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Les cartes, planches, tableaux, etc., peuvent Atre filmte d des taux de reduction diffirents. Lorsque le document est trop grand pour Atre reproduit en un seul clich«, 11 est film« A partir de I'angle supArieur gauche, de gauche A droite. et de haut en bas, en prenant le nombre d'images ndcessai'e. Les diagrammas suivanta illustrent la m^thode. 1 2 3 1 2 3 4 5 6 . MiaoCOnr RiSOlUTION TKT CHART , NS! and ISO TEST CHART No 2) ^ APPLIED IM/IGE I nc JBC . '655 €03l Main Streei B^S Roc^ieater. Naw Yorl< l*609 uSA ^^S (^'6) *82 - 0300 - Phor-e ^^S (716) 2S8 - 5989 - Fa« Reprinted iron Th* Bthmital GattUf, 4t; 8i-io«, AnguX, 1906 THE NASCENT FOREST OF THE MIS- COU BEACH PLAIN CONTRIBUTIONS TO THE ECOLOGICAL PLANT GEOORAPHV OK THE PROVINCE OF NEW BRUNSWICK. NO. 4 (WITH FO'JKTBBN nOURES) W. F. GANON<; PSUrrU) AT m UMIVBRUTV Or CHICAGO PKKSS VOLUME XIJI NUMIIKK 2 ! i Botanical Gazette AUGUST, igo6 THE NASCENT FOREST OF THE MISCOU BEACH PLAIN. CONTRIBUTIONS TO TIIK KCOI.OOICAL PLANT (JEOORAI'HY OF THK I'ROVINCK OF NKVV HRUNSWICK, NO. 4.' \V. F. Canon o. (with fourtkkn figures) The extreme northeastern angli of the Province of New Hruns- wick, as the accompanying map will show, is formed by the island of Miscou. The northwestern margin of this island is an extensive sandy beach plain, growing rapidly by action of the sea, locally called Grande Plaine. On this plain there is developing a forest which exhibits every stage of formation from the salt plants of the open sea beach to the heterogeneous vegetation of the mixed woods. The conditions are unusual and the phenomena of proportional interest. In August 1905 I wa.s able to give the place some two weeks of oteervational study, with results which follow. In all such studies as this the correct identification of the plants is of first importance, and idcntificalun is becoming a matter of such difficulty that only a professional systematist is competent authority. Accordingly I have sent all of my a llections, including a .specimen of every plant I found at Grande Plaine, to Professor M. L. Fernald, of the Gray Herbarium of Harvard University, who has been so kind as to determine their identity, and, as well, to give me the names they should bear in accordance with the recom- mendations of the Vienna Congross. I wish here to express my indebtedness to him and my best thanks for this invaluable aid. Such is the origin of the nomenclature of this paper. ■ No. 3 is in the Box. Gazette 36:161-186, 280-302, 349-367, 4*9-455. 1903. 81 I BOTANICAL OAZHrTE (ai'uuit An li» prcviiius litcraturt- of thi(» partiiular bubjtit, thcrt' i.s none. In 1886 Dr. (J. U. Hay made a colliiiion i>f Miscou plants for the Ucologiial Survey of Canada, but no aicounl of them was ever published, and no other liotanisl has heretofore been tin the island. In many resfx its, however, as the reader will observe, the vegetation of this bea. h plain resembles closely the vegetation no. I.— Outline map of part of the Gulf of St. Lawrence, to show the geographical position of Miscou Island. of the sand beaches and dunes of Lake Michigan as described in CowLKs's well-known memoir,' and many of the conclusion^ of that work are also applicable here. Grande Plaine extends along the v.est side of Miscou. Begin- ning on the south at Eel Brook (see the accompanying map, fig. 2), >B0T. GAMTTE 37:95-117, 167-JOJ, 281-308, 361-391. 1899. looft) C.tNOXa-XASrF.XT FOKEST OF UlSCOf BF.ACll H where it is but a f-w yards wide, it rapidly > r..adtns northward until it reaches Mune half a mile ;i ross, and ihen narr -wi» URain tuward-H its mtrthern end, which is also the Btty CAnitur northernmost p.»int ol ihi island. Though nearly level as a whole, .! is by no means t^at, for it is com. IK)8cd of a series of ap- proximately concentric dune beaches, which, two or three in numlK-rat Kel BnK)l(, in crease to over forty opposite Lac Krye. In height these dune be.rhcs vary from tv;o to five or six or even seven feet (o.6-a"') and in breadth from eight or ten up to forty or fifty paces. At iis widest part, wnich '•omprises s»)me thirty or more of the beaches, new ones are plainly being ra-^idly added, while at its nortnern end the entire plain is being washed away by the sea, which is cutting sha.7>ly across the ends of the old Ijeachc Aiiout two -fifths of the plain, in- cluding the older parts n;xt the upland, are lorested; about two-fifths, incluriinjr, all the outer and mv.er parts, are open, cloi..ed i nly by the waving beach grass; the intermediate zone, a sr .11 one-fifth of the area of the plain, is a »4 BOTASICAL liAZETTK (AI'OVIT trutiMlion iimc whtn- the- f-.rusl is pushinj? iu advamc into the upcn Kruiiml. Thf mntU of f..rn»ali..n of this nmarkahk' |(lain, invulv itif,' ihf an.nnaly nf fXliriMvc land huildinu <,n a -inkinn ^'mM, I ha\i ili's( riU'd w.miwhat fully tlMwhiri'. ' Uriilly, ihf fa(W arr ihtM'. As tin- risull ot pcculiariliijt c-f iho ti.i)<.Kra()hy, wiml, anil liiUs .if this rif^iim, ihcro i-. f..rmcil .m iho shallow north- wi-sti'rn Hiilf ..f Mihc.ii a kimi of (jnat .cMy iti whirh all mo\at)lf nialtrials, sand ami uravil from thf wtiir of the rapitlly crumhIinK adjaifnt coasts, as will