e p r o d u c t i o n and c o m p e t i t i o n . Jour. F orestry 20:253-254. M c C u llo u g h , H. A. 1 94 8 P l a n t s u c c e s s i o n or. f a l l e n l o g s i n a v i r g i n sp ru ce-fir f o r e s t . E cology 29:508-513. 45. 46. 47. 48. 49. 50. 51. Pomeroy, 1949 K. B, Eke g e r m i n a t i o n a n d i n i t i a l e s t a b l i s h m e n t o f l o b l o l l y p in e under v ariou s su r fa c e s o i l c o n d itio n s . Jour. F orestry 47*541-543. P o t z g e r , J . E. 1937 V e g e ta tiv e rep ro d u ctio n in c o n if e r s . Am er. M i d i . B a t . 18:1001-1004. E i g g , 6 . B . a n d E. S . H a r r a r 1931 Ehe r o o t s y s t e m s o f t r e e s g r o w i n g i n s phagn um . Amer. J o u r . B o t . 18:391-397. S i e g l e r , H. E. 1937 W i n t e r r o d e n t damage t o game c o v e r . J o u r . Mamm. 18:57-61. S o c i e t y o f Am erican F o r e s t e r s 1940 F o r e s t co v er ty p e s o f the e a s t e r n U nited S t a t e s . 3rd e d i t i o n , r e v i s e d . S o c i e t y o f A m e r i c a n F o r e s t e r s , W a s h i n g t o n , D. 0 . S t e w a r t , G. 1925 F o r e s t t y p e s o f t h e n o r t h e r n swamps. Jour. Forestry 23:160-172. S w i f t , E. 1946 A h is t o r y o f W iscon sin deer. W i s c o n s i n Cons. D e p t . P u b . 323* 52. 1948 53* 54. 55. 56. 57. 58. 59. 60. — W i s c o n s i n ’ s d e e r damage t o f o r e s t r e p r o d u c t i o n survey — f in a l rep o rt. W i s c o n s i n Cons. B e p t . P u b . 3^7* T r i p p e n s e e , E. E. 1948 W i l d l i f e management. M c G r a w - H i l l Book C o . , I n c . , New Y o r k . U. S . D e p a r t m e n t o f A g r i c u l t u r e 1941 C l i m a t e a n d man. Yearbook. County S o i l S u r v e y s , M ic h ig a n : A l g e r , B a r r y , B r a n c h , C a l h o u n , C l i n t o n , E a t o n , H i l l s d a l e , Ingh am , L i v i n g s t o n , J a c k s o n , Macomb, M u s k e g o n , O c e a n a , S c h o o l c r a f t , S t . C l a i r , T u s c o l a , W as htenaw . W a t s o n , E. 1936 Northern w hite ced ar. U. S. F o r e s t S e r v i c e , E e g i o n 9» m i m e o . , 4 4 p p . W e a v e r , J . E . a n d F. E. C l e m e n t s 1938 P l a n t e c o l o g y , 2nd e d i t i o n . M c G r a w - H i l l Book C o . , I n c . , New Y o rk . W e s t v e l d , E. H. 1939 A p p lied s i l v i c u l t u r e in the U n ited S t a t e s . J o h n W i l e y & S o n s , I n c . , New Y o r k . W h e r r y , E. T. 1922 S o i l a c i d i t y p r e f e r e n c e s o f some e a s t e r n con ifers. Jour. F orestry 20:488-496. W iesner, J. 1907 Der l i c h t g e n u s s d e r p f l a n z e n s . L eip sig . 61. 62. W i l d e , S . A. 19^6 3Torest s o i l s and f o r e s t g r o w t h . O h r o n i c a B o t a n i c a C o . , Waltham, M ass. Zon, B. and J . L. A v e r e l l * 1929 D r a i n a g e o f swamps an d f o r e s t g r o w t h . U. o f W i s e . A g r . E x p . S t a . R e s . B u i . 