Butler University Botanical Studies

Volume 7 Article 3

A Biological Spectrum of the Flora of the Great Smoky Mountains National Park

Stanley A. Cain

Follow this and additional works at: https://digitalcommons.butler.edu/botanical The Butler University Botanical Studies journal was published by the Botany Department of Butler University, Indianapolis, Indiana, from 1929 to 1964. The scientific journal eaturf ed original papers primarily on plant ecology, taxonomy, and microbiology.

Recommended Citation Cain, Stanley A. (1945) "A Biological Spectrum of the Flora of the Great Smoky Mountains National Park," Butler University Botanical Studies: Vol. 7 , Article 3. Retrieved from: https://digitalcommons.butler.edu/botanical/vol7/iss1/3

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Butler University Botanical Studies (1929-1964)

Edited by

Ray C. Friesner

The Butler University Botanical Studies journal was published by the Botany Department of Butler University, Indianapolis, Indiana, from 1929 to 1964. The scientific journal featured original papers primarily on plant ecology, taxonomy, and microbiology. The papers contain valuable historical studies, especially floristic surveys that document Indiana’s vegetation in past decades. Authors were Butler faculty, current and former master’s degree students and undergraduates, and other Indiana botanists. The journal was started by Stanley Cain, noted conservation biologist, and edited through most of its years of production by Ray C. Friesner, Butler’s first botanist and founder of the department in 1919. The journal was distributed to learned societies and libraries through exchange.

During the years of the journal’s publication, the Butler University Botany Department had an active program of research and student training. 201 bachelor’s degrees and 75 master’s degrees in Botany were conferred during this period. Thirty-five of these graduates went on to earn doctorates at other institutions.

The Botany Department attracted many notable faculty members and students. Distinguished faculty, in addition to Cain and Friesner , included John E. Potzger, a forest ecologist and palynologist, Willard Nelson Clute, co-founder of the American Fern Society, Marion T. Hall, former director of the Morton Arboretum, C. Mervin Palmer, Rex Webster, and John Pelton. Some of the former undergraduate and master’s students who made active contributions to the fields of botany and ecology include Dwight. W. Billings, Fay Kenoyer Daily, William A. Daily, Rexford Daudenmire, Francis Hueber, Frank McCormick, Scott McCoy, Robert Petty, Potzger, Helene Starcs, and Theodore Sperry. Cain, Daubenmire, Potzger, and Billings served as Presidents of the Ecological Society of America.

Requests for use of materials, especially figures and tables for use in ecology text books, from the Butler University Botanical Studies continue to be granted. For more information, visit www.butler.edu/herbarium.

Vand Dr. E5ther pel and Dr. Alice ler Adams ('25), A BIOLOGICAL SPECTRUM OF THE FLORA OF Mrs. Noe Higin- I THE GREAT SMOKY MOUNTAINS NATIONAL t, Texas. ]. E. P. PARK* B')' STANLEY A. CAIN The University of Tennessee the Dotan}' De­ The present study of life-forms of the Great Smoky Mountains in that the chief flora is based on the system of Raunkiaer (1934). Realizing the operation of the difficulties involved in correlation of meteorological and climatological capable students data with the natural occurrences of plants, Raunkiaer designed his a reasonable de­ life-form system as a means of defining what he called phytoclimates. sincere apprecia­ The theoretical basis was a familiar one in plant geography (Cain, ted" the celebra­ 1944) and may be expressed as follows: ( 1) Plants are limited in ~ ~ents, and to my their capacity to endure different environmental complexes. (2) lerously gave of There is usually a correlation between the morphology (growth-form, ~ .rRJES~ER. life-form) of an organism and its environment, i. e., there is a mor­ phological basis for adaptation in many if not all cases. (3) A plant, in its successful existence, represents what may be called an automatic physiological integ-ration of all the factors of its environment. It follows, if these are general truths, that the life-forms of the plants of an area are a measure of the environmental conditions, especially climate. It remains only to find the key to the plant-climate interre­ lations. Raunkiaer decided that the significant relationship was to be looked .for in the seasonal climates. (and all climates. but that of the constantlJ warm-humid tropical rainforest do have a seasonal rhythm in precipitation. temperature, or both). When growth is slowed or dormancy forced upon a plant by cold or drought the most critical tissues are the meristematic. Therefore, the amount of protection provided embryonic growing tissues and their success in enduring the unfavorable period represent a crjtical adaptation. It is for this reason that Raunkiaer selected the protection afforded the perennat­ ing buds as the principal basis for his life-form system. Raunkiaer's life-form system met with ready acceptance and has been applied widely, if sporadically, Over the world. This is because