as driftwood, watirwcrds, and othtr tlolsam, ofini from a ^riat distami-, find to (olkii, and ihincf an- driven ii>horf by the prevailing wisltrly wiruU. I'ornurly the island ixlinded fartlur north than n-.w, tarrying wilh it Ix.th i-ddy and jilain; but ihf >r<" low northerly i nd beneath the sea, thus fori inj,' the eddy and the adompanyinjj plain buildin)? >,'radually southward. The northern end of (Irande I'laine tixlay is Ininf? rapidly washed away (com|)nre map), to be redeiK)sited farther south, and the plain as a whole is thus rolling by its outer margin southwaril along the eoasl. The subsidence of the lanil has pr.dured another effeet U|)(.n the plain, and one of lonsiderable eonse<|uen(e to its ve^e tat ion, namely, its inner and oliler |)art averages somewhat lower, that is, less above sea level, thiin the outer and newer part, thus lead- ing to a settling of water towards the older inner parts, and a rela- tively higher water table in them. That we have here a Ix-aeh plain, instead of a series )f lofty sand dunes, is the result of l\ • fact, appar- ently, that the dry sand of the beach is blown rshore o faster than the Inach grass can fix it. .At l>oth the northern and s(,uthcrn enis of the plain, however, there is some ajjproach to a building of true, though low, dunes. My brief study of the vegetat-on of Orande Plaine was entirely observational, not at all instrumental, nor do any meteorological or other exact physical data for this region exist. Orande Plaine lies at sea-level in latitude 48°, Ix-side a shallow sea, warm in summer but frozen over in winter. The summer climate is remarkably equable, of a temperature most comfortable for man, with no fogs and but little iloudy weather. The rainfall must lie not far from tBull. Nat. Hist. S.v<;-,v.i.vt>:.vy tnnKsr of- mi. Mxn HI t< // 45 iiKlus. Hiiivy wind;, irinn ilu' wi>i prtsiiil JMiil is of imn- or ;rt/, nanil tk-riwil in.m tb in siininu r. Ilu ifcroii!* Mimlstnms <>f ih »• W(iii II ihf ){ray larU. I ii'Kion, this naml havi nx. ' f iht usual rdalion.-. Id waur Mipph, minira) miinVnN, .1.. N,, .ili.r »|>i-iial faiti.rs with .1 UaririK u|n n \V v.xiiaiioii ,,,,|,t.ar (.. Ik' promint-nt. Wc iiirii now to ton-.i(liT the Mnitalion. AlihoiiKh it |.ri>tniH rwry Krudatjon fr. m Inin.l.lf lurh- . | ilu' (.|Hn Ua. h i<. ilu' (knsfM W(hkI,s, nivirlhcless ihi- lyi- Ih, ( , a.tiihl.nuil to riT.Kni/i. anignali', (.rtaiii .Klmilr M'^ilational rr>;i.ins. 'I'ln'M' ri-priHnt the mn.li.., „r .limaxis, as it w.ri', in thi> MKitati..p , urM' -Ihi' parts whiih ixhibit a iliMin. live ('lara.tiT in the ,,hvsio«- nomy of thi- whole Ihiy a v the iVllosinj;; (i) ihr new Ina.h, (2) ihi' ^-rass plain, (.;) the swaks, (4) ilu- samlv w,m«I>, (,) ihv cU>8f.re is niorliiied in- Imal c ndi lions of erosion and nsive to these conditions. It is extremely scanty, the plants growing widely isolated, while manv square yards do not show any vegetation at all. Thus c. mpetiti.in among the plants seems not to exist, and the struggle is wholly with the 86 BOTANICAL CAZETTh [august physical environment. The most characteristic plant by far is the small, radiate-decumbent, succulent, annual saltwort, Salsota y "/i\^^'' "nir //||„,'' '///n"y//ni>"%//iiii>'';„„. '/llllV7/(lll"'"///lillW"''«|,>'' mm 2 o e ^ .2 2 a I U. it; Kali, which occurs but rarely and for the most part in the lee of 1906] OANONG-XASCJlXr FOREST OF MISCOC liE.lCll 87 some shelter, such as a hollow or large piece cf (Iriftwood. Xcxt in abundance, though but scarce, is the little fleshy, nsette-like, annual sea rocket, Cakile edentula (C. amcricana). Third in abun- dance is the low haio-n sctte, perennial sea lungwort, Mtrlinshi marUima, here seemingly growing as an annual, also m( sllv in places of some shelter. Rarely, and then only in a sheltered ],< si'tion, occur tiny radiate-creeping plants of the beach pea, Luthyrm marl timus, growing apparently only as an annual, and sometimes show- mg a marked difference in the windward-creeping and leeward- FlG. 4.— lypical open, or new, beach, l.wking north; among the drift«i»(l occur scattered tufts of saltwort and Ijeach grass. creeping shoots on the same plant, the former being much shorter and smaller-leaved. Even rarer is the radiate-creeping, small- Ifaved, halo-scurfy annual, Atriplex patiila haslala. Here and there, however, especially in sheltered places, arise the tufted culms of the beach grass, Ammophila arenaria, the characteristic sand- binding perennial of the dune beaches next to be considered, here seemingly growing from seed. I was able to find no other plants on^ the new beach. Thus we see that this vegetation is distinctly adjusted to the physical conditions, for it is (,f great jjaucitv, of small and slow growth, annually renewed, closely ground-apprcssed, and strongly xerophytic. 