Vegetation Structure and Environmental Conditions of Forest Edges in Panama Author(s): Guadalupe Williams-Linera Source: Journal of Ecology, Vol.

78, No. 2 (Jun., 1990), pp. 356-373 Published by: British Ecological Society Stable URL: http://www.jstor.org/stable/2261117 . Accessed: 27/03/2011 12:20
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Journal Ecology(1990), 78, 356-373 of

VEGETATION STRUCTURE AND ENVIRONMENTAL CONDITIONS OF FOREST EDGES IN PANAMA

GUADALUPE WILLIAMS-LINERA* Department Botany,University Florida,Gainesville, of Florida32611, U.S.A. of SUMMARY
structure and treemortality five in (1) Micro-environmental conditions, vegetation to forest edgestenmonths twelve wet yearsold werestudiedin thetropicalpremontane forest Panama. of (2) Along transects from to of clearings theinterior theforest, greatest the changein and occurredbetween2 5 and 15 m into the forest. temperature relative humidity The forest border.At themostrecently canopywas mostopen at theclearing-forest cleared this farther theforest into site, opencanopyextended where took edgethanat sites clearing place earlier. at and basal area oftrees < 10cmdiameter 1 3 m high)weretwiceas great (3) Density ( of at at theforest edgescomparedwiththeinterior forests thesiteswheretheboundaries werecreatedfiveto twelveyearspreviously. was (4) Floristic composition unchanged from forest the boundaries to along transects theinterior forests light-demanding of and specieswerenotmoreabundantat forest edges interior. comparedwiththeforest (5) The edge:interior ratioof treesthatdied after edgeswerecreatedwas 14: 1. the (6) Beyond 15-25 m into the forest, neither environmental conditions nor the forest structure tree and wereinfluenced proximity theforest to mortality Between by boundary. 0 and 15 m, however, structure vegetation changedwithboth distancefromthe forest and timeelapsedsinceclearing. This study the boundary indicates ecologicalsignificance of edge vegetation a buffer as protecting forest from in vegetation conditions adjacent clearings.

INTRODUCTION Whenforests cutand abrupttransitions thevegetation are of result between clearings and the remaining the conditionsat the boundarychangemarkedly forest, environmental (Lee 1978; Lal 1987; Kapos 1989). Canopy removalincreasesthe amountsof solar radiationand rainfall thatreachthesoil surface. The highrelative humidity commonly observedin a forest interior decreasesin clearings because of increased temperature air and windvelocity, decreased and ratesoftranspiration (Schulz 1960;Grubb& Whitmore 1966;Raynor1969;Lawson,Armstrong-Mensah Hall 1970;Lee 1978;Lawson,Lal & & Oduro-Afriyie 1981; Fritschen1985; Lal 1987). Environmental conditionsat forest boundariesare intermediate betweenthoseprevalent clearings in and forest interiors. However,thereis littleinformation the concerning distanceinto the interior tropical forest whichtheseboundary at conditions prevail. structure floristic Vegetation and at composition theedge,and somedistance intothe interior theforest, of reflect changesinabioticconditions (Wales 1972;Ranney, Bruner & Levenson1981;Lovejoyetal. 1983;Miller& Lin 1985).For example, structural a analysis oftemperate deciduousforest edgesshowedthatbasal area and stemdensity trees of and decreasefrom edgetoward interior a distance about 15m (Ranney, the the saplings for of Bruner Levenson 1981). In an Acerrubrum maple) forest & (red edge,leafdensity was twiceas large on the forest boundaryas thatin the interior (Miller & Lin 1985). In
* Present address:Instituto Ecologia, Apartado Postal 63, Xalapa, Veracruz91000,Mexico. de

356

G. WILLIAMs-LINERA

357

branches on often developlow and thick boundary forests, treesat theforest temperate (Wales boles thatlean towardtheclearing and thesideoftheclearing, also asymmetrical as have been described boundaries 1972;Ranney1977;Levenson1981).Tropicalforest by are interior. former characterized The thoseoftheforest unlike marginal communities, and vines, woodyclimbers heliophilous withmanysmalltrees, of densestrips vegetation of the five 1974).Within yearsfollowing creation a forest (Longman& Jenik ground herbs 10 in Amazoniareportedly developeda zone ofvinesbetween m and edge,forests central shaded the forest whichacted as a windbreak, 25 m wide and secondaryvegetation comparedwithconditions and increasedthehumidity, interior, decreasedthe sunlight edges the (Lovejoy et al. 1986).Although tropical-forest following clearing immediately with occurrence increased has and preservation, their are potentially important forest for structure theseedges are lacking. of deforestation, on vegetation data of edgesand of thatthespeciescomposition thevegetation forest It has beensuggested to of are edges are reported have more Temperate-forest theinteriors forests different. (Wales 1972; Ranney, individualsthan those of the forestinteriors shade-intolerant edges speciesinvadeforest Bruner& Levenson 1981). In centralAmazonia, secondary of among the trunks dead and fallentrees vegetation and createa band of secondary (Lovejoy et al. 1984). in of the forest mayaffect speciescomposition vegetation to Proximity theremaining may initial invasionof shrubs intotheedgeofa prairie overtime.For instance, clearings for a subsequent successfulinvasion of hardwood forestspecies be a prerequisite that fruits frugivores and (Petranka McPherson1979).Plantspeciesthatproducefleshy & forests (Thompson& dispersetheirseeds are abundantalong the edges of temperate of donnell-smithii (Apocynaceae),thefruits which Willson1978).Treesof Stemmademia at are dispersed birds,producemorefruits theedge of tropicalpasturesthanin the by & probably interior adjacentto forest forest (McDiarmid,Ricklefs Foster1977).Pastures farther from forest the of (Augspurger morepropagules treespeciesthanpastures receive to activities necessary maintainpasture are & Franson 1988). Continuedagricultural close to forest boundaries. treesat a newly the of forest Whenisolatedfrom protection neighbouring vegetation, interior of createdboundaryhave higherprobabilities dyingthan those in the forest conditions (Brokaw 1985; because of theirexposureto wind and otherenvironmental Lovejoy et al. 1986;Hubbell & Foster 1988; Lawton & Putz 1988). In thenorth-eastern of edges uprootmore UnitedStatesforests Abies balsamea,treesclose to theclear-cut In of the increasedmortality treesoccursat a readilythantreeswithin forest. general, clearing (Gordon 1973;Sprugel& Bormann1981). creatededge following newly to the of boundaries createdforest This investigation designed study effects newly was in conditionsand the structure tropicalvegetation Panama. The of on environmental tree mortalityand environmental between vegetationstructure, interrelationships at the of wereexamined theforest edges.In addition, importance cattlegrazing gradients in of and vegetation and agricultural composition practices themaintenance thefloristic at structure theforest edge was investigated. METHODS Studyarea of The study area is locatedneartheborderof theProvince Panama and theSan Blas Comarca, Republic of Panama, along the El Llano-Cartiroad (901 5'N and 901 8'N,