.. A contribution (Botanical Laboratories, The University of Tennessee. N. Ser. No. 75) in recognition of the 25th Anniversary of the Botany Depart­ ment of Butler University. 11 the system is homo~eneolts. the life-forms are relatively few and hope that the whole flora evcn easily determined, and the results can be employed statistically in the this point of view, \lost of th l comparison of floras and climates. \Vhen the fl01'a of an area is between gcophyte and hemic.rjl>t analyzed and it is found that a certain life-form percentage exceeds and challlaephyte. .\nothrr n tile proportion which that life-form is of the normal spectmm1 the nect ion with snbcJasses, partic phytoclimate is designated hy that superabundant or predominating chamaephytes. Finally,life-fo Ii fe-form. Thus Raunkiaer spoke of the. phaneropbytic climate of :\jnuntains can not be reiined the tropics, the hemicryptophytic climate of the humid temperate zone, cerning the altitudinal occt1rren the chamaephytic climate 0 f arctic and alpine regions, and the thero­ in formation concerning the flo ~'()mmttnitics. phytie climate of deserts. Since such a th sible for years to come, I ha\'e After forty years there are still inSIl fficient data for a close de­ analysi~ is worth doing for it lineation o[ the major world climates, hilt certain general correlations The five principal classes 0 orig-inally pointed out by Raunkiacr have been confirmed. at least for {based, as we have said, 011 tb certain regions. lIuds dt1ring the unfavorable $ n following Raunkiaer's system in making a biological spectrum r crcasjn.~ protection: phaneroph of the Great Smoky Mountains flora, T uscd the preliminary cata­ (low perennials with buds du IOg'lle o[ the flowering plants of the Great Smoky A·fountains Na­ phytes (huds at the soil surf tional Park, a checklist in preparation ior several years ulHkr the Sll­ soil or under v,,'ater), and th pen'ision of i'dI'. Arthur Stllpka, Park !\aturalist. The principal ~eeds). These classes are suhj iield and herbarium work was carried on hy the late Professor IT. M. are so few in number that no Jennison and more recently by Professor .:\aron J Sharp. T have tion concerning the hemicrypt' not employed the list completely in its present form, haying- omitted treatment o( Sllbclas;;es. The I f 1'0111 con~ideration all varieties and forms except in cases where phytes) were elassified accordi a ~pccies is represented in the area only by a variety. l\lso, numerous hearing the percnnating buds ~scapes from agricu1tnre and gardens have been omitted where there tuhers, or roots. but the "ariol!. is any uncertainty as to their establishment. The plant list and my for present purposes. The as~jgnment of Ii fe-forms to the individual species are not here pub­ ,t reated according to f0\11' sube lished because the incomplete natme of the catalogue prevents its phytcs exceed 30 mders; mel' release at this time by the Park Naturalist. meters ; microJlhanerophyte.~ ~ Tn cases where I am not familiar with a species, its assignment l,hytes are less tall than 2 to a certain life-form often has been on a basis of previously published (abo\\t 25 cm.). classification, usually by Ennis (1928) or McDonald (1937), Cases The ana'lysis of the flora if of Cjuestionable life-form status for the Smoky !\fountains area and including well-estahlished ad" cases of disagreement between authors have been settled by reference to occupy the highest altirudi to herbarium material and the literature, I am aware of the probability the tops of the mountains at of incorrect assignment of certain species to life-form classes and corresponds in general with subalpine forest Tt is pene l The normal spectrum, based on 1,000 carefully selected species, is no more in the "alleys and lower ~ap than a yardstick, a statistical .approximation of the life-form percentage composi­ tion of the flowering-plant flora of the world as a whole. heath bald:; (Cain, 1931). I 12 relatively few and hope that the whole flora eventually may bc studied in the field from ,d statistically in the this point of view. \rost of the doubtful cases fall on the boundary flora of an area is between geophyte and hemicryptophyte and between hemicryptophyte percentage exceeds and chamaephyte. Another need for further fichl work is in con­ trmal spectrum' the nection with suhclasses, particularly among hemicryptophytes and It or predominating chamacphytes. Finally, life-form studies of the flora of the Smoky ~ lfoP h"ytlc c1ullate 0'f \1ountains can not be refincd greatly without more knowledge con­ fnid temperate zone, ccrning the altitudinaL occurrences of the species and Inore complete lons, and the thero­ inforlllalion concerning the floristic composition of the major plant cOlllmunities. Since such a' thorough-going study may not be pos­ lata for a close de­ sible for years to COllIe. I have assumed that the present preliminary general correlations analysis is worth doing for its immediate value. firmed. [It least for The five principal classes of the life-form system of Raunkiaer (based. as we have said, on the protection afforded the perennating iological spectrum buds during the l1nfavorable season) are arranged according to in­ ~ j)reliminarv cata­ creasing protection: phanerophytes (trees and shrubs), chamaephytcs ~ ­ kv Mountains Xa­ (low pel-cllnials with buds close to the ground snrface), hemicrypto­ phytcs (buds'at the soil surface), cryptophytes (buds beneath the ~~ars uncleI' the SII­ '."t. The principal soil or under water), and therophytes (annuals, buds within the I seeds). These classes are subject to subdivision. The chamaephytes te Professor H. ~f. are so few in number that no breakdown was made. 1\ly informa­ Sharp, I have IJ tion concerning the hemicryptophytes is inadequate for the detailed fl, having omitted treatment of subclasses. The geophytes (the major group 0 ( crypto­ pt in cases where phytes) were c1assi fied according to whether the subterranean organs ,. Also, numerous bearing the perennating buds are rhizomes, bulbs, stem tubers, root ~itted where there tubers, or roots, but the various groups seen1 to have little significance plant list and my for present purposes. The phanerophytes, however, were easily are not bcre pnb­ treated according to four Sll bclasses based 011 height. NT egaphanero­ ogl1e prel'ents its phytes exceed 30 meters; mesophanerophytes are between 8 and 30 meters; microphanerophytes are between 2 and 8 meters; amI n

HaUJlkiacr'~ ':\onnal spectrum TOlal flora High altitude species Norma) Life·form No. spp. Per Cellt No. spp. Per cent Spcctr\lln 1,000 species