88 BOTANICAL GAZETTE [Arousi THE GRASS PLAIN. Of atlV^' ';"' "! "'^"" ^''^ '^^'■"^ ''^'^ ^'^"d plain, composed !l fhT "T °^/PP'--™-'<^ly parallel dune beach s. forming smoothly swelhng ridges and hollows r.f elevations and breadth! already descnbed. Every dune beach, I believe, originates wU a core of dnftwood. As the tidal beach is built outwards byle par until rif' '"''"'''' '^™'""" '" ^*« "" ''^ "PPe-oM part, until finally some unusual combination of great winds with high t.dcs sweeps it up beyond reach of further disturbance Thn the dnvmg sand from the beach is caught among it; the beach grass gams a foothold in the shehered places, spreads, and cheks he further movement of this sand. Then more sa;d is dr'en shoreward, and it grows into a low dune which is fixed bv the beac^ ST-fT ^' r \""" '''' ""'* '^ ^^-'-d -ly -hen' a new 1 ne of dnftwood has been formed outside and begins to stop the sand or its own growth. The resultant dune beach offers severe cond' ttons for plant he, for its surface is swept, especiallv on the summ t and wmdward slope, by heavy winds; it is heated 'intensely bv the sun; >t .s readdy movable; and it forms a soil extremely p^r in mmeral nutnents.^ It lacks the salt of the newer beaches.'ho:e -er! or this ,s soon removed by the rain; and it possesses an ampl supply of moisture a foot or two beneath the surface for the sud^v brought by the rain drains but slowly away, owing to the Lw ^ad em of the water-table. These conditions, especially at their extreme on the summits and windward slopes of the Lch dunes. enlrTd by practically but a single plant-the herbaceous-perennial Tb terranean-creeping xero-culmed, deep-rooting beach^rass. iZo- fn t""' 7'"^' """""■ "''•^""^ '-y --I-titor ;hatever m open sea te^d tussocks, only partially coverinrthe ground ^ wel shown in fig. 5, and in closer view in figs. 6 and 7. iZLZ that this grass is of considerable economic value to the neighC mg farmers, who cut it and haul it for hay. and whose catt^'grte oTthe Jut^' btaT"" "•'"' ^'' ^^"^^^ ^" '-^'- -po- of the outer beaches, permitting them to be irregularly cut by the wind. It IS for this reason, I have no doubt, the newer outer beaches ^^^^^As indicated by Ke.k..Vs receni studies: Box. Gazkxtk 37:4.^436. .906] GAmxC-NASCnXT FOREST OF MISCOU BEACH gg are SO much more irregular in their various character, than the older inner beaches, which antedate the advent of man esped li?£hfnd'^"f '"? '" "" ^""P^"'"^' '^ ^'^-^•^ ^ shelter. oH ^e nit T- uT"""' P''""'"'"^ '''' «^"^-"^ "^ - ""-"be 1 InHr^ ' ' ^''''''''"' ^"'■'" ''"' ^" '-nsignificant part of Inothe m"':'''""' ''"' "''^'^ ^" -^^•»>' -P-ated from o e Catr'smT rr""' "' ''"^' P'^^^P^' '-^ '"^^ beach sedge aZ t t \ ^"""^^ '" '^^"^^^'^ '"««"^ks here and there among the beach grass, and it is indeed the only other plant Jhth theblrchS'''"' "'^ """■ """"'^ "°"'^ P"*^"^""-^ "° P"^"' -""e except seems at home in this situation. The remainder of the plants. aU sparsely or rarely represented, are obviously stragglers from the most diverse habitats, many of them quite^unex^ct d tTdem Lh anVt'eT"H ''^"^' ^^"^^ ^'^-^ stra^ in from the on i t L u IT '' ^'"^ '°'"^^'''^' """"^ flourishing than on the open beach. Then there are greatly dwarfed individuals of cjrtam ubrquatous forms able to endure a wide range of phyid hist S,-r "■" : °"'" ""^" '' '""^ P'^'"' ''^^ Canada thistle, Cmcus arvens^s; the sow thistle. Sonchus arvensis; the 90 BOTAXICAI. GAZETTE [august field sorrel, Rumex Acetosella, in dwarfcd-rosctte, very red forms; the evening primrose, Oenothera biennis; and the moonwort, Botry- chium ternatum intermedium. There are also some forms usually characteristic of rather a dry habitat, such as the pearly everlasting. Anaphalis margaritacea, and a western yarrow, Achillea lanulosa. In addition there are others, generally in more sheltered spots and also greatly dwarfed, which usually prefer a moister habitat, such as the two westc n roses, Rosa acicularis (R. Sayi, R. Engelmanni) and Rosa lucida; a western violet, Viola adunca; a stitchwort, Stel- laria long! pes laeta; a silver weed, Potentilla Anserina concolor; the stellate false Solomon's seal, Smilacina stellata; and one of the vetches, Vicia Cracca.' The great diversity of natural habit ()f these plants, their scanty and irregular occurrence, and their dwarfed size and rosette-forming tendency all unite to show that none of them are here at home. Obviously they are the ones which, of all the many kinds which must be brought to this plain by natural modes of dissemination, are sufficiently tolerant physiologically to be able to germinate „..der, and then to withstand, these extreme physical conditions, eking out here a starved and precarious exist- ence. The conditions for germination upon the sand must be extremely severe, and it is very likely that other kinds of plants could exist here as adults, could their seeds develop; and further It IS probable that the individuals which do exist on the plain are those whose seeds happened to fall in especially favorable spots, or became properly buried by the moving sand. Else why are they so few? The universal dwarfing is due in all likelihood not to the heat and dryness of the surface, nor to any salt content in the soil, and certainly not to a scarcity cf soil water, but to the paucity of mineral nutrients in the sand. This is in harmony with another feature they mostly show in common-very deep and, I think, much-branching roots. The fact that they come 5 The following Grande Plainc plants appear to he new to the flora of New Bruns- wick: .!./:,/,« lanulosa. Viola adunca, Rosa acicularis, Stdlaria