358

edgesin Panama Tropical forest

zone (Tosi 1971).The Wet Forestlife 78?55'Wand 78?58'W)in theTropicalPremontane is altituderangesfrom220 m to 325 m a.s.l. Averagetemperature 26 ?C but diurnal variationis greaterthan annual variation. Mean annual minimumand maximum Annualaveragerelative is are humidity 86%. temperatures 21 ?C and 29 ?C, respectively. is Mean annual rainfall about 3500 mm;thedrierseason (< 100 mmmonth-1)extends to fromJanuary March. Soils are veryacid (pH <4-6) and clayey,and containlarge of Al and low concentrations exchangeof quantities exchangeable and organicmatter able P, K, Ca and Mg. primaryforeston the Pacific side of the Five clearingsadjacent to undisturbed in sitesare presented for Details of thestudy dividewereselected thisstudy. continental The local inhabitants. age oftheedge is by Table 1. Siteage was determined questioning was similar all the in history was cut. The disturbance thetimeelapsed sincetheforest of hectares forest of begins withthecutting several pastures sites.The processofcreating With dries. after vegetation the a the thedry seasonand burning area within month during the first rains (April and May), rice or corn is usuallyplanted for one or two years During the finalcrop, seeds of the exotic grass 'ratana' dependingon soil fertility. are indicum (Houtt.) Merrill sown,and thepasturethatdevelopsis grazedby Ischaemum at wereonlypresent theseven-yearand between forest thepasture the Wirefences cattle. wheretrailsstartor wherewood has been old site. Cattle enterthe edge of the forest Therewas no edgeoftheforest. Cattledid notpenetrate beyondtheimmediate extracted. at evidenceof fire the penetrating edges of theforest studysites. gradients environmental Pasture-forest and weremeasured alongone humidity Maximumand minimum temperatures relative 25 from m into was sites.The transect 70 m long,extending transect each of thestudy at at were for Sun thepastureto 45 m intotheforest. shelters thethermometers constructed wereplaced at 10-m shelters Eightweather usingpalmfronds. 150cm above theground, pasture-forest and intervals alongeach transect, one morewas placed on each immediate were border.Temperature relative and humidity (measuredusinga slingpsychrometer) timesa week fortwo weeksat each siteduringthedryseason recorded noon) three (at this weremadeone siteat a time; explainssomeof 1987).Measurements (January-March in and humidity among sites. thedifferences absolutevalues of temperature relative

sites the of and TABLE 1. Characteristics disturbance history thestudy near border
of theProvinceof Panama and theSan Blas Comarca, Republicof Panama. Cleared area Edgeage AltitudeSlope of Disturbance clearing history adjacent (m) (?) Aspect (ha) (years) out the was when study carried rice field 5 5-15 NW 330 08 15 for years convertedpasture, cattle, then to 20 rice corn two and 315 10-30 NE 5 1 horse 2 to 8 then converted pasture, cattle, horses for 10 rice a year 5-15 N 7 325 20-30 cattle, to 10 riceforone yearthenconverted pasture, 340 10-25 SW *10 of 4 horses most theyear *12 20 340 10-30 NW * Ten-year-old twelve-year-old were sites and separated a deepforested by adjacent alongthesameridge, gorge.

G.

WILLIAMS-LINERA

359

was of Overstorey density trees estimated hemispherical by canopyphotographs (taken lens witha fish-eye placed 60 cm above thegroundin February1987).The photographs weretakenat 0, 5, 15 and 25 m along two transects fromthepasture-forest extending intotheforest each study border at site.The negatives wereanalysedforcanopyopenness of (percentage thehemispherical imageunobscured vegetation) by usinga computerized canopyphotograph analysisprogram (Becker,Erhart& Smith1986). Forestvegetation structure At each studysite,vegetation was surveyed along fourrandomly located 10-m-wide stripsfromthe pasture-forest borderto 20 m into the forest. The belt transects were dividedinto5-mx 10-mplotsparallelto theedge of theforest. each plotnumber In and diameter breast-height at (dbh:diameter 1 3 m) ofall trees lianas( > 5 cmdbh)were at and In recorded. a 5-mx 5-msubplotrandomly selected each of the5-mx 10-mplots,the in number and dbh of all woodyplants(< 5 cm dbh but > 2 m tall)wererecorded. Woody and herbaceousplants < 2 m tall wererecorded lifeform a percentage by as coverin a randomly located 2-mx 2-m subplotin each of the 5-mx 10-m plots. All plantswere identified. Forest vegetation heightadjacent to pastureswas measuredwith a Haga inclinometer. To assess the depth of penetration edge effects of into the forestvegetation, the transects wereextended 60 m intotheforest to interior. Plotsof 5 m x 5 m, as described above forplants < 5 cm dbh,wereplaced at intervals 20 m alongtwotransects. of Pointquarter samplesof trees (Cottam& Curtis1956)weretakenat 20-mintervals alongeach 20-mtransect. dbhand distance nearest The to neighbouring wererecorded thesize tree in classes:trees5-9 9 cm dbh, 10-19 9 cm dbh and ,20 cm dbh. Voucherspecimens were depositedat theHerbarium theUniversity Panama. of of Effect agricultural of practices thepasturevegetation theboundary on at Permanent plots were establishedto assess the effect cattlegrazingand pasture of maintenance near-edge on in vegetation thepasture.Adjacentto edgescreated5, 7, 10 and 12 yearspreviously, seriesof 2-mx 2-mplotsweresetup from edge to 20 m two the intothepasture. excludecattle To and agricultural activities barbed-wire a fence built was aroundone series plotsat each site,and theother of was used as a control. The fence was located 1 m outsidetheplots. Percentage cover,averageplant height lifeformand by floristic compositionwere recordedbeforeconstruction the exclosures(Augustof September 1986) and again in February, Juneand November1987. of on Effect edgeproximity dead treedensity To assess theeffect proximity theforest of to boundaryon treemortality, fallen, all or dead treesin a 50-mx 60-mplot at each of thefivestudysiteswere broken, standing counted.The shortaxis of the plot was the forest-pasture boundary.Each plot was dividedintostrips m wideby 50 m long.In each strip number dead trees 5 the of and the apparentmode of death (broken,uprooted,or died standing), diameter class (5-9 9, 10-19 9, 20-29 9 and > 30 cm dbh), and decay stateof each wererecorded. The effects edges on treedeathcannot,however, assessed usingall dead trees, of be becausesomeundoubtedly priorto thecreation theedge.Therefore, died of decayclasses were used to estimatewhen the treeshad died. The decay classes used in this study Maser & Trappe 1984) wereas follows: (adapted from

360 Class 1 Class 2 Class 3 Class 4 Class 5

Tropical forest edgesin Panama intactand undecayedtrees, smalltwigsand evenleaves present; mostly bark intact, branches present; but piecesof bark startto sloughoff wood is stillhard; bark generally absent,wood starts rot; to bole easilybreaksdown intosmallpiecesof rotten wood whenit is struck.