8 9 . 4FPSBDIXES Table 35* — .Abundance and frequency o f se e d lin g reproduction under 2 f e e t in h e ig h t in v a rio u s l i g h t c o n d itio n s F oot-candle power 5-7 7-10 10-14 14-20 20-28 Abundance Number of stems 3 .2 10.0 6 .5 17-0 6 .6 Frequency Number of m ilacre p l o t s in c la s s P ercent 38. 86. 90. 82. 77- 8 14 19 28 31 28-40 40-53 58-81 81-114 114-162 2.9 9 .0 6 .8 9 .6 21.0 67. 76. 53. 76. 69. 15 25 19 17 13 162-232 232-331 331 P lu s 23.8 5 .5 1 .2 7550. 33. 8 4 9 Table 3 6 . — Abundance and frequency of seedlin g re-production in swamp stands at various pH l e v e l s 1 / • • pH Abundance Frequency : 5 Humber of m ilacre p l o t s in c la s s • • 4. 0-4.4 4.5-4. 9 5.0-5.4 5.5-5.9 6.0-6.4 6.5-6.9 7.0-7.5 Humber o f stems Percent 1*5 0.8 1.2 0.9 7.3 17.4 39.1 54. 33. 4o. 3366. 81. 92. 1 / T a l l i e s are based on seedlings from 0 to 2 f e e t in height. 13 6 10 15 61 52 13 A n a l y s i s o f v a r ia n c e . Germination o f cedar seed i n the l aborato r y under d i f f e r e n t l i g h t c o n d i t i o n s . O r i g in a l data p res en te d i n t a b l e 6. Source________ D. P. Treatment 1 Time S. S. Variance______ P________ 5^_____ l £ 60 60 H 1377 Treatment x Time b Replicates 1.50 1+.21 7 .6 8 3U4 8 . 60 ** 2.7 3 U .ll 1806 1+52 11 . 30 ** 2 .73 U .ll 3 83 28 8 .6 2 2 6 .5 0 Error 27 1076 *J0 Total 39 U1402 A n a l y s i s o f variance pH c o n d i t i o n s . Source 1.^3 Germination o f cedar seed under various O rig in a l data presen te d i n t a b l e 7. D. P. S. S. Variance P & 1* PH 5 3229 6 U5 .8 35.5** 2 .3 0 3 .2 0 Time 3 I 625 U 5^18.0 297.6** 2 .7 0 3 .9 8 Replicates b 12 3 .0 6 . 1* 5 .6 6 13.57 Time x pH 15 50U5 3 3 6 .3 1 8 . 5 ** 1 .7 8 2 .2 5 Error 92 1671 1 8 .2 Total 119 26211 L. S. D. a -/ Error variance x N x 2 a J 18.2 x 6 x 2 = / 218.1+ = 29 . U x x 1.985 1.985 x t 0j x y 7 Treatment T otal Comparisons pH 3 .0 92 14.0 3*17 L ess than 7 .0 , 8.0 5 .0 362 L ess than 8 .0 6 .0 293 L ess than 8 .0 7 .0 378 8 .0 Uoo L ess than U.Q, 5 . 0 , 6 .0 , 7 .0 , 8.0 An alysis o f variances. Germination of cedar seed i n the laboratory on various types of germinating media. Original data presented i n t a b l e 8. Source D. P. S. S. Variance P 5# 1* Treatment 3 75*4 251 1.37 3 .8 6 6.99 Replicates 3 238 79 2 .3 2 8 .8 1 2 7 . 3*1 Error 9 16 H6 I 83 Total 15 2638 Correlation between the number of cedar Btems per acre at present (19*19) and the percentage o f browse available in Cusino Browse P l o t s . (X) Plot Cedar Stems (Y) Browse Percentage (X)^ (Y)^ XY 5 103 3 .0 1 0 ,6 0 9 9 .0 0 3 0 9 .0 8 155 6 .9 211,025 147.61 1 , 0 6 9 .5 1 I 87 23.3 3^,969 5 *12.89 *4,357.1 7 299 U5 .2 89,1401 2 , 0143.014 13,51*4.8 H6.68 . / 2 0 ,6 2 0 x 1 .105 = .978 Appendix B: Common Baines o f Trees and Shrubs and th e ir S c ie n t if ic E quivalents. Alder Amalanchier Ash, B lack Aspen (Poplar) B a lm -o f-G ilia d B irch , W hite (Paper B irch ) B irch , Y ellow Cedar, n orth ern White (A rh orvitae) Cedar, Southern White Cherry, B lack Cranberry, High-bush Dogwood, R ed -o sier Elm, American P ir , A lpine P ir , Balsam P ir , D ouglas Hemlock Maple, Red P in e, E astern White Spruce, BlaCk Spruce, Engelmann Spruce, Red Spruce, White Tamarack Willow Alnus Inc ana W illd . Amalanchier canad en sis ( 1 . ) Med. P raxinus n ig ra .Man sh Ponulus trem u loid es Michx. Ponulus b a lsa m ifer a 1 . B e tu la n a n y r ife r a Marsh. B e tu la lu te ^ Michx. Thuja Occidenta l i s L. Chamaecynaris th v o id e s L. Prunus s e r o tln a Ehrh. ^ yiburnum onulus americanum ( M ill.) A it. Oornus s t o lo n if e r a Michx. UlmuB americana L. A bies lasiocar-p a (Hook.) B u tt. A bies balsam ea (L .) M ill. Pseudotsuga t a x i f o l i a (La Marck) B r it. Tsuga canadensis ( L .) Carr Acer rubrum L. P in u s strobus L. P ic e a mariana ( M ill.) B r it . P ic e a eiigelm annii Engelm. P ic e a rubra Link. P ic e a glau ca (Moench) V oss. L arlx la r ic in a (DuRoi) Koch. S a lix s p p . . . . Append!s 0: Common ITames o f Birds end Mammals and th eir S c ie n t if ic E quivalents. Birds Barred Owl Oreat Horned Owl Mammals Bobcat Chipmunk, Le ast Coyote Deer, White-tailed Hare, varying (snowshoe) Lynx Shrew, Masked Squirrel, Red Vole, Meadow Vole, Red-backed Weasel, Short t a i l e d Wolf, Timber Str ix v a r ia v ar ia Barton Bubo virginianus v i r g i n ! anus (Gmel.) Lynx rufus rufus Schreber Eutamius minimus .lackson! Howell Oanis latrans l a tr a n s Say Odocoileus virgin!anus b o r e a lis Miller. Leuus Americanus nhaeonotus Allen Lynx canadensis canadensis Kerr. Sorex cinereus cinereus Kerr Tamiasciurus hudsonicus loquax Bangs. Microtus nennsylvanicus -oennsylvanicus Ord. Olethrionomys ganneri genneri Vigors Mustela cicognanii cicognanii Boneparte Canis luous lycaon Schreber . A p p e n d i x D. Method o f S e e r Browse A n a l y s i s In trod u ction The method d e s c r i b e d h e r e i n i s a rather r a d ic a l m o d i f i c a t i o n o f t h e A l d o u s method o f d e e r y a r d a n a l y s i s 1 / t h a t was u s e d i n d e t e r m i n i n g "browse c o n d i t i o n s d e s c r i b e d i n th is d isserta tio n . been in u se fo r The f u n d a m e n t a l c o n c e p t s i n v o l v e d h a v e so m etim e by the Game D i v i s i o n , M ichigan D e p a r tm en t o f C o n s e r v a t i o n . F i e l d Procedure F i e l d p r o c e d u r e i s b a s e d upon t h e Game D i v i s i o n Browse S t u d y Form No. 1 , a copy o f w hic h i s d escrip tio n . i n c lu d e d w it h the C o l l e c t i o n o f d a t a was made i n p e r m a n en t p l o t s one c h a i n i n l e n g t h and 10 l i n k s (6.6*) in w idth. Stem c o u n t s were made o f t h e d om in a nt b r o w s e s p e c i e s — northern w h ite A ll stems c e d a r — i n th e y a r d u n d e r s t u d y . o f t h e d o m in a nt b r o w s e s p e c i e s o v e r two f