Phanerophytes 223 19.5 64 21.2 ~6.0 Referring again to table 1 (Mega-H) ?!) 2.5 2 0.6 1'ion of the total flora of the (Meso-H) 73 6.4 18 G.O (Micro-") 70 G.l 22 73 of the normal spectrum. Cr. ( \'ano-") 51 4.5 22 7.3 one-half times the normal. Chamaephytes 20 1.7 7 2.3 9.0 percentages of the normal H emicryptophytes 595 52.1 170 5G.5 260 phanerophytes, which are I CrYlltophylcs 173 15.1 52 17.2 G.O That this spectrum is tYPI Thcrophytcs 131 11.5 8 2.G 130 ate climates is seen from th Totals 1142 99.9 301 99.8 of the same pattern anel var" ~mokies in the latitudinal TABLE II Cape Breton is somewhat rni Some life-form spectra for eastern North America represcnting areas of a mountainous area in whit:: humid mesothermal and microthcrmal climates charactcri;t;ed as hemicrypto­ dimates are characteristic pbytic. \Iountains spectrnm. excee ]Ireton, and the spectrum i· Hemi· Flora and alHhor .Phanero~ Chall~i1c· crypto· CryplO­ TtlCI"O· accentuates certain charact r\umhcr of species ph)'!es phytes phYlCS phytes phytcs other classes of life-forms .'\ Jabama. Ennis, 1928 The high position of phan 2,012 species 17.0 3.1 47.8 17.1 14.4 the series, of cot1r~e) is an Mississippi, Ennis, 1928 area together with the vari~ 1,724 species 17.7 3.1 49.4 16.2 128 14 thern hardwoods go to TABLE II-(Continued) )ted by grassy balds. Some life-form spcctra for eastern North Amercia representing areas of ~ I for comparison. humid mesotherl11al and microthermal climates characterize.d as hemicrypto­ phytic. s conspicllonsly repre­ Hemi· d is of a type generally Flora 3.nd author Phancro­ Chalnae· crypto· Crypto· Thero· perhaps more accurate Number of species IJhy{e~ phl'tes phYles phytcs phytes ~-form percentage clis­ Great Smoky Mountains ~ries of closely related 1,142 species 19.5 1.7 52.1 15.1 11.5 :fefinite to pronOllnced Dull RUIl, Virginia, Allard, 1944 1son. This can be iI­ 980 species . 18.2 1.4 51.7 11.3 17.u Conneeticut, Ennis, 1928 'at Smoky l'vfol1ntains 1,453 species 15.0 1.9 49.4 21.7 11.7 ~a, table II. Cape Breton, Ennis, 1928 637 species 14.1 1.8 51.3 25.6 6.7

Indiana, McDonald, 1937 Ihl: ilora ni thl: highest 2,109 species 14.3 1.9 49.0 18.0 16.7 Iowa, Ennis, 1928

~----~-- 1,320 species 14.8 1.0 48.6 20.9 14.2 R:=tLll1l.:ja.cr'~ I allitude :'ipccics Normal Norma) spectrum spp. Per cent Spectrum 1,000 species 46.0 9.0 26.0 6.0 13.0

64 21.2 ~6.0 U.6 Referring again to table I, it is seen that the hemicryptophytic por­ 6.0 tion of the total flora of the Smokies is 52 per cent, just double that 7.3 of the normal spectnull. Cryptophytes, with 15 per cent, are two and 7.3 one-balf times the normal. These excesses ovel" the corresponding 7 _? .• "l 'i.U percentages of the normal spectrum are primarily at the expense of 170 5(,.5 26.0 52 17.2 (l.O phanerophytes, which are less than half the normal for the world. S 2.6 LUI That this spectrum is typical of the spectra for the humid temper­ ---~~ ate climates is seen from the data in table II. These spectra are all 301 99,8 --_._------_._­ of the same pattern and vary only in small ways. The position of the Smokies in the latitudinal series from Alabama and Mississippi to Cape Breton is somewhat misleading because we are here dealing with repre~enting areas 0 r a mountainOllS area in which much of the vegetation and the 'higher tcrizerl as !Je111icrYIllll­ climates are characteristic of higher latitudes. Thus the Smoky ------_._-- 1V[ollntains spectrum exceeds in hemicryptophytes even that of Cape Ii· Ereton, and the spectrum for high altitudes in the Smokies (table I) to· Crypto­ Thera. ~s pJ1J'tt'S J)ilYle~ accentuates certain characteristics of the flora as a whole. All the other classes of life-forms are increased at the expense of therophytes. The high position of phanerophytes in the Smoky spectrum (within 17.1 144 the series, of course) is an expression of the southern position of the