Classes 1 and 2 wereassumedto includetrees thatdied after creation theedges; of the classes4 and 5 includetrees thatwerepresumed havediedpriorto clearing, to regardless of thetimethathad elapsed.Class 3 was used as an intermediate class fortreesthathad in probably diedbefore of felling theyoungest clearings after creation theedgesin the the oldestclearings. Data analyses To estimate distances whichtheedgeeffect be detected boththeclearings in at can and the forest, data of maximumtemperature in were fitted a post-hocpiecewiselinear regression model: y = bo + b1D; (yo < D < Y1).
Co; (y > Yi) (ao; (Y < Yo)

(1)

To estimate distanceintotheforest to whichedge effects manifest, the are up canopy basal area and density woodyplants( < 5 cm dbh), and dead treedensity of openness, in werefitted a post-hoc model: piecewiseregression y = bo + b1D; (y < y1). (2) that: Both modelsare subjectto theconstraints
at y = yo, ao = bo + blDo, and at y = yl, co = bo + b1D1.

In model 1,D is distance alongthepasture-forest gradient, is theestimated distance yo into thepastureat whichmaximum temperature startsto decrease,y, is theestimated distance intotheforest where variableis notaffected theproximity theedge,ao is the of by thepredicted at temperature yo,and cois thepredicted at temperature y . In model2, D is the distancealong the forest borderand the interior the forest, is the estimated of y, distance intotheforest to which are and up edgeeffects manifest, bois thepredicted value ofthevariable(canopyopenness, basal area,density, dead tree or at density) theclearingforestborder. Piecewiselinear models were fitted with SAS (SAS 1984) and SYSTAT (Wilkinson1986). ANOVAANDt-tests wereperformed SYSTAT with (Wilkinson 1986). All proportions were before to arcsine-square-root-transformed beingsubjected parametric tests. Means were comparedusingthe Waller-Duncan Bayes LSD (Petersen1985). Variationabout the mean is reported mean+ one standarderror, as unlessotherwise specified. RESULTS environmental Pasture-forest gradients In the pastures25 m fromthe forestborderthe average maximumand minimum temperatures duringthe dry season of 1987 were 1 5-3 1 ?C and 0-10 ?C higher,

G. WILLIAMS-LINERA of on distance, estimated regression by model, to TABLE 2. Theeffect edge-age the maximum temperaturethe in to boundary affects which proximity theforest the maximum temperature at clearing and in theforest ao, (yo) (yi), and predicted border distance and coat distance The clearing-forest is at 0 m. Valuesin yo yl. parentheses standard are errors.
Edge age (years) 08 5 7 10 12 Forest Clearing Maximum Maximum Distance Yo temperature DistanceYI temperature ao cO (m) (OC) (m) (OC) -94 (304) -11.0(6 11) - 105 (384) -07 (022) -90 (1 73) 279 299 264 276 28 1 69 (271) 25(375) 150 (394) 65 (088) 67 (1 64) 257 274 252 252 254

361

values of 15-45 m into the forest.Midday relative respectively, than corresponding than 15-45 m intotheforest. humidity 7% loweron theaveragein theclearing was Maximumtemperature therelative and changed alongpasturehumidity progressively in interior transects all sites.Because relative forest humidity relatedto temperature, is linear modelto estimate of were temperature usedina piecewise measurements maximum of conditions wereaffected theproximity theedge. by thedistance which at environmental in humidity) temperature (and consequently relative The greatest changein maximum 2 and between 5 m and the 0 occurred between 7 m and 10 5 m from edgeintothepasture changesin (Table 2). Beyond 15 m into the forest, 15 m fromtheedge into the forest of were irrespective thetimethathas elapsedsince maximum temperature notsignificant, clearing. of the into Alongtransects theinterior theforest canopywas mostopenat theclearingsite indicatedthattheten-month-old forest border(Fig. 1). Piecewiselinearregression at of of had a higher percentage canopyopennessand thattheeffects conditions theedge where tookplaceearlier thanat sites farther theforest into wereevident clearing (Table 3).

which proximitytheforest the to into interior, and boundary detected theforest is values the at forest of basal of border canopy openness, areaanddensity predicted x and of (5-m 50-m strips parallel to woody plants 5 cmdbh), density deadtree (< in are errors. edge).Values parentheses standard
Distance (m) intoforest (yi) and predicted value ao yi ao Yi ao y' ao

by model, to of distance estimated regression TABLE 3. Theeffect edgeageon the

Condition Canopy (%) Basal area (m2ha-') Density(stemsha-') Dead trees(number 0-025ha- )

08 96 (0 86) 430(1 93) 0.0 (1 06) 20(021) 1-4(1-66) 3515 (828) 13.0(1-00) 17-0(1-21)

Edge age (years) 7

10

12 6 0 (0 39) 104(038) 9 8 (1-31) 55(050) 12 7 (0-97) 9692 (783) 9 4 (1-41) 15 5 (2 12)

5 4 (0 15) 5 3 (0 42) 31 8(066) 153(086) 14 4 (2 15) 7 5 (2 34) 51(044) 32(075) 6-1(1 20) 6-7(2 80) 7770(906) 6062(860) 7-6(1-77) 6.7 (0-60) 10.5(2-63) 10-9(2 17)

4 9 (0 53) 128(1 72) 7.4 (0 78) 61(054) 12 9 (3-5) 8092 (1251) 6-8(0-93) 13 8 (2-72)

362

Tropicalforest edgesin Panama

40
[

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20 _ 40: (b)

a1)

40 20.

(c )

a)

-0
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40-

(d)

40. 40:
20

(d,

(e))

5 15 25 0 Distance into the forest(m)

at of FIG. 1. Changesin thepercentage canopyopennesswithdistanceintotheforest thesitesin and (d) Panama whereboundaries were(a) tenmonths, five (b) years, sevenyears, tenyears, (c) one (e) twelve yearsold. Errorbars represent standarderror.

structure Forestvegetation with distance from the Densityand basal area ofwoodyplants(< 5 cmdbh) decreased borderinto the forest edge of themost pasture-forest interior, exceptalong theforest the sitesolder clearedarea (Fig. 2). Between m and 20 m from edgeoftheforest 0 recently valuesfordensity basal area of trees(< 5 cm and thanfive yearsold showedthehighest were estimatedto extend6-14 m into the forestinterior dbh) (Fig. 2). Edge effects and valuesfordensity class (Table 3). Treesin the5-9-9-cm-diameter also showedhigher basal area up to 20 m fromthe pasture-forest boundarythan beyond20 m into the ten werecreated at where boundaries the of seven, and twelve years interior theforest sites in diameter classes(10-19-9and > 20 and The previously. density basal area oftrees other cm dbh), however, not changewithdistanceinto theforest did (Table 4). Stem density woody plants(< 5 cm dbh and > 2 m tall) was 49-55% less in the of of and interior theforests thanat theedges,exceptat siteswhich wereclearedtenmonths sevenyearspreviously present when (Table 4). At themostrecently clearedsite,saplings thefield created was in recruitment seeds from grew height, apparently and there little was or seedlings. in Based on differences stem density basal area forwoodyplants(< 5 cmdbh),and and relative and edge' was defined as temperature, humidity openness thecanopy,a 'forest of a 20-m-wide of were strip forest measured from trunk thetrees> 10cm dbhwhich the of closestto thepasture.Data from20-60 m fromthe forest to edge wereconsidered be of representative theforest interior.