longipts laeta, and PolenUlla Ansenna concolor. Certain others are new in name, the species having been recently more exactly defined and segregated: .ilnus mMs, Myrica carolin- rns,s,Vaccnuum VUis-Idaea minor. Others are new in name txrause made to con- form to the riiles of the Vienna Congress, but in these cases the names of Gray's Man- ual, 6th edition, have Ixen given in brackets. «9o6] CANONG-NASCRNT FOREST OF UtSCOV BEACH 91 from such a diversity of natural habitats, and yet hve in this peculiar situation upon an equal f(K)tinp, shows how far we are from under- standing the real bases of physiological adaptation, and further shows that in the study of the physiological life-histories of plants lies the most important and attractive field for the ecologist of the near future. So much for the exposed parts of the dune beaches. But in addition they offer, upon their inner or leeward slopes and in the hollows, situations more sheltererl, not so much from the sun, since Fic. 6.— Typical hollow between outer dune beaches; the tall grass is all beach grass, but the small plants among it are the common strawberr)-. their average course is nearly north and south, but from the westerly winds. The older inner dune beaches also are protected to some extent by the newer outer ones, as well as by their slightly lower average level. The diiference between leeward slope and hollow is not simply one of degree of shelter, however, but also of physical condi- tions, for the hollow is much nearer the source of water supply, the free table of which is not over a foot or two beneath the surface! In consequence of these differences we can recognize three distinct phases of vegetation: prst, a larger development on the leeward 9» BOTANICAL GAZETTE [aitoust li- slopcs of plants which are small and rare on the outer slopes; second, a distinctive vegetation of the hollows; and, third, a distinctive vegetation of the inner slo})es. As to the first phase, it is enough to note that several plants, in particular the pearly everlasting and the wild roses, small, scarce, and scattered on the outer slopes, become larger, frequent, and even patch-forming on the inner; and this is true alsD in less degree of other species. The beach grass persists in all situations. The second phase is the vegetation of the hollows. The very first plant to appear in this situation, and that too near the outer beaches, is always, strangely enougii, the common wild strawberry, Fragaria virginiana, apparently of normal size and form, seem- ingly quite at home, and spreading abundantly by runners, so that it forms considerable patches. The appearance of the nascent patches is wel' shown in fig. 6. As soon as the patches reach an appreciable density, such that they afford a cover to the ground, then turf-forminf? grasses, of which the first is the red fescue, Festuca rubra, appear and initiate the turf-formation which is so important a feature of the swales to be described below. The strawberry, of course, is one of the most tolerant, and hence ubiquitous, herbs of our flora, and its situation here is partially explained by the near- ness of the abundant water supply. Yet it is surprising to find it taking so important a part in a vegetation in so peculiar a position. The third phase of this vegetation is that characteristic of the sheltered slopes. First of such plants to appear, and the most common and characteristic, is the dwarf creeping juniper, Junip- erus Sabina procumbens, of which single plants originate just below the beach dune crests, and creep radiating, more to leeward than to windward, in a close dense mat covering many square feet. A young plant is show" in fig. y, in characteristic form and position. On the inner beaches ihese plants occur upon the oaier as well as the inner slopes, and the shelter of the mats thus formed affords in reality the principal starting-point for the development of other plants which lead gradually to the development of the forest, as will be noted under the transition vegetation. In a similar situation, but independently, arise patches of two other characteristic plants, a bright-green, leathcry-lea\ed, tufted shrub, the wax berry. iQOft] CA!VO\'a—NASCEXT FOREST OF MISCOU BEACH 93 Myrica carolitunsis, which comes to form dense discoid (sometimes almost fairy-ring like) masses (.n the crests and inner slopes; and the less frequent, low, dense-tufted, white hairy shrub, Hudsonia tomenlosa, in irregular close patches. All (.f these plants are pro- nounced xerophytes, which amply ex;>lains their ability to live in this situation, and even their preference for the somewhat drier upper slopes of the dune beaches. Their xen.philism, in common with that of many other evergreen sand plants, is, as I guess it, an 8W!i ''^N 't^^;^**^ t ,i«(»*r'ir*.. ■^^^ Fig. 7. — Typical upper inner slope of a dune beach; the grass is beach grass, but in the center is a typical plant of dwarf creeping juniper. adaptation to the physiological dryness which results when, as must often be the case in spring and fall, the ground water is of low temperature and hence slowly absorbed, while the leaves are exposed to high tianspiration conditicns from the bright sun, heat reflected from the .sand, and high winds.