162 128 area together with the variation of conditions resulting from the alti­ 15 tudinal range. Its low position for cryptophytes is due entirely to able with close study as can the absence in the mountains of marshes, ponds. ane! lakes and the alliances, as follows: Ae. consequently v~ry small number of helophytes and hydrophytes. sugar maple-silver-bell, yel It is not within the purposes of this paper to discuss life-form Tsugion, including the hem spectra in general, especially how the spectra for the hemicryptophytic segregates. I have statistio climate differ from those of steppe, desert. tropical and other climates, each sampled by a plot of i but the similarities of the spectra in table II indicate the close similar­ the shrub and fidel layers ity in climate of the areas of the deciduous, summer-green forest re­ under spring conditions, a1 g-ions of eastern United States. They do not differ as to the fUnc];L­ conditions. At each statiol mental type of climate, but only in details of length and coldness of rats of one sq. m. area. wI' winter, etc. quadrats of six sq. m. are, The type of life-form statistics employed in the preceding section contains tbe results of the COl1stan~ depends on the use of total Boras 0 f whole areas. In such an analy­ all the species by sis one species counts as much as another irrespective of its role in tables will not be repeate the structure of the vegetation of the area. The other use of Ii fe­ subsequent life-form spec forms is their employment in the description of vegetation types in­ The constancy percent; cluding communities' of all sociolog-ical rank. The description of relationship between the \'cgetation partly at least in terms 0 f life-form and especially the Ii fe­ number of stations in whi forms of the dominants is an ancient practice in plant geography, as Frequency is the same s( witnessed by such terms as woodlancl, bushland, steppe, etc. In com­ vidual quadrats. rather tl plex communities the whole phytocoenosis may be referred to in The following lists of terms of the life-form of the dominating layer. Lippmaa (1933) wo~ds as determined by has developed a system of vegetation description which' depends upon form. The nomcnclaturl the separate analysis of each synusia of the phytocoenosis, the syl1usiae menL in each group is being single-layered communities each composed of plants of one or percentages. The result (,f two closely related life-forms. Raunkiaer, however, illtrodnced the most useful, graphically descriptive employment of life-forms in PllANER01'llYTES. ?I cOllll11Unity analysis. I say this becau:-;c his method uses :-;imultan­ C III us odan.ch'n., Bct,f!a eously the complete li fe- form data for the community and statistical Fagus gmndifolia.. Fra information on the quantitative roles of the species. That is to say, Pad us 'iJirg·i,.,,·ia.no, CastaJ he developed life-form spectra for plant communities in which the Tulipastl'lr.l1£ aelllllillatrtl percentages for eacb life-form are based on their total frequenc)' .\IesophanerophyLes: H points resulting from quadrat analysis. For the Smokies it i:-; possible r Yaseri, A err /'cnnsyl7; lle~; for me to apply this methotl only to the cove hardwood forest complex lo.cv'is, opaca, 11cI ulc fur which some quantitative data recently ha\'e been published (Caill, Clo.dmstis 1,.11 eo , O.ryc/CJ 1943a) . thenoeissus (juillqur.j'olia The cove hardwood forest complex is frequently consitlered to Mierophanerophytes: .A be a unit, especially by foresters, and is sometimes designated as virg'iniano, llex tlwnti( undi fferentiated or mixed mesophytic forest by ecologists. Even in the limited Greenbrier area of the SlI10kies it is, however, reeogniz­ , /\ brief nute 011 the ;;< 16 tophytes is due entirely to able with close study as consisting of seven minor forest types in two • ponds, and lakes and the alliances, as follows: Aesculion, including the buckeye-basswood, bytes and hydrophytes. sugar maple-silver-bell, yellow birch, and beech segregates; and the s paper to discuss Ii fe- form Tsugion, including the hemlock-beech, hemlock tuliptree, and hemlock tra for the hemicryptopl1ytic segregates. I have statistical data for 31 stands of this complex forest, , tropical and other climates, each sampled by a plot of ahout one acre area. Sample plot data for. 1I indicate the close similar­ the shrub and field layers were obtained from 10 of these stations tiS, summer-green forest re- under spring conditions, and fr0111 nine other stations l1nder summer not diffcr as to the funda­ conditions. At each station the vernal flora was sampled by 10 quad­ of length ane! eoldness of rats of one sq. m. area, whereas the aestival ftora was sampled by 10 fluadrats of six sq. m. area.' The cove hardwoods paper cited abo\'e y in tbe preceding scdilJl1 contains the results of the quadrat study and presents the results for Ie areas. In such an anal v­ all the species by constancy and ·f requency percentages. These long irrespective of its role in tables will not be repeated here, but they provide the data for the The other use of life­ subsequent life-form spectra. ion of vegetation types il1­ The constancy percentage of a speeies for a community type is the rank. The description of relationship between the total number of stations studied and the Ie nn and especially the 1i fe­ number of stations in which the species occurred in the sample areas. tice in plant geography, as Frequcncy is the same sort of concept, but it is based on the indi­ hland, steppe, etc. In (,Olll­ vidual quadrats, rather than on the data for stations. . is may be referred to ill layer. Lippmaa (193.1) The following lists of species present the flora of the cove hard­ 'ption which depends upon woods as determined by the above procedure. and arranged by life­ phytOl'oenosis, the synusiac form. The nomenclature follows Small's Manllal and the arrange­ posed of plants of one or ment ill each group is one approximately according to constancy naer, however, introduced percentages. The resulting life-form statistics compose Table lIT. p)oyment of lifc-forms in . PUANEROPHYTES. 'Megaphanerophytes: Ts/./ga mnadcllsis, Acs­ hi. method uses simultan­ culus octanell-a, Bctula alleghcnicns':s, Saccharodendrun barba/1I111, Dtnmunity and statistical [7o.gus gmndifolia) F1'axiJtl~s americana, Lir·iodc·ndron t-ulipifera, e species. That is to sav, , Padus ,-irgill.iana, Casta:nea dcntata, P-ieca l'ltbe11,s, RuIo.·eer 1"[tbru1'Jl, nunnnities in which the TuJr:pastnl.m ae'U'ntinatuln, QUfrcus maxin.w, Q'ltcre-us montana.­ on their total f requcncy l\Tesophanerophytes: Halesia monticola, Ti/ia negleeta, Magnolia r the Smokies it is possibk rrascri, AreI' penns'ylvanieum, Hicoria cordifo·rm·is, Amclanehier hardwood forcst complex laevis, !lex opaea, BetuLa len.ta, Wallia eince-rae, Cynoxylon floridtf.1n, live been published (Cain, . Cladmsll:s lutea, Oxydendrum arbareum, Robinia pseu.doacaeia, Pa1'­ thel'1Oeissus qltinquefol-ia (liana), Al'istoloehia macrophylla (liana).