(a)

(a)

sooflfpL
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01111l

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iSlul (c)

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(e) 5000:

(e)

05

10 15 2040 60510

Distance into the forest (mi)

150 i5204060

in FIG.2. Changesindensity basal area with and distance intotheforest Panama for woodyplants the were(a) tenmonths, five (< 5 cmdbh) at thesiteswhere boundaries (b) (c) years, sevenyears, one (d) tenyearsand (e) twelveyearsold. Errorbars represent standarderror. TABLE4. The effect the time that has elapsed since clearing on stem density and of basal area at the forest edge and within the forest. Values in parentheses are standard errors. Values at the forestedge are averages of sixteen plots for the tenmonth and five,ten and twelve-year-oldboundaries and thirteen plots forthe sevenyear-old site. Forest interiorvalues are averages of six point-quarter samples (for trees > 5 cm dbh). dbh category Edge age (cm) (years) <5 08 5 7 10 12 08 5 7 10 12 08 5 7 10 12 08 5 7 10 12 Stemdensity (stemsha-') Edge Interior 0-20 m 20-60 m Edge: interior 3621 (387) 6030(439) 5186 (482) 6126 (564) 5553 (692) 709 (94) 811 (121) 959 (100) 1036(122) 1086(167) 438 (52) 370 (74) 478 (68) 443 (82) 681(118) 250(51) 312 (58) 239 (63) 341 (56) 285 (53) 3343(662) 3455(407) 4157 (169) 3044 (481) 2749 (423) 636 (126) 782(102) 617 (105) 676 (38) 649 (54) 356 (24) 376 (47) 289 (35) 506(71) 497(61) 211 (19) 243 (22) 202 (23) 300 (37) 317 (29) 1 08 1 75 1-25 2 01 2 02 1 11 1-04 1 55 1 53 1 67 1 23 0 98 1 65 0-88 1-37 1 18 1 28 1 18 1-14 0-90 Edge 0-20 m Basal area (m2 ha- 1) Interior 20-60 m Edge:interior 1 60 (0 35) 1 86 (0-21) 1 88 (0 27) 1 38 (0 35) 1 47 (0 06) 2 97 (0-64) 3 32(047) 2 73 (0 46) 2 85 (0 17) 2 96 (0 24) 6-05(0 56) 6 21 (0 91) 4 69 (0 54) 8 70(1 24) 796(092) 23-45(290) 36-25(6-81) 26-54(4.89) 27 35 (4 56) 50 88 (9 68) 1-44 1 85 1 30 2 57 1 88 1-00 098 1 48 1.51 1 48 1 04 0-82 1-47 0 74 1 16 1 07 0.99 1 13 1-12 0-63

5-9 9

10-19-9

2 30 (0-21) 3 45 (0 36) 2 44 (0 31) 3 55 (0 39) 2 77 (0-39) 2 96 (0-42) 3-24(046) 4 04 (0 44) 4 31 (0-55) 4 39 (0-73) 6 30 (0 75) 5-09(1 05) 6 89 (1-07) 646(1 15) 920(1 51) 25.03(7.77) 35-74(8 77) 2989(8-92) 30-70(6-93) 31 82 (7 97)

)20

364

Tropicalforest edgesin Panama

Within forest, the averagedensity trees 10cmdbh)was 659 stems of (> ha-1, basal area was between30 m and 35 m, withemergent averaged35-9m2ha-1, and canopyheight a trees 45-50 m tall.Woodyplants(< 5 cmdbh) formed very conspicuous layer5-8 m tall Dead branches, along all forest edges,exceptat siteofthemostrecent clearing. however, the wereobservedon theside of tall treesfacing pastureat theforest edge. Understorey cover percentage Coveragebyplants(< 2 m height) was sparse;no significant differences detected were in of relation distance intotheforest to interior, irrespective theage oftheclearing (one-way cover was 15 8 (n = 97 plots; S.D. = 5.17). ANOVA, P > 0-05). The average percentage Coverbylife-form treeseedlings 6%, palms4 4%, Zingiberales Araceae3 2%, was: 4 and vines0 9%, shrubs 5%, grasses0-5%,liana seedlings 2% and 0 0 ferns 1[4%,herbaceous herbs0 1 %. Life-forms Dicotyledonoustreesrepresented 75-88% of the woody plants (<5 cm dbh) and decreasedin density withdistancefromtheforest edge. Palms represented 4-5%, and lianas2 8-13 0% ofall individuals 5 cmdbh),respectively. shrubs (< Few werepresent in forest interiors (Fig. 3). The tree: 9: palmratioofapproximately 1 was observed woodyplants(> 5 cmdbh) for both in forest interiors and along forest edges. Lianas (>5 cm dbh), however, were recorded onlyat theboundariesof forests (Fig. 3). Floristic composition A total of fifty-six families was represented woody plantsgreaterthan 2 m tall by was similarin the forest (Table 5). Species composition interiors and the forest edges; light-demanding speciessuchas Cecropiaspp. (fourteen individuals of2738), Vismia out were Kunth(fourindividuals), Palicoureaspp. (thirty-eight macrophylla or individuals) not abundantalong forest In edges and in the forest interior. the forest interior these to specieswererestricted gaps encountered along transects. on at of practices thepasturevegetation theboundary Effect agricultural Pasturevegetation near forest boundariesfive,seven,ten and twelveyearsold was combined intothree herbaceous groups:plantedIschaemum indicum, plants,and woody plants.The mostcommonherbswereferns, Lycopodium spp.,sedges,grassesotherthan and I. indicum vinesbelonging thefamilies to Passifloraceae, Rubiaceae,Smilacaceaeand Sapindaceae. The mostcommonwoodyplantswereshrubsof thefamily Melastomataceae, Solanumspp., Vismiamacrophylla, Cecropiaspp. and Piperspp. Withinsix to tenmonths, percentage the coverand height thepasturevegetation of as in groups, wellas thespeciescomposition, changed theexclosures from which cattle the wereexcluded, unlike controls the the plotswhich cattlegrazed(Fig. 4). Percentage cover ofwoodyplantsincreased and I. indicum in decreasedsignificantlythefenced plotsin the pastureadjacentto theten-year-old edges.All vegetation groupsin pastures adjacentto the seven-year-old edge and ten-year-old edge, as well as herbsand ratana in pasture in adjacentto thetwelve-year-old weretaller thefenced edge, plotsthanin thenon-fenced P plots(t-tests, < 0 02). The five-year-old pasture edgedid notdiffer significantlycover in orinheight between In treatments. all pastures, grazedplotssupported fewer speciesthan non-grazed plots.