e e t in h e i g h t w e re t a l l i e d a c c o r d i n g t o t h e breakdown o f t h e f i e l d sheet. P reviou s ex p erien ce 0 to 2 f o o t in d ic a t e d th a t the i n c l u s i o n o f the c l a s s w o u ld i n t r o d u c e a marked e r r o r i n t h a t t h e stems w i t h i n t h i s h e i g h t c l a s s a r e i n a v a i i a h l e a s w in te r food f o r t h e w h i t e t a i i due t o snow c o n d i t i o n s . O f f i c e Procedure Summarize th e f i e l d d a t a . i n each browse 1/ category. O b t a i n t h e t o t a l number o f ste m s C a lc u la te the p e r ce n ta g e o f ste m s i n A l d o u s , S. E. 1 9 ^ . A Deer Br owse Su rve y Method. J o u r n a l o f Mammalogy 2 5 *1 3 0 - 1 3 6 e ^ c h "browse c l a s s . B r o w se C l a s s N o. o f S tem s No "browse P e r c e n ta g e I n C la s s 50 5 1 .0 ( 5 0 /9 8 ) 0 - 33$ 28 2 8 .6 (28/98) 34 - — 66$ 6 6.1 ( 6/ 98) 67 - 1 0 0 $ 14- 1 4 -.3 ( 1 4 -/9 8 ) TOTAL 98 W ith t h e p e r c e n t a g e i n a "browse i n d e x . o f a v a ila b le each c la s s , we a r e p r e p a r e d t o In d o i n g s o , we m u st e s t a b l i s h b row se f o r each o f th e a s in g le ta lly - c l a s s e s . o f e a c h c l a s s w as c h o s e n a s m o st r e p r e s e n t a t i v e The v a l u e s fo r th e v a r io u s c la s s e s C la s s No - 34- 67 - By m u l t i p l y i n g v a lu e The m ean o f th e c l a s s . w o u ld t h e n b e : B ro w se P a p t o r b r o w se 0 o b ta in , 0 .0 33$ 16.0 66$ 50.0 100$ 84- . 0 in each c la s s by t h e b r o w s e f a c t o r we a r r i v e a t a “ b r o w se i n d e x 11. U s in g t h e same e x a m p le s as p r e v io u s ly , th e p e r c e n ta g e we w o u ld o b t a i n No browse fo llo w in g r e s u lt s . B ro w se P a c to r P e r c e n ta g e I n C la s s C la s s th e B r o w se I n d e x 51.0 X 0.0 m 0.0 0 - 33$ 28.6 X 16.0 - 4- . 6 34- - 66$ 6.1 X 50.0 = 3 .0 67 - 100$ 14 - . 3 X 84-. 0 3 12.0 19.6 The "b ro wse index" 1 9 . 6 ma y be used c o m p a r is o n "between y a r d s o r "between s u c c e s s i v e y e a r s i n yard. H ow ever, it w as fo u n d m ore s a t i s f a c t o r y " p e r c e n ta g e o f a v a ila b le th a n th e It same to u se th e "browse" a s a c o m p a r a tiv e v a lu e r a t h e r " b ro w se i n d e x " . s h o u l d b e rem em b ered t h a t on a maximum v a l u e o f 8 4 , u p p er b row se c l a s s in d e x " "by t h i s a v a ila b le a r r iv e th e b r o w se " f o r a v a ila b le " b row se in d e x " of b r o w se " f o r ( 8 4 .0 ) , b a sed By d i v i d i n g t h e we a r r i v e a t t h e " p e r c e n t a g e th e in d ic a t o r 23.3 is t h e mean p e r c e n t a g e o f b r o w se i n o n I*orm 1 ( 6 ? - 1 0 0 $ ) . v a lu e a t a f ig u r e th e s p e c ie s . ( 1 9 .6 /8 4 .0 ) t h e e x a m p le th e " b row se of Thus we w o u ld a s th e " p e r c e n ta g e of c ite d . D is c u s s io n The a n a l y s