* The juniper, while perfectly at home here, apparently is so cnly thrcugh coincidence, for its original habitat is seemingly dry reeky hills. But the other * This principle, which from its discoverers we may call the KtHLUAN-GoEBEL principle, seems to me deserving of much more recognition than our students are inclined to give it. At least it calls for careful experimental investigation. 94 BOTANICAL GAZETTE [auovst i i|' two, the waxberry and the Huclsonia, are characteristic of just this situation, in and to vhich they have apparently been adaplively developed. Towards the inner dune Ix'aches another low shrub comes in on the slopes, though dwarfed and not abundant, the common blueberry, ,ccinium pennsylvanicum; it is evidently not here at home, bu Us somewhat xerophytic habit permits it to exist. As these various plants grow older and extend their patches, they run together more or less, sometimes two, sometimes three, and even all four. Later others are addi' to them, initiating the juniper mats and the wcwds carpet, later to be considered. The contrast lx;tween the vegetation of the outer and the shel- tered slopes of the dune beaches comes out with striking clearness a few hundred yards north of Eel BrcHik, where it happens the entire plaii. is very narrow, and sIcjIjcs in both directio. s from a central higher crest. Outside of this can be seen only the l)cach grass ami its accompanying forms as listed above, while inside the various xerophytic shrubs show to great perfection. THE SWALES. Between the open grass plain and the woods occurs a transition zone marked n')t only by an intermediate vegciation but also by distinctive physical features as well. First of all it is characterized by the presence of several great lurf-carpeted and tree-bordered swales, morphologically hollows l 'ween the dune beaches which here spread much farther apart than usual. They are well shown in figs. 8, 10, II. They are best developed in the widest part of the plain, hardly cccurring towards its southern or northern ends, and outside of them runs a line of higher dune beaches, which indeed can be traced through most or all the length of the plain (fig. z). The swales are narrow southward, but broaden northward, deepening as they go, until in some cases they dip beneath the water-table (thus exhibiting pools), after which they rapidly narrow and rise to disappear northward. Again, the trees of this zone, occurring always along the slopes of the dune beaches, do not exhibit a transi- tion of size and age to those of the sandy woods, but are always so much smaller and younger as to be sharply marked oflf from them, the case shown in fig. lo being very exceptional, and that of fig. 8 «9o6) GAXOXa-ffASCEXr FOKEsr OF MlSCOt' BF.iCIl 95 more typical. Again, the transition fn.rn the bnmd swales t.. the beaches of the sandy vmkIs is m,.st abrupt, f„r the latter are regular narrow close together with scarcely any hollows between, and also exhibit a curious barre-ness on their summits in marked en- tra.st to the better-dothed summits farther out (cmpare /igs. H and 13) Unfortunately the full importance of these features did no stnkc me m time for a study , f ,hcm on the ground, but ,uch data as I possess in n<.tcs and maps lead mc to believe that the swales arc much newer in origin than the beaches imme.liately inside them', ard that they mark the transition from an , Idcr series <,f Jx-aches which formed part of the original Grande Plaine extending far to withS'Ll^mT^'''")'/™"''""" """'■ '""'''"« """''• "•""^■'"8 <» »*»'<' °n the right, with Its sharp Unc of tmns.Uon to the woods; the trees are all white spruce. the'northof the present i.land, and a newer series formed by the rolhng of the plain down the coast, as described earlier in this paper. Al the facts I possess both as to geography and vegetation are con- sistent with this view.' Aside from the question of age, the swr'e zone differs physically from the grass plain by its greater shelter from the west winds, its lower level and greater nearness to the water-table, a probable increase of mineral nutrients derived from decaying driftwood and diffusion from the upland, and some slight accumulation of humus. T And it is sustained by the tradition of the R-sidents who say that the plain has X thi: Tu^d^r T" """"^ """°^' *''•"" "'^ memory'^f ^^su" 24 4?6. ,906 '^ ^'■" '""'^ '""^' '" ''""• ^'''- "'^'- S"^- N. B. No. BOTANICAL GAZETTE [^vavn becoming the bn,ad expanse of mea.low turf of the .wales an.l the jumper mats extemling anM\y with the a.Miti, „ f white «nni.-.. . c !• • '"'^ ""'"'•"'''"'"" <'f many young 2 ;-7 r Tr* '""" ""^' """"'^•^' "^"' 'here re.ult gla les "u inX t '^ ' """ '■'"""■"« "'^"^•''' "-^ ^"-■" -p^^i""y .. this plant, or else variously unite.! an.l cmbined wi-h patches ..f waxberp:. Hu.Isonia, an.l bluefxTry, exten.l greatly in .n'm t " a.verjng the crests as well as the slo,x.s of the .lunc Ua .7 i thus form a w,KKly net in the shelter of which several otheLms especially the beach ^., .ss, ,H-arly everlasting, and yart.w The new forms which a,,.ar .re, first of all. the common cn^wl, rry E. M- «.^r«. and the rock cranberry, Vacciniun, Vi,i.I,aJ.