G.
90 *

WILLIAMS-LINERA >5 cm dbh

(a)

365
(a)

<5 cmdbh 60 -

30 where thebounarieswee(a)temonth 601 3090

0 0

(b) fi

(b)

) twelve

(c)

g- 600E

30 60-

90 60-

(e)

30 TrePalm Liana Shrub Tree Palm Liana

FIG. 3. Changesinlife-form of and >5cm dbh (right) proportions woodyplants <i5cm dbh(left) at (w) theforest interior edge (0-20 i), and (0) in theforest (20-60 i), at thesitesin Panama wherethe boundarieswere (a) ten months,(b) fiveyears,(c) seven years,(d) ten yearsand (e) twelve yearsold.

on of Effect edgeproximity dead-tree density The density dead treesdecreasedwithdistanceintotheforest. of Dead trees(> 30 cm The ratio dbh) were 2-6 timesmore abundant at the forest edges than in the forest. decreasedforsmaller-diameter the of categories (Fig. 5). In theforest, averagedensity dead trees > 5 cmdbh) was 104trees 1. Treesin theforest hadiedmainly snapping of ( by trunks (55%), byuprooting (33%), orwhilestanding (12%). Most ofthedead individuals weredicotyledonous trees(84%); palms represented 16% of all deaths. dead treeswereseparatedinto the fivedecay classes listedearlier. Edge and interior Whendead-tree was examined relation decayclassesthere in to density weredifferences betweenforest interior. edge and theforest When thefirst two classes of least decayed trees werecompared, averageedge:interior the ratioforall siteswas 14:1 fortrees greater than5 cm dbh.Whenthelasttwoclassesofmostdecayedtrees wereused,thenumber of

366

Tropical forestedgesin Panama

TABLE5. Familiesand speciesofwoodyplants(> 2 m tall)inthestudy area nearthe

of border theProvince Panama and theSan Blas Comarca,RepublicofPanama. of Nomenclature followsCroat (1978). Family Palmae Species
Socratea durissima Wendl. H. IriarteagiganteaH. Wendl. Calyptrogyne spp. Palicoureaspp. Faramea spp. Pentagonia spp. Cephaelisspp. Psychotria spp. Brosimum sp. Sorocea spp. Cecropiaspp. Perebeaguianensis Aubl. Tovomita spp. Rheediaspp. L. Symphonia globulifera F. Bonafousia undulata (Vahl.) Markgr. Aspidosperma sp. Stemmadenia sp. Inga spp. Macrolobium sp. Pterocarpus sp. Bauhiniaspp. Manilkarazapota (L.) Van Royen Pouteriaspp. Miconia chrysophylla Urb. Miconia trinervia Don D. Triana Miconiasimplex Miconia spp. (14 species) Clidemiaspp. Xylopiabocatorena Schery. Annonaspp. Dialyanthera otobaWarb. Virolaspp. Protium spp. Tetragastris sp. Mabea occidentalis Benth. Richeriaspp. Eschweilera R. pittieri Knuth Gustaviadubia(Kunth) Berg Gustaviasp.

Number of individuals Percentage 196

54 24 21 179 42 20 16 14 161 54 13 14 8 145 71 18 7 122 89 16 5 116 53 15 12 7 112 17 9 93 8 4 4 70 7 83 23 9 83 24 24 75 71 3 65 48 8 62 15 11 9

7 16

Rubiaceae

6 54

Moraceae

11

5 88

Guttiferae

5 30

Apocynaceae

4-46

Leguminosae

4-24

Sapotaceae Melastomataceae

4 09 3-40

Annonaceae Mystiricaceae Burseraceae Euphorbiaceae Lecythidaceae

3 03 3 03 2-74 2-37 2-26

G. WILLIAms-LINERA
TABLE 5 (continued).

367

Numberof individuals Percentage 62 2-26 Myrtaceae 22 Eugeniaspp. 11I Calyptranthes spp. 6 Myrciaspp. Lauraceae 59 2-15 Ocotea cernua(Nees) Mez 5 Flacourtaceae 43 1-57 25 RyaniaspeciosaVahl Pittier 8 platyptera Carpotroche 4 Casearia arborea(L. C. Rich.) Urban Otherfamilies 371 13-55 Non-identified non-collected or individuals 711 25-97 Family Species Total 2738

of all was ha which considered, edges 206trees -', average density deadtrees along forest
to wereestimated extend foundwithin forest. the is twicetheaveragedensity Edge effects site at farther theforest the 10-month-old (13 m) thanin theoldestedges(Table 3). into whenan edge is thatthere an increasein treemortality is These results suggest strongly created.
(a) (C)
60

at to was the dead trees theedgewith respect theinterior roughly same.The edge:interior ratios were 14:3, 11:2 and 9:0 for treesthat died snapped, uprooted and standing, and (decay classes 1 and 2). Decay classes are respectively, whichdied most recently and depend on bole probablybiased because woody decay rates are species specific whendecayclasseswerenot diameter (Lang & Knight1979;Harmon1982).Nevertheless,