i s o f f i e l d i n f o r m a t i o n i n a c c o r d a n c e w it h t h e m e th o d d e s c r i b e d made p o s s i b l e of fie ld ta llie s ta k e n a t th e h ig h e s t d e g r e e o f u t i l i z a t i o n t h e t im e o f p lo t e s ta b lis h m e n t. A n u m e r ic a l e s t i m a t e o f b r o w se c o n d i t i o n s w as o b t a i n e d by t h i s m e th o d . th e A t h ir d a d v a n ta g e th a t t h i s fa c to r s m eth o d p r e s e n t e d w as t h a t o f n a t u r a l p r u n in g and l o s s sn o w sh o e h a r e b r o w s in g w ere i n t e g r a t e d It is a c k n o w le d g e d t h a t t h e r e a r e d is a d v a n t a g e s in h e r e n t in is made f o r in to th e v a lu e s o b ta in e d . sev era l t h e m eth od e m p lo y e d . th e v a r y in g p e r c e n ta g e o r d o m in a n t b r o w se s p e c i e s n o r i s d iffe r e n c e s o f b r o w se d ue t o No a llo w a n c e c o m p o s it io n o f t h e i n d i c a t o r a l l o w a n c e made f o r p o s s i b l e i n b ro w se p r o d u c t i v i t y o f th e s i t e s e r io u s on v a r i o u s fe lt th a t u n ifo r m ity an d f o r e s t th is so u r ce o f e r r o r to a la r g e d e g r e e . s ite s . I t was t y p e w o u ld c o n fo u n d A f u r t h e r d is a d v a n ta g e o f th is m eth o d l i e s in th e f a c t fa c to r s ape not a v a ila b le f i n a l v a lu e - p er acre th e l a t t e r c r u is in g t h a t w o u ld a l l o w p e r c e n ta g e o f a v a ila b le o f a v a ila b le th e u lt im a t e ly th a t — a t p resen t — "browse, a f i g u r e d e s ir e d d is a d v a n ta g e m e th o d s i n u s e in te g r a tio n , c o n v e r s io n o f th e b r o w se — d ir e c tly in to pounds r e la te d c a r r y in g c a p a c it y . se e m s t o "be i n h e r e n t i n a n to d a y . s a tis fa c to r y to H ow ever, t h e "browse MICHIGAN DEPARTMENT OF CONSERVATION Game D i v i s i o n BROWSE STUDY FORM Name o f Yard_ Location: T_ Study P lo t No;.________ Average Maximum Snow Depth N R Sec, P roject No- Sub_ Investigator__ '""I* SPECIES 0 -2 ft, 2* J 3> 2-5 f t , j 5~g f t . No Browse Height o f Browse Line_____ Date I 5. ! b. ; 8 ?-3 » DBH I 3^6" DBH j 6 -9 “DBS j j ____ o -J li3^ - 6 # _ -- — No Browse1 l 6 - 100 No Browse 2 - = - 2 I .k . „ 2L =J§J. 67 - 100 % No Browse lija jL -7- ~ J ; 0 0 .^ „ BCJ:aet ------------------ Appendix E: A s s o c ia te s o f Cedar from C o lle c tio n s and O bservations. S c i e n t i f i c ifame A bies balsamea (L .) M ill. Acer rubrum L. Acer -pennsyivanicom 1 . Acer saccharum Marsh. Alnug incana ( L .) Moench. Am elanchier canadensis ( L .) Medic. Andronogon fu rca tu s M. A r a l!a n u d ic a u lis L. B etu la a lb a L. B etu la lu te a Michx. f . Botrychium virginlanum (L.) Sw. Carex f la v a r e c t lr o s t r a Gaudin. Carex f o l l l c u l a t a L. Carex s t r i c t a Lam. Carex trisoerm a Dewey Chamaeda-phne c a ly c u la