­ frequcntly considcred to Microphanerophytes: Aeer spieatu11't, Sv·ida alternifolia, Hammac/is slmetimes designated as virginiana, llex monticola, A·ral.ia spinosa, Vibtwnum lantanoid.es) t by ecologists. Even in it is, however, recogniz­ 1:\ brief note on the sampling problem is appended at the cno of this paper. 17 ErtOnymus americanus. Sambucus pubens.-Nanophanerophytes: Eu.­ ~'ery rich forest community (' OWVlflltS obm'al/{s, Hydrangea a.1'borcscens, Pyntlar-ia pubera, Gros­ nanophaneropbytcs) in prime sularia Cynosbati. climatic and soil conditions u CI-n,r"EPBYTES. Alsiue Icnnesseensis.. Mitchella repens, Phlox .second conspicuous change { stolonifcra, Scdum ternatum, Cymophyllus Fmsel'i. from 15.1 to 25.8 per cent cr I-IE:I1ICRYPTOPHYTES. Tia:rella cordifolia, Astel' acu1'n;nalus, Vi­ increased, from l.i to 4.4 pe:r ola s01'01'·ia, Viola blanda.. Solidago CU1'tisii, Eupat01rittm urtieaefolhun, be of much consequence. Nabalus sp., Viola hastala, O:ralis montana, Ratluncnlus 1'ecurvatus, hcmicryptophytes, which drop Monarda didj'11W, Poa eHspidata, Osmorrhiza Claj'toni, Viola Ca1~a­ pbytes, from U.S to 3.4 per c dCIiSis, Viola 1'ostmta, Galiltnt tl'iflorum, Care:.: flexlwsa, Carex plan­ taginea, Rudbeckia lacini(/Ia, Hepat·iea aOtta, Cm'cx ausl1'o-caroli~tiana, Viola rotundifoha, Crjlptotaenia wnadensis, Viola e'l'iocarpa, Cam­ pan.u1astrul1·t m'Jlt!'rieanuUt, Ge1,(m wnadcnse, Pera'l1Iiunt ophioides, Viola pallens, Viola eucullala, .lolitella diplryl1a, Geranium maculatum, Zi.;ia Bebbii, luncoides bulbosu11't, Carex prasina, Ranu:nculus fasi­ Spectra based on species c'lrlaris .. Aiicranthes '/'Ilicranthidifolia, Junco·ides saItuense, Ranu11C1tl/{S I. Total flora allor/i,'us, Heuchera ame'l'icmw, Senecto Rugelii, Taenidia'i1ttegenima, 2. field layer, vernal aspect lUeph'ilia hirsula., Solidago axillaris, Hystrix Hjlstr'i~', Houstonia P/'{.1'­ 3. Field layer, aesti\'al aspect !'ltrca, Meibo11w nudiflora, Juncus te'/lui~·. Thahrtnon dio'/cum, Pani­ Spectra bascd on constancy Clt/'ll sp? Carex stellata, Asclepias exa1tata, Lys-itmachia q~tadrifolia. 4. Total flora 5. Field layer, vernal aspect CRY PTOPHYTES (all geophytes). Erjllhronium americanl.l1n, 6. Field layer, aestival aspect Del1.laria d-iphyl1a, Anemone qu£nquefolia, UTticastrum divo.:riwtu111-, Spectra based on (requetlc)" 13/cuculla canadensis, PMta:r trifolium, Caulophyllum thalictroides, 7. Field layer, vemal aspect Clay/onia 'i:irginica, Tlrillium erectum var. albun~, Cimicifuga a11~er·i­ 8. Field layer, aestival aspect ((lIW. Poljlgonatu,11't bij'lol'u1n, Disporum lmtgu.ginos·/./.'nt, TO'vara 7nl'­ .'Jinian.a... Hj,drophylll./1-n canadetlse, Mcdcola virginiana, Validallium \Vhen the phanerophytes I con~ide tri('o(( m·l'l .. J7 era.trum 7Jin'd e, Arisael'JUI quinat-u'n'/" Podophyllum pelt(/.­ herbaceolls syllllsia is tmn. X eniatrm·1'/. u1nbelluJatl.ll'n, Arisae'/'/l.a /riphjlllwltL, Chrosper-ma changes, so comparison willi JIIu.\·weta.l'1cU'I'I'I, SYlldesmon thalictroides, Vagncra l'aCeltlOSa, Lilin'ln aestival aspects of this layerl supcrbH111. Trillium gmndiflo-rum, Actea alba, Diphylle-la cy'l11osa, Ilounced representation of (j Collinsonia ranadt'1'/.sis, C'i"caea lat·ifolia, Bicuwlla cucullaria, Clin­ per cent, and all geophytes ultim~ Ionia boreahs, Glycine A pios, M onol'ropa uniflora, CareX' pennsyl­ slowly developing but ,'a·n.u·(( . the aestival flora (61.3 per c.! TH EROPI-) YTES. Impatiens pallida, Ga.lil.lllt eircaezans, Cuscuta cryptophytes for the normal sp., Adi('ea pumila, Phacelia fimbriata, Gahum aparine. Smoky Mountains flora, an Lines 1-3 of table III present life-form spectra for the cove hard­ vernal flora of the cove hal' woods in which the species as such form the basis of the statistics. Seven geophytes in the v' The total flora spectrum, line 1, differs strongly from that for the val aspect, apparently beca.u" Park as a whole, table 1. Phanerophytes increase from 19.5 to 36.3 cycle and retreated again to p"r cent, a change that would seem to be due to the fact that this is a iU.1lt a11tericanu-ln, Bicuculla 18 Iphanerophytes: £14­ very rich forest community (14 mega-,.15 meso-, 8 micro-, and 4 tlaria puhera, Gros­ nanophanerophytes) in primeval condition with rather tlni form micro­ climatic and soil conditions under the arborescent dominance. The· ltella repe11.s, Phlox second conspicuolls change from the spectrum for the whole Park is / ~n. from 15.1 to 25.8 per cent cryptophytes. Chamaephytes likewise are tel' ac/{.]-nillalus, V i­ increased, from 1.7. to 4.4 per cent, but these figures are too small to wiutn /lrticac!oliHII't, be of much consequence. The increases are at the expense of the u:nc/ill/s 1'ecur-vallls, hemicryptophytes, which drop from 52.1 to 30.1 per cent, and thc1'o­ 'a:'J'toni, Viola W1W­ phytes, from 11.5 to 3.4 per cent. ~.1:uosa, Carex pIa1/.­ t austro-cal'ohn·iana, TABLE III 10. criocarpa, Ca11'l­ Life-form spectra for the primcval cove hardwoods of the Sl11okie~. ramium ophioidcs, I . IraJ/1·lt1n 1J!acula/llm, Ph ell H Cr Til , RanunCl/lus fasi­ 'uense. Rallullcllllls Spectra based on speeies Spccie, cllidia inlcgerrima, I. Total flora 36.3 4.4 30.1 25.8 3A ]]3 ',ix, Houstonia pur­2. fo'ield layer, vema) aspect .. .. 7.0 47.2 40.3 5.5 72 3. l'ield laycr, acstival aspect .... 6.6 61.3 29.3 2.6 75 11 1 '/1- dio'icw!1t, Pani­ Spectra based on constancy Points ~achia quadrifolia. 4. Total flora 3Ll 5.9 331 26.9 J.O 4,052 111m (luter-tcalllf.nt, 5. l'ield layer, vernal aspect 9.0 48.0 39.0 4.0 2.790 tnou di'l'aricatu11t, 6. Field layer, aestiva( aspect ... . 8.1 58.1 31.1 2.7 2,877 Ilu.m /halictroidcs, Spectra basw on freql1ency Points 7. Field layer, vernal aspect .... 11.0 43.3 41.9 3.8 1,309 al1~e1'i­ 7imidfuga 8. Field layer, aestival aspect . . .. 14.2 58.6 23.9 .13 J,25S )//111. Tovara ,'ir­ ialla, Vahdallill1n \Vhen the phanerophytes are left out of consideration and the 'doPhyllm/t pella­herbaceous synusia is considered alone there natmally are percentage u"m, Chrospr.r1l1il changes, so comparison will be made only between the vernal and {aCC11I o.W, Liliu1It aestival aspects of this layer. The most striking result is the pro­ ph.I'llcia CylllOsa, nounced representation of cryptophytes in the vernal flora (40.3 cllcullaria, CNn­per cent, and all geophytes) and the preponderance of the more Cal'cx pennsyl­slowly developing but ultimately rank-growing hemicryptophytes in ~he aestival flora (61.3 per cent). The contrast between six per cent (lc:::ails, Cuscu/a I cryptophytes for the normal spectrnm, 15 per cent for the whole Ie. F Smoky I\:J ol1ntains flora, and 40 per cent of geophytes alone for the r the cove hard­ vernal flora of the cove hardwoods is a 'very striking phenomenon. •f the statistics. Seven geophytes in the vernal flora which were absent in the aesti­ 1m that for the val aspect, apparently because of having rapidly completed their life­ IIIl 19.5 to 36.3 cycle and retreated again to their subterranean organs, are El'ythl'Ol1­ ct that this is a iu,m americanU111 , Bl:cuculla canade1tsis, Bic·c{.culla cU('ullaria, Panax 19 trifolium, Cla}'tonia '(.if,rginica, Trilliwlt :graHclifloru11l, and Actea alba. not at hand for the cove hard The seven other geophytes unsampled in the aestival flora are the on frcquel?cy..points are .more longer enduring Veratrum viride, Podophyllum peltatu11l, Lilium various life-forms than are the superlmm, Diph),zlc·ia C)/1fwsa, Collinsonia canadensis, Circaea tatifolia, alone. and Clinlol/ia borealis. The six geophytes sampled in the aestival In table IV are spectra or. c flora and not in the vernal'flora are Tovara virgi11iana, Chrosp'enna data for which I helieve to be' /ltuscactoxicu-m, Syndesmon thalictroides, Glycine Apios, i\lIonotropa' mesophytic climax are strikin u/liflora, and Can.t' pennsyh·anica. Although the differences in appear to be wholly comparable geophyte listing for the vernal and aestival aspects of the field layer cally, the higher altitude of the are partly due to sampling and partly due to normal variability in the higher latitude of the Ohio s composition of such a rich community, it still seems that the first \:ion segregate" has a conspkl' group of species listed immediately above represents a distinct excess is maintained by the Laurenti of geophytcs in the vernal flora. Furthermore, that the cryptophyte­ to which is added a striking . hcmicryptophyte relationship in the vernal and aestival societies is trends at the expense of ph a true one is substantiated by the constancy and frequency studies within ~he deciduous forest en: (lines 4-8, table III) where very similar ratios reoccur. expectation according to Rauo experience with regional sped It often happens that the use of quantitative data for species in ~oci~tion,s also produce simila the development of life-form spectra produces strikingly di fferent fragmentary, there is a sugg' n:sults from spectra based solely on species with each species having be used better to distinguish c the same weight. In this case, however, the various field spectra are than areal and regional speet all of the same pattern, as shown by lines 2, 5, and 7 for the vernal floras as large as those of state aspect and lines 3, 6, and 8 for the aestival aspect. This reslilt would the Great Smoky Mountains, .1 seem to be due to the fact that the cove hardwoods flora is a very rich climate and habitat for any bu one in which no small number of species is clearly predominant. This One technical point conC(' situation is in strong contrast to the more impoverished hut compar­ added here as a ;ort of ;lppend' able field layers of the l11esophytic deciduous forests of Europe ent sizes of quadrats in the sam! (Lippmaa. 1938, and Raunkiaer, 1934). The most interesting lIew of the field layer. I have di. feature of the compared spectra (obtained from species alone, con­ 1932, 1938, 1943b), but thes st,ancy points, and frequency points) is the steady increase of chamae­ in table III that the constancy phytic percentages from 7.0 to '9.0 to 11.0 for the vernal aspect and and for the aestival aspect 2,8: from 6.6 to 8.1 to 14.2 for the aestival aspect. tel' was about six times that microphytocli~ Theoretically the most signi fkant spectra for the holds for the frequency points mate of the field layer of the cove hardwoods are those hased on fre­ respectively. No such close al quency points hecause they provide better data on the roles of the constant-size quadrats been ~ species in the community. Raunkiaer showed that frequency points The larger size for the aestiv~ approach density values when numero.us .small quadrats are used. stature and area of the sum~ The best way of preparing statistical life-form data for. spectra would prohahly be by the use of dominance data because of. the biological significance of dominance in a community, but such information is 20· ?YUnt, and Aciea alba. not at hand for the cove hardwoods. At any rate, the spectra hased lesti"al flora are the on frequel}cy,points are more revealing as to the role played hy the 1m pcltatu1n, LilimH various life-r'orms than are the spectra based on species composition nsis, Cifcaca lati.folia, alone. npled in the aestival In table IV are spectra or, certain American forest associations the 'giniana. ('hvosp'cnna data for which I believe to be comparable. The two spectra for mixed 1C A pius, J1onoifO pa . mesophytic climax are strikingly similar. The communities would I tbe differences in appear to be wholly comparable ecologically and probably are climati­ cts of the field layer cally, the higher altitude of the Tennessee stands compensating for 10nnal variability in the higher latitude of the Ohio stands. The Long Island oak "associa­ seems that the first tion segregate" has a conspicuous increase in chamaephytes which ~nts a distinct excess is maintained by the Laurentian maple "association segregate" and that the cryptophyte­ to which is added a striking increase in hemicryptophytes. These aestival societies is trends at the expense of phanerophytes and cryptophytes, and ld frequency studies within the deciduolls forest climax formation, are in accord with reoccur. expectation according to Raunkiaer's theory of phytoclimates and data fill' species in experience with regional spectra. The two studies on Populus as­ strikingly different sociations also produce similar spectra. Although the data are each species baving fragmentary, there is a suggestion that association spectra may )liS field spectra arc be used better to distinguish climatic differences' and delimit types ~nd 7 for the vernal than areal and reg'ional spectra. Specifically, spectra based upon This result would floras as large as those of states, or even areas like Cape Breton and flora is a very rich the Great Smoky ?vlountains, include too much variability of micro­ predominant. This climate and habitat for any hut the most general comparisons, :rishecI but compar­ One technical point concerning the quadrat technique may bf' forests of Europe added here as a ~ort of appendix, and that concerns the use of di'Her­ I. . bst I1l terest1l1g new ent sizes of quadrats in the sampling of the vernal and aestival aspects species alone, con­ of the fielel layer. I have discussed this problem elsewhere (Cain, ;lcrease of chamac­ 1932, 1938. 1943b), but these data offer a new approach. Notice vernal aspect and in table III that the constancy points for the vernal aspect are 2,790 and for the aestiva1 aspect 2,877 although the sampled area in the lat­ the lIlicrophytocli­ ter was about six times that of the former. The same relationship hose hased on f rc­ holds for the frequency points where the numbers are 1,309 and 1,258, the roles of the respectively. No such close approximations could have resulted had frequency poinls constant-size quadrats been l1sed in the sampling of both aspects. .iadrats are tlsed. The larger size for the aestival aspect was necessitated by the larger for spectra would stature and area of the summer plants. oL the biological :h in formation is