501
(b) :05
50

I fl~~~~~
*

40
(d)
~~~~~~~~60-

10

15 0 6 Time since exclosure (months)

10

15

(~ )and Ischaemum FIG.4. Changeswith time coverand height woodyplants(El,0), herbs in of indicum (ratana grass) (U, U) in (a, c) fencedand (b, d) grazedplots in thepasturein Panama adjacentto theedge createdtenyearspreviously.

368

forest edgesin Panama Tropical

I0

(b)

0~

oen10
a)

VI
k

hE

H!SE;
(e)

10 | I0

Distance into the forest (m)

30

60.

the edgesat thesiteswhere parallelto theforest strips in of FIG. 5. Number dead trees 5-mx 50-m years years,(c) sevenyears,(d) tenyearsand (e) twelve (b) were(a) tenmonths, five boundaries are: dead trees5-9 9 cm dbh (E3); 10-19 9 cm dbh (E); 20-29 9 cm dbh old. Diametercategories (0); and > 30 cm dbh (U).

DISCUSSION conditions border,environmental the from pasture-forest Beyond20 m into theforest of not by wereapparently influenced proximity the of and thestructure thevegetation dependsupon theorganisms of edge,however, The definition a forest forest boundary. animalsfarther affect thatforest boundaries Thereis someevidence underconsideration. of studieson predation birdnestsshowed'an thanplants.For instance, into theforest (Wilcove,McLellan & that extendedas far as 200-500 m into the forest edge effect' that'the Gates & Gysel(1978) found 1988),although & Dobson 1986;Andren Angelstam Studiesofplants only45 m intotheforest. of on edgeeffect' predation birdnestsextended on indicatethatthe edge effects basal area, stem deciduousforests, in temperate-zone & (Ranney,Bruner extendonly 10-15 m beyondclearings and leafarea density density Levenson 1981; Miller& Lin 1985). distances into the forest, reach different on Edge effects the micro-environment of on depending the timethathas elapsed sinceclearingand subsequentdevelopment conditions at vegetation theboundary.At thesestudysites,a changein environmental and relativehumidity, occurredwithin2-5-15 m of the boundary.The temperature, wet beyond15-25m from to opennessofthecanopyweresimilar thoseoftropical forests the thatin tropical forests, suggested Kapos (1989), however, of theboundary theforest.

G.

WILLIAMs-LINERA

369

Micro40 in extended m intotheforest. on edgeeffect themicroclimate theunderstorey in Panama are concordantwith at along transects the sites differences environmental and that the relative are temperatures higher, reportsthat maximumand minimum comparedwith and forests and the bordersof clearings is humidity lowerin clearings (Schulz 1960;Grubb& Whitmore forests oftropical data corresponding fortheinteriors & 1966; Lawson, Armstrong-Mensah Hall 1970;Longman& Jenik1974;Thompson& 1981; Lovejoy et al. 1986;Kapos 1989). Pinker1975; Lawson, Lal & Oduro-Afriyie vegefromforestclearanceaffect conditionsresulting Alteredmicroclimatological thanfive greater boundariesof clearings For example, at tationstructure theboundary. of twice density woodyplants the the yearsold (except one thatis sevenyearsold) support developed at mostvegetation theedgeoftheforest Within years, five intheforest interior. These from conditions theclearing. of a wall of woody plantsthatinsulatedthe forest edgevaluesof edges,where deciduousforest temperate from to are results similar reports of stemdensity were about twicethose in the interior the forest(Ranney, Bruner& Levenson 1981). to composedof treesaplingsand palms.Contrary was The wall of vegetation mostly of proportion plantsalongedgesas in lianas (< 5 cm dbh) showeda similar expectation, wereonlyfoundalong edges of the interior. Lianas (> 5 cm dbh), however, theforest and plantsgrow are that Thiscontrasts with observations liana seedlings abundant forest. in of taller significantly alongtheedgesofgaps thaneither thecentre gaps or in theforest interior (Putz 1984). edge are probablyrelatedto major rates of treesat the forest Increasedmortality edgethe In conditions. the5-mband closestto theforest changesinmicro-environmental of edge thenumber treesthatdied and At of number dead treeswas greatest. theforest diedin in The thanthenumber theinterior. waysin whichtrees werestanding higher was weresimilarto those observedon Barro Colorado Island, Panama, the forest interior had proportions been uprootedor had wheremostdead treeshad snappedand smaller treesand palmswere died and werestillstanding (Putz & Milton1982).Dicotyledonous diameter forest edge.Treesin thelarger created to equallylikely die whenclose to a newly of edgesthanintheinterior alongtheforest diedin a greater proportion classes,however, died standing of and thefactthatmoretrees The number dead trees theforests. increased stress is of of at theforest edgethanin theinterior theforest a reflection theenvironmental of the (Lovejoy et isolatedfrom protection neighbouring vegetation by experienced trees al. 1986;Waring1987; Hubbell & Foster 1988; Lawton & Putz 1988). confera peculiar structure and increasedratesof treemortality Alteredvegetation vegetation on edges. Observedfromthe pastureside, forest-edge physiognomy forest and tall 5-8-m-tall woody vegetation layerof understorey consistedof a well-defined less Crownsofwoodyplants, wereeasilydistinguished. trees. Thesetwovegetation layers at of crowns talltrees theforest towardtheclearing. However, than8 m tallweregrowing side. weredead on theclearing and the away from clearing, branches edgeweregrowing zone whichgrow thosein thetemperate from differ apparently Dead crownsof talltrees createdforest edges (Gysel 1951; Trimble& Tryon 1966; toward artificially laterally of forests Abies Levenson 1981; Ranney,Bruner& Levenson 1981). In high-altitude & is deathofexposedtrees common(Sprugel1976;Sprugel Bormann balsamea,however, are Amazonia dead trees abundantalong forest edges(Lovejoy et 1981).Also, in central of (and highal. 1984). The prevalence dyingtreesalong the edges of tropicalforests at conditions theforest thatenvironmental boundary altitude suggests forests) temperate tropicalforest regions.Alternatively, are more severein the tropicsthanin temperate

370

Tropicalforest edgesin Panama