^ aH of them plants characteristic ..f dry upland rocky situations Less frequent are the little gentian. Genliana Amarel/a acutJZ t^^ corn f: •'%"'"-^ "^ ^^p^^'^"^ -" ^-^•^^•'^ Palustre (so much more highly developed in the woods) another curious mixture of xerophytic and hyZphytic f.,n^rVe ha" lro7f.r d " T^'T' ' '■"'' '^'^^-geneous assem! Wage of forms drawn from diverse natural habitats all the wav from r.,cky hills to bogs. They do not exist here th rctre tn forms of he flora .>f this region whose adaptations happen to fit he^ conditions, or wh.>se range of physiological ...leratioThappen o be great enough to permit endurance of the conditions here Of helps to emphasize h..w large an element of accident or incident •906] ovi.vo.V(;-.v.i.vt/.;.vr tonicsT oh .viscoi bi:.u„ 97 then, i, in „,,ap,a,i..„, „„., ,,.,, y^^^^.^^. ., .^ „,^^, .^^^ u . matdy pn.vc ... U- a ma..cr ..f ,hc 1.h..sc. an.l lar^c rather ,h«n ol thf ix ut ami mmutc. Finally it is in this same situation. u|.,n the u,.,Kr s1,.,h^ „f P r<-a ./«. develop. Stan.linK in ..,,en f..rmatio, . ,hey d , no ."terferc wah one am.th.r's «n.w,h, and in o.n...,u.n,; In-eome nunllr f~'^yy''"'l '"8^ J""'P«^ ■""' "" a ^l"pe an.l crest ot , dune Ix^a.h with a number of assocmted planU n..|e,l in the text; k >king s.,uth. ™!/"'Tr'"' ^•^^^•'\^>™'"^''"-' '" """ine and clothed to the Zrl' I 7 'T^- "' '''"""■"" "" ''""''^' ^"^ ^'^'^ '^'^^ reason hat the shrubs do as a compromise Ix-tween the greater wetness ... the hollows and the greater dryness of the beach summits. This habtt of growmg thus upm the sloj^^s. and not on summits .,r h.,1- lows has a most important effect uiK.n the physiognomv of the veg^ta ton m t ,s zone; for to it is due the ...nness of the swales, ^th thetr regular borders of trees, and as well the openness of the beach summits m the sandy woods later to Ik. noticed. Toward the sea the spruces are small and dense, and often show, as in fig n BDT.i.MCAL li.iZt.rTH (4U0IWT pronounud wind cffwls. In phucs many stwIlinK 'rif* m»y In- foumi, thouKli ihf «liMri»)uti<)n of ihisc U curiously irrcf{ular. In one plait- only did I find any . thir Irtr, und tha». was a (tingle .s|K.ti. mi-n (.f the prinu's pine, Pinm Punksiima. If It be axkctl why the white spruce is the first tree to develop on these plains instead of some other of those growing on the upland near by, I can only .>uy that an answer must wail until we know something alxiut the physiology of the wh" iinue and of other trees of the viiinity. We turn next to the swales, those long ojKn hollows car|K-ted by tt close turf, ami l)orderewk-<|f(i- of thoM- ki^>u|mi« I ovi-rltMtkfil. On anil iimnnn thit»f plants iKtur uthtrs, anions whkh I have colU'ilcd the followinn: the cytl»ri}(ht. Euphrasia ameriiiiHti lE. i>t)'uinali\); the huKle wird. I.yiopus uniflttrus (L. I'irniniiinii ; a liny everlasting, Anlinmirin miHlioua; a }K'arIwi>rt, Sagina profumhtm; the plantain, Plunlano mnjttr; the two eonim4)n ein<|uefoils, Polinlilla mmrgka and Annrrina; the fall dandelii n, Ltontodon aulumnair; and the while i lover, Trijolium rrhfns. These forma, in tomnion with the Kraskes, are all greatly d. e>iplur (the white spruce among it Ix-ing on a lixal elevation). and do!ived from diverse habitats, and are evidently a collection of heterogeneous stragglers from the neigh lK)rhood, held together by no stronger bond than ability to ckc out e.xistence in this inhos- pitabl" position. The majority belong to somewhat moist places, and they find an ample supply of water; for the water-table even in the driest summer is within a f(x)t of the surface, and of the sweetest water. Evidently it is not dryness which stunts the forms, but most likely, is I believe, paucity of mineral nutrients. The tuif represents the first closed formation we have met with, and lOO BOTANICAL GAZETTE [august competition may therefore determine some of its minor features, but to these I gave no attention. The turf reaches its climax in the open swales like those shown by fig. 10. In the woods it disappears, as will be noted under the next section; but towards the lower levels, especially towards the pools of standing water, it gives way gradually, by definite steps, to an assemblage of true swamp plants. The very first of these to appear in the k wcr i)laces in the swales is always the common blue flag, Iris versicolor, and characteristic scattered clumps of this plant may be seen in the foreground in fig. ii, in the distance Fig, I j.