21 TABLE IV c. Spectra obtained frem entirely of the pattern of ones Life-form spectra of certain American deciduous forest associations. the large nnmber of species inv No. in numbers and Illass by one Of S»t:cies Ph eh H Cr Th 4. A comparison of certai close similarity between clo~l Cove hardwoods mixed mesophytic climax, Great Smoky Mountains 113 36.3 4.4 30.1 25.8 3.4 Southern Ohio and Eastern Ten Mixed mesopbytic climax, Cincinnati and the Alberta and.\! ichigart [. area. Withrow, 1932 127 33.6 3.9 34.4 23.4 3.9 Aar "association sc~regate" l"e QuercetuLll montanae, Long Island, position' in its relatiYc1y high Ch :.iew York, Cain, 1936 92 34.8 10.9 32.6 20.()" 1.1 Aceretum saccbaropbori, Laurentian a regional spectrum in the cas reg-ion. Dansereau, 1943 346 17.0 10.0 56.0 15.0 2.0

AS[len associatioll, Northern Lower '\1.1. \ ltD, H. ,\. All analysts of th Michig-an. Gates, 1930 310 22.9 3.9 47.J 16.1 to.3 of Virginia using Rannkiaer's PO[llar association, Central Sci. 34 (4): 112-119. 194-t ..\Iherta. Moss. 1932 170 25.8 l.R 48.2 17.1 7.0 C\IK. STANLEY A. Ec()lo~ical .1r ~orth a~ ,. Tneluding 2.1 % Monotro[laceae. Mountains of Carolina 1931. SUMMARY Concerning i\~onogr. 2: 475-501\. 1932. 1. The Iife- form spectrnm for 1,142 species 0 f the Great Smoky The compositij j\[ot1l1tains National Park is entirely similar in pattern to other spectra S[lring Harbor, LOIlg- Islant!, \l for humid mesothermal and microthermal climates and the eastern .'\mer. MidI. :-':at. 17 (4): 725-J American deciduous summergreen forest region. Hemicryptophytes ____ The species­ predominate, being the life-form of 52 per cent of the total flora. 573-581 1938. ____ The Tertia!"}' 2. Species of the area known to exceed 4,500 feet elevation and or the Great Smoky:\lo\1l1ta grow in what is essentially the sprnce- fir belt produce a similar spec­ Club 70 (3): 213-235. 1943a. I rtlm [0 that of the Park as a whole, but with all classes slightly in­ ____ Sample-plot creased at the expense of therophytes which drop from 11.5 to 2.6 \Vyorning. !\mer. Jour. Bot. 30 ____. FOlfl/(!atirnls ~ per cent. & Bro., l\e"... York. 1944. 3. Statistical data (on constancy and frequency) for the flora D \ :-'·SERF..·\ ~r, PIERRE. I.'Frablie of the virgin cove hardwood forests of the Greenbrier region of the especes. Contr. Inst. Bot. Unl Park allowed a special analysis of life-forms in that community: El\.'\ls. B. The life JOHns of relation to climate. Conn. S a. In comparison with the Park as a whole, the cove hardwoods G \T[S, F. C. Aspen associatioll show an exceptionally high percentage of geophytes (25.8%), and 90 (3) : 233-259. 1930. a high percentage of phanerophytes (36.3%) for the latitude. L,PPM,\A, T. La methode des ass b. Comparison of the vernal and aestival aspects of the herba­ des associations. .'\cla lllst. 1933. ceous layer of the CO\'e hardwoods showed the importance of geo­ ___. :\real und phytes in the vernal flora (40.3%) and of hemicryptophytes in the Problem del' Charakterarten aegtival ilora (61.3%). Univ. Tartuensis G (2): I-I!.. 22 c. Spectra obtained i rem constancy' and i requency points arc us ioresl associations. entirely of the pattern of ones from species per se. This is due to -----,--- the large number of species involved and the lack of preponderance