conditionswhen isolated from treesmay be less adaptable to the new environmental neighbouring vegetation. in is Development vegetation theedge of theforest faster tropical of at forests thanin five temperate forests. understorey The vegetation layerwas welldevelopedwithin years and basal areas of stemsweredouble thoseof the and stemdensities following clearing in it At forest interior. the edges of temperate deciduousforest, contrast, takes about densities leafdensities twice largeas thoseoftheforest and are as before stem twenty years & interior (Ranney,Bruner Levenson 1981; Miller& Lin 1985). at interior consisted of Species composition theforest edgesand in theforest mostly shade-tolerant species with few light-demanding species. Tropical forestspresent'a continuum regeneration of patterns' (Augspurger 1984;Brokaw 1987;Hubbell& Foster 1987). For convenience, treespeciesare classed into a fewregeneration groups.Two and secondarytree species,or else lightare common classifications eitherprimary demandingand shade-tolerant species (Whitmore1975). The shade-tolerant species or saplingsin theunderstorey untila canopy gap opens as suppressed persist seedlings and permits accelerated above or nearthem growth (Brokaw 1987).The light-demanding in be in These speciescan rarely foundgrowing speciesgerminate gaps and growrapidly. or theforest understorey, exceptperhapsunderdeciduoustrees close to gaps (Hubbell& Foster 1987). Althoughvegetationat the forestedge was expectedto be dominatedby lightshade-tolerant demanding plants, specieswereabundantat boththeforest edgeand inthe interior. forest wereabundantinthepastures intheadjacent but Light-demanding species foresttheywere mostlygrowingin light gaps. In temperate deciduous forests, by often the differs between forest comparison, speciescomposition markedly edgeand the interior theforest. of Forestedgesare morespecies-rich, shade-intolerant and speciesare abundant along forestboundaries(Wales 1972; Ranney,Bruner& Levenson 1981). of Likewise, abundantregrowth light-demanding speciesis observed alongtropical forest boundariesin central Amazonia (Lovejoy et al. 1984). The scarcity light-demanding of plants on forestboundaries in Panama indicatesthat besides the requirement for increased lightpenetration, secondary specieshave otherrequirements germination for and establishment (Williams-Linera 1990). Quality of light(Vazquez-Yanes & Smith 1982;Vazquez-Yanes & Orozco-Segovia1985)and soil disturbance (Wesson& Wareing the 1969; Bell 1970; Putz 1983) may influence presenceof pioneer species at forest In boundaries. addition, faster of forest the boundaries growth existing plantsin tropical of maycurtailtheestablishment light-demanding species. Whencattle wereexcluded from pastures adjacentto forest edges,thegeneral effect was coverand height a reduction pasturegrassand an increasein number, in of percentage woodyplants.Studiescomparing grazedand ungrazedvegetation usuallyindicatethat in is speciesdiversity higher moderately grazedareas (Harper 1977;Collins 1987).Other decreased as a functionof authors, however,have reportedthat species diversity increasing grazing cattle by (Waser& Price1981),other largemammals (Belsky1986)and small mammals(Dickinson& Kirkpatrick the 1986). In thisstudy, fencedplots had a number speciesthanunfenced of higher plots,and thepasture grasswas shadedbywoody six Growthof secondary specieswithin to tenmonths. specieswas so fastthateventhe shortperiod of cattleexclosureindicateda significant change. In the unfenced plots An of woody plantswerebrowsedor cut whenpasturesweremaintained. implication theseobservations thatabruptpasture-forest is boundariesare a resultof grazingand pasturemaintenance.

G.

WILLIAMs-LINERA

371

at boundarieschangeswithboth structure forest This studyshows thatvegetation and over time.When edges are created,micro-environmental distanceinto the forest are close to thepasturemargin modified. conditions theforest in

ACKNOWLEDGMENTS that and Thanks are due to JackPutz forsupport,encouragement usefulsuggestions J. and thismanuscript, P. Feinsinger, W. Ranney,N. V. L. Brokaw, greatly improved A on for referees valuablecomments themanuscript. J.R. Etherington anonymous and discussion. Many people helpedin Panama, butI to specialthanks JackEwel forhelpful me plantspecies indebted Gordon McPhersonforhelping to identify to am particularly canopy photographs.I analysis of hemispherical and Alan Smith for computerized from James the NoyesFoundation the received support gratefully acknowledge financial Tropical ResearchInstitute. through Smithsonian the

REFERENCES
in ratesas an edge effect habitatislands:experimental Andren, & Angelstam, (1988). Elevatedpredation H. P. evidence.Ecology,69, 544-547. of of treeseedlings: comparative and C. a Augspurger, K. (1984). Lightrequirements neotropical study growth survival. Journal Ecology,72, 777-795. of seeds into light-gaps forest and sitesin a Augspurger, K. & Franson, E. (1988). Inputof wind-dispersed C. S. Neotropicalforest. Journal TropicalEcology,4, 239-252. of D. A. Manual CanopyPhotoAnalysis Smithsonian Becker, Erhart, W. & Smith, P. (1986). Operator's P., System. Tropical ResearchInstitute, Balboa. and A Bell,C. R. (1970). Soil distribution germination experiment. TropicalRain Forest(Ed byH. T. Odum & R. F. Pigeon),pp. 177-188.U.S. AtomicEnergy Oak Ridge,TN. Commission, in of National Park, disturbances grasslandsof the Serengeti Belsky,A. J. (1986). Revegetation artificial Tanzania. Journal Ecology,74, 419-437. of in Brokaw,N. V. L. (1985). Gap-phase regeneration a tropicalforest. Ecology,66, 682-687. Brokaw,N. V. L. (1987). Gap-phase regeneration three of pioneertreespeciesin a tropicalforest. Journal of Ecology,75, 9-19. Collins, L. (1987). Interaction disturbances tallgrass S. of in prairie: field a experiment. Ecology, 1243-1250. 68, J. Cottam,G. & Curtis, T. (1956). The use of distancemeasuresin phytosociological sampling. Ecology,37, 451-460. CA. Croat,T. R. (1978). Flora of Barro ColoradoIsland.Stanford University Press,Stanford, K. J. in burnedand Dickinson, J. M. & Kirkpatrick, B. (1986). The impactof grazingpressure clearfelled, undisturbed eucalyptforest. Vegetatio, 133-136. 66, of L. fore, environmental Fritschen, J. (1985). Characterization boundaryconditions affecting phenomena. TheForest-Atmosphere Interaction byB. A. Hutchinson B. B. Hicks),pp. 3-23. D. ReidelPublishing (Ed & Company,Boston,MA. L. and successin field-forest ecotones.Ecology, Gates,J. E. & Gysel, W. (1978). Avian nestdispersion fledging 59, 871-883. windand other causes in mixedfir-red standsadjacentto clearcuttings. fir D. Gordon, T. (1973). Damage from ForestServiceResearchPaper,PSW-90. StatesDepartment Agriculture, United of P. T. of and lowlandrainforest Ecuador. II. The in Grubb, J. & Whitmore, C. (1966). A comparison montane climate on and itseffects thedistribution physiognomy theforests. and of Journal Ecology, 303-333. of 54, Gysel,L. W. (1951). Bordersand openingsof beech-maplewoodlands in southernMichigan. Journalof Forestry, 13-19. 49, Harmon,M. E. (1982). Decompositionof standingdead treesin the SouthernAppalachian Mountains. Oecologia,52, 214-215. Harper,J. L. (1977). Population Biologyof Plants.AcademicPress,London.