ta ( L .) Moench Chelone glabra L. Cornus a l t e r n i f o l i a L. f . Cornus can ad en sis L. Cornus s t o lo n if e r a Michx. Corylus r o s tr a ta A it . Cyorinedium a c a u le A it. Cyprioedium uubescens (W illd .) Knight. Eauisetum spp. Evxoatorium uerfoliaturn L. Euog.torium ■ouraureuia maculatum (L. ) D ari. E ragaria v esca americana P orter Fraxinus nigra Marsh Galium asnrellum Michx. Gault her i a -orocumbens L. G entiana nrocera Holm. Habenaria d ila t a t a (pursh) Gray. H eracl eum lanatum Michx. Inroatiens b i f l o r a Walt. I r i s v e r s ic o lo r L. L arix la r ic in a (DuRoi) Koch. Ledum groenlandicum Oeder. L ilium •ohiladel-phicum andinum (H u tt.) Ker. Linnaea b o r e a lis americana (Porbes) Rehder. L o b elia Kalmii L. L onicera canadensis Marsh. L onicera h ir su ta Eat. Lycouus americanus Muhl. Malanthemum canadenoe D esf. M ic r o s ty lis mononhyllos ( L .) L in d l. Monotrooa u n iflo r a L. P a rn a ssia c a r o lln ia n a Michx. P ice a canadensis (M ill. ) BSP, P ic e a mariona ( M ill.) BSP. Pinus Strobus L. Common Name Balsam f i r Ped maple S trip ed maple Sugar maple Speckled a ld er S e rv ice berry Beard grass Wild s a r s a p a r illa White b irch Y ellow b irch R attlesn ak e fern Leather l e a f Balmony A lte r n a te -le a fe d dogwood Bunch berry R ed -osier dogwood Beaked hazelnut S tem less la d y 's s lip p e r Larger y ello w la d y 's slip p H o r se ta il Boneset Joe-Pye weed Black ash Rough bedstraw Teabeiuy Gentian Fringed o rch is Cow parsnip S potted touch-m e-not Larger blue f la g Tamarack Labrador te a Wood l i l y Twin flow er American f l y honeysuckle Hairy honeysuckle Adder's mouth Indian p ip e Grass o f Parnassus White spruce Black spruce White p in e Polygonatum commutatura (R. & S . ) D ie tr . Pouulug b a lsa m ifer a L. Pouulus trem u loid es Michx. P o t e n t i li a f r u c tic o s a L. P r u n ella v u lg a r is L. Prunus s e r o tin a Ehrli. Ranunculus a c r io L. Rhamnus a l n i f o l i a L’Her. Ri~bes spp. Rosa spp Rubus t r i f l o r u s R ichards. S a lix spp. S arracen ia purpurea L. S ciro u s a tro v iren 3 Muhl. S c u t e lla r ia g a le r lc u la t a L. Senecio aureus L. Sisyrinchium a n g u stifo liu m M ill. S m ilacln a racemosa ( L . ) D esf. Solidago se r o tin a A it . Solidago u llg in o s a N u tt. Snhagnura spp. S t e ll a r i a l o n g i f o l i a Muhl. Thalictrum uolygamum Muhl. Tsuga can ad en sis ( L . ) Carr. Typha l a t i f o l i a L. Vaccinium spp. V alerian a u lig in o s a (T. & G .) Rydb. Viburnum c a s s in o id e s L. V io la spp. Great Solomon’ s s e a l Balsam p op lar American aspen Shrubby c in q u e fo il H e a l-a ll Wild black cherry T a ll buttercup Buckthorn Dwarf raspberry P itc h e r -p la n t Bulrush Skullcap Golden ragwort B lu e-eyed grass F a lse spikenard Golden rod Golden rod Chickweed T a ll meadow rue Hemlock Common c a t - t a i l Swamp v a le r ia n W ild r a is in