Ch II Cr Th in numbers and mass by one or a few species. 4. A comparison of certain iorest-association spectra rel'eals close similarity between closely related associations. sllch as the -1.4 30.1 25.8 3.4 Southern Ohio and Eastern Tennessee mixed mesophytic associations :1.9 3-14 2:>.4 .1.9 and the Alberta and IVlichigan PapH/us associations. The Laurentian AceI' "association segregate" reveals the influence of more northern 10.9 32 Ii 20,(," 1.1 position' in its relatil'ely high Ch and H percentages better than does a regional spectrulll in the case of Cape Breton. 10.0 56.0 15.0 20 ~._---_.. -'--- .. _-.­ LITERATURE CITED

3.9 47.1 lli.1 10.3 '\l.I.IIW. H. A. An analysis of the flora of the Gull Run '\<[ountail\ regi01l of Virginia 11sing Rallt1kiaer's "life-form" methorl. ]OUrIl. \

23 McDoNALD, E. S. The life-forms of the flowering plants of Indiana. Amer. MidI. Nat. 18 (5): 687-773. 1937. Moss, E. H. The vegetation of Alberta IV. The poplar association and THE STEM 8Ml related vegetation of Central Alberta. Jour. Ecol. 20: 380-415. 1932. RAUNKIAER, C. The Life Forms of Plants a,"d Pia,,! Gcoyraph)". 632 pp. IN NORTH A: Oxford Press. 1934 (This book contains translated papers of Raunkiaer since 1903 on statistical plant geography, and is the source of all reference to Rauukiaer in this paper.) \VrrHRow, ALICE P. Life forms and leaf size classes of certain plant communities of the Cincinnati region. Ecology 8(1): 12-35. 1932. an Over much of th Range," comprising s' O~yzopsis are promin These are commonly as 15-20 per cent of t as 90 per cent infecti~ grasses as component manifest in the nattl seemed desirable to r soon became apparel stem smut on Stipa a as occurring in l\ortt than 80 years stem 51 and curated in this (Schlecht.) Fr., wh one variety, in two g' phologic aspects of I the name U. hypod)!1 already have been pi treatise on the knowl in North America. '

l Cooperati\,e im'es of Forage Crops and D Engineering, Agricultl a [ Agriculture, ill COOl meut Station, Pullmal rector as Scientific P; '" A contribution Department of Rutlel • Pathologist, Di, Industry, Soils, and