372

Tropicalforest edgesin Panama

Hubbell, P. & Foster, B. (1987). La estructura S. R. espacialen granescala de un bosque neotropical. Revistade 35 Biologia Tropical, (Suppl. 1), 7-22. of forest. PlantEcology(Ed Hubbell, P. & Foster, B. (1988). Canopy gaps and thedynamics a neotropical S. R. by M. J. Crawley), 77-96. BlackwellScientific pp. Publications, Oxford. of Kapos, V. (1989). Effects isolationon thewaterstatusof forest patchesin theBrazilianAmazon. Journal of TropicalEcology,5, 173-185. Lal, R. (1987). TropicalEcologyand PhysicalEdaphology. JohnWileyand Sons, New York. D. in Lang, G. E. & Knight, H. (1979). Decay ratesforboles of tropical trees Panama. Biotropica, 316-317. 11, K. moistsemi-deciduous forest Lawson,G. W., Armstrong-Mensah,0. & Hall, J.B. (1970). A catenaintropical near Kade, Ghana. Journal Ecology,58, 371-398. of Lawson,T. L., Lal, R. & Oduro-Afriyie, (1981). Rainfallredistribution microclimatic K. and changesover a clearedwatershed. Tropical Agricultural Hydrology byR. Lal & E. W. Rusell),pp. 141-151.John (Ed Wiley and Sons, New York. in R. and Lawton, 0. & Putz,F. E. (1988). Naturaldisturbance gap-phaseregeneration a wind-exposed tropical lowermontanerainforest. Ecology,69, 764-777. Columbia University Lee, R. (1987). ForestMicroclimatology. Press,New York. J. islandsinSoutheastern Wisconsin. Forest in Levenson, B. (1981). Woodlotsas biogeographic IslandDynamics New Man-Dominated Landscapes(Ed by R. L. Burgess& D. M. Sharpe), pp. 13-39. Springer-Verlag, York. Longman, A. & Jenik, (1974). TropicalForestand itsEnvironment. K. J. Longman,London. J. H. of Lovejoy,T. E., Bierregaard, O., Rankin, M. & Schubart, 0. (1983). Ecological dynamics tropical R. RainForest:EcologyandManagement byL. Sutton, C. Whitmore A. C. T. forest fragments. Tropical (Ed & Oxford. Chadwick),pp. 377-384. BlackwellScientific Publications, of A. Lovejoy,T. E., Bierregaard, O., Rylands, B. et al. (1986). Edge and othereffects isolationon Amazon R. Conservation forest fragments. Biology:Scienceof Diversity by M. E. Soule), pp. 257-285. Sinauer, (Ed MA. Sunderland, R. K. L. M. T. J. Lovejoy, E., Rankin, M., Bierregaard, O., Brown, S., Emmons, H. & Van derVoort, E. (1984). remnants. Extinctions byM. H. Nitecki), 295-325. University Ecosystem decayofAmazon forest (Ed pp. of Chicago Press,Chicago. donnell-smithii McDiarmid, R. W., Ricklefs,R. E. & Foster, M. S. (1977). Dispersal of Stemmademia (Apocynaceae)by birds.Biotropica, 9-25. 9, StatesDepartment Maser,C. & Trappe,J. M. (1984). The Seen and UnseenWorldoftheFallen Tree. United of PNW-164. GeneralTechnical Agriculture Report, Miller,D. R. & Lin, J. D. (1985). Canopy architecture a red maple edge standmeasuredby a pointdrop of method.TheForest-Atmosphere Interaction byB. A. Hutchinson B. B. Hicks),pp. 59-70. D. Keidel (Ed & Publishing Company,Boston,MA. Petersen, G. (1985). Designand Analysis Experiments. R. of Marcel Dekker,New York. Petranka, W. & McPherson, K. (1979). The role of Rhus copallinain thedynamics the forest-prairie J. J. of ecotonein NorthCentralOklahoma. Ecology,60, 956-965. of Putz,F. E. (1983). Treefall and mounds, pits buried seeds,and theimportance soildisturbance pioneer to trees on Barro Colorado Island,Panama. Ecology,64, 1069-1074. of Putz,F. E. (1984). The naturalhistory lianas on Barro Colorado Island,Panama. Ecology,65, 1713-1724. K. rateson BarroColorado Island. TheEcologyofa Tropical Forest Putz,F. E. & Milton, (1982). Treemortality Institution (Ed byE. G. Leigh,A. S. Rand & D. W. Windsor), 95-100. Smithsonian pp. Press, Washington, DC. No. 1069. Forest Island Edges-their Sciences Division Publication, Ranney,J. W. (1977). Environmental and to Structure, Development, Importance RegionalForestEcosystem Dynamics.Oak Ridge National Oak Ridge,TN. Laboratory, J. J. of of Ranney, W., Bruner, C. & Levenson, B. (1981). The importance edgein thestructure dynamics M. and forest islands. ForestIsland Dynamicsin Man-Dominated Landscapes (Ed by R. L. Burgess& D. M. New Sharpe),pp. 67-95. Springer-Verlag, York. in G. structure a coniferous forest and a contiguousfield.Forest Raynor, S. (1969). Wind and temperature Science,17, 351-363. SAS (1984). SAS/ETS User'sGuide,5thedn. SAS Institute, Cary,NC. in Suriname.Verhandelingen Koniklijke der Schulz,J. P. (1960). Ecological studieson rainforest Northern van Wetenschappen Nederlandse Akademie Natuurkdunde TweedeReeks,53, 1-267. Afdreling of in D. structure wave-generated Abiesbalsameaforests thenorth-eastern United Sprugel, G. (1976). Dynamics States.Journal Ecology,64, 889-910. of F. D. and balsam fir Sprugel, G. & Bormann, H. (1981). National disturbance thesteadystatein high-altitude forests. Science,211, 390-393. in 0. R. characteristics a tropicalevergreen Thompson, E. & Pinker, T. (1975). Wind and temperature profile in forest Thailand. Tellus,6, 562-573. of J. and fruits. Thompson, N. & Wilson,M. F. (1978). Disturbance thedispersal fleshy Science, 200, 1161-1163.

G.

WILLIAMS-LINERA

373

de Tosi, J. A., Jr(1971). Zonas de vidaen la Reptublica Panama. FAO, Rome. cut into E. G. Trimble, R. & Tryon, H. (1966). Crownencroachment openings inAppalachianhardwoodstands. 62, of Journal Forestry, 104-108. in controlof seed germination the tropicalrain forest H. Vazquez-Yanes,C. & Smith, (1982). Phytochrome 92, New Phytologist, and itsecologicalsignificance. and pioneertreesCecropiaobtusifolia Piperauritum 477-485. de del A. C. Vazquez-Yanes, & Orozco-Segovia, (1985). Posiblesefectos microclima los clarosde la selva,sobrela y Piper donnell-smithii Heliocarpus de germinaci6n tresespeciesde arbolespioneros:Cecropiaobtusifolia, Mexico II (Ed by A. G6mezsobrela Regeneracion Selvas Altas en Veracruz, de Investigaciones auritum. Pompa & S. Del Amo), pp. 241-253. INIREB. EditorialAlhambraMexicana, Mexico. and southern edges in a matureoak-hickoryforest. Wales, B. A. (1972). Vegetationanalysisof northern 42, EcologicalMonographs, 451-471. 37, to of R. Waring, H. (1987). Characteristics treespredisposed die. Bioscience, 569-574. of of Waser,N. M. & Price,M. V. (1981). Effects grazingon diversity annual plantsin the Sonoran Desert. Oecologia,50, 407-411. of of in occurring populations buried P. G. Wesson, & Wareing, (1969). The roleoflight germination naturally 20, Botany, 403-413. of weed seeds. Journal Experimental T. Whitmore, C. (1975). TropicalRain Forestsof theFar East. ClarendonPress,Oxford. zone. in Wilcove,D. S., McLellan, C. H. & Dobson, A. P. (1986). Habitat fragmentation the temperate MA. (Ed Conservation Biology:ScienceofDiversity by M. E. Soule), pp. 237-256. Sinauer,Sunderland, The System Statistics. L. SystatInc., Evanston,IL. for