— Marshy swale, looking south; in the center a permanent pool with margin irodden by catUe; behind it are cattails and rushes, and back of them a thicket of poplar; on both right and left is sweet gale, and in the foreground is the blue flag. on the swale in fig. 8, and on the left margin of the swale in fig. lo. Next follows always the sweet gale, Myrica Gale, and after that low bushes of the balsam poplar, Populus balsamijera, a plant which forms very dense thickets and grows larger as the situation is more sheltered. Finally the pools of standing water arc reached, and on their margin occur cat-tails, rushes, and mare's tail, Hippuris vulgaris, with some other forms which I have not attempted espe- cially to study. The plants may be variously combined according to local circumstances, but a \cr>- typical arrangement is shown in fig. 12. It is plain that we are dealing here simply with an ordinary 1906] GANONC— NASCENT FOREST OF MISCOV BEACH lOI swamp, offenng nothing peculiar unless it be the small size of some of the plants, notably the poplar. But these places develop yet farther in time, and there come in after the poplar three willows : ScUix bdsamijera, S. lucida, and 5. Candida, forming very dense thickets, and apparently under congenial conditions. Finally comes in the alder, which appears to be mostly a form of the green alder, Alnus mollis, giving us the culmination of the swale thickets. THE SANDY WOODS. Inside the swale zone, through almost the whole length of the plain, extends a narrow zone, only some four or five dune beaches Fig. 13. — Typical sandy woods, just inside the swales, looking north; in the cen- ter a dune beach, bearing scanty beach grass and reindeer lichen, while on the slopes are small juniper mats with white spruces. wide, of remarkable sandy woods, whose characters are well shown '^y 1^&- ^3- Their most striking feature is perhaps the relative bare- ness of the tops of the beaches, which remain far m( re clear of vege- tation than do most of the beaches outside of them ; and this bare- ness, in conjunction with the presence of trees on the slopes and in the hollows, gives rise to curious vistas as shown by the photi graph. The bareness must have some physical basis, but I was not able to discover it. These dune beaches, further, are very narrow, low, and regular, with hardly any true hollows between, so that the turf i02 BOTANICAL GAZETTE [aogvst from the swales is very scanty, almcst wanting, in this Z' ne. Further, the trees, all of them white spruces, are much elder than those of the swale zone, the transition being commonly of the most marked abruptness. All of these features tend to emphasize the conclusion earlier given, that there is an abrupt physical difference between the beaches of these woods and those outside, a difference which, I feel sure, is one of age. The position of the zone would indicate that it possesses more favorable physical conditions as to water, mineral supply, and shelter than the zone outside, with which the large size of the trees is in agreement. But the bigness of the trees makes the barrenness of the beaches all the harder to explain. In their vegetation the sandy woods e.'hibit three divisions: the sparse, scattered beach grass and reindeer lichen (fig. ij) on the beach crests already mentioned; a few and sir ill patches of turf which can hardly obtain a foothold where the hollows are so small; and the juniper mats in the slopes and hollows with their well-grown white spruces. The mats, however, are no longer entirely creeping, for the junipers send up numerous erect shoots. With them persist several of the plants from the transition zone, especially the rock cranbeny, the three-toothed cinquefoil, the pearly everlasting, and a few others. But in addition new forms come in, especially and characteristically the bearberry, Arclostaphylos Uva-ursi, a rocky-hill plant, here creep- ing radiately over the sand, apparently under congenial conditions. Beneath the shelter of the trees appear some plants of the woods carpet which we may best consider under the next section. The trees themselves are of moderate size, rarely if ever over 20 feet in height. A fact of interest about the juniper mats, applying also to a less degree to the forest mat which succeeds it in the closed woods, is its very slight hold upon existence on the sand, for where teams cross and disturb it, the entire mat dies and soon disappears. Such instability shows forcibly how hard are the conditions of life in this situation, and how narrow the margin between success and failure. THE CLOSED WOODS. The climax of the sand-plain vegetation is reached in the dense though dwarfed mixed woods extending between the sandy woods and the upland, A typical view of the closed woods is shown by fig. 14. 1906] GANOXG—XASCEXT FOREST OF MtSCOU BEACH 103 Physically the situation is> much more protected than the zones out- side of it, and, lying at a slill lower level, it has a moister soil. The soil, however, is slill of sand, though it contains some humus from the decaying vegetation and must derive some mineral matter by diffusion and drainage from the upland. Very likely also the sand is shallower here than farther out (fig. 2), and hence some influence of the minerals of the underlying s^il may be felt, while in places an appreciable enriching of the soil must result from the decay of Fic. 14. — Typical closed woods, chiefly of white sprue . i 'it with some deciduous trees in the background; the closed forest carpet shows in the glade of the left fore- ground. the bodies of the walrus, formerly slain here in great numbers, as manifest by their semi-fossil bones.** These additional sources of mineral nutrients, however, by no means furnish a supply sufficient for the proper growth of the woods, for in every <'eature they exhibit marked depauperation as compared with the same species on the neighboring upland. In relation to the preceding zone, the closed woods consist essen- * Described more fully in a note in Bull. Nat. Hist. Soc. N. B. No. 24:463. 1906. 104 BOTANICAL CAZETTU [kvovn tially of a greater development of the juniper mats, which unite to form a complete unbroken carpet, together with a greater devel- opment, both in number and size, of the white spruce trees, to which are added some deciduous trees and shrubs. And where the hollows dip lower than usual, and towards the upland in places, this forest merges to alder and cedar swamp. We consider first the woods carpet. Morphologically it is a direct development of the juniper mats of the outer zones, though but little juniper, aside from occasional erect shoots, is left. With it persist some of its earlier associates, the rock cranberry, three- leaved cinquefoil, some grasses, the bearberry, and the reindeer lichen, varying in their respective development according to situa- tion. To these are now added dwarf plants of the bunchberry, Cornus canadensis, the twin flower, IJnnaea borealis amer'.cana, Pyrola chlorantha, the pipsissewa, Chimaphila umbellala, and an abundant brown moss, which has been identified for me by Mr. A. J. Grout as Aulacomnium paluslre, a typical swamp moss. Upon this carpet develop a few larger forms, especially the abundant wild sarsaparilla, Aralia nudicaulis, the gooseberry, Ribes oxy- acanihoides, the dwarf raspberry, Rubus Iriflorus, with others less conspicuous. We consider next the trees of these woods. First in importance and size, far surpassing all others in both respects, is the white spruce. It attains a height of perhaps 7. 5*", a diameter near the ground of perhaps 45"", and it exhibits over 100 annual rings, though per- haps some may be much older than those I counted, which were cut by the residents for wood. The next to appear is the balsam fir, Abies balsamea, becoming somewhat abundant and character- ized by a spruce-like arrangement of its leaves all around the stems. Then follow the red maple, Acer rubrum, the aspen, Populus trem- ulaides, the paper birch, Belula alba papyrijera (in very small trees however), and the mountain ash, Pyrus americana; while the common undershrubs are the red dogwood, Cornus slolonijera, and the black alder. Ilex verticillata. There are probably some others, but these I believe arc all that are notable. In especially low places, such as in certain hollows, and at the contact of plain and upland, the conditions verge towards those of t{ igo6J CANONO-NASCENT lOREST OF MISCOU BEACH lOS a swamp, and swamp plants apjK'ar — the iris, the sweet gale, some mints, species of Ga'.'um, and the dewberry; while the spruce gives way to the white cedar. Thuja occidenlalis, and the alder beiomes abundant, forming a dense jungle. But this is of less interest than the vegetation of the outer zones, and hence I gave it little study. Thus it appears that these woods present no features, size of the plants alone excepted, markedly different from those of woods preponderatingly coniferous in the neighboring upland, and they arc evidently tending towards the typical woods of this region — the mixed coniferous-deciduous forest. We have thus another illustration of that principle so important in physiognomic ecology, that vegetation, no matter under what immediate physical conditions it may be, is always tending towards a climax type, determined primarily by climate. CONCLUSION. In this paper I have tried to state the facts about the vegetation of a somewhat rcinarkable place, adding thereto some ecological comment whose chief value is to illustrate our ignorance of that subject. As I understand it, such descriptions as this aims to be may have three values. First, they can present to all who have interest in such matters a series of pictures, as vivid and real- istic as possible, of the vegetation of special places, and they are the more valuable according as they are the more clearly and attract- ively written and the more aptly illustrated. Second, they should help to supply information, badly needed by all of our manuals, about the natural habitats of the lommon or important species of plants. Third, they can form storehouses of facts about vege- tation upon which the future student can draw as the advance of physiological ecology gradually makes possible an understanding of the principles underlying physiognomic ecology. Such descrip- tive work can be done to profit by the studt.it whose work is perforce confined to his summer vacations, if he but bring to it time and care enough; but he should be content to describe well and to leave interpretation to the field physiologist yet to come. Speculation cannot of itself advance knowledge, and it can bring a subject into disrepute. It is only, I believe, through field physiology, the study left BOTANICAL GAZETTE [AUGUST M in field laboratories of fundanjcntal plar ' namits, that ecological knowledge can really be advanced. And the dynamical problems, as I see them, fall under these heads, in the ordT of importance: (a) physiological life-histories of species, (b) physics and chemistry of the soil, (c) nature of plant competition, (d) a better correlation of meteorological data with ])hysioiogical phenomena. Surra Coliege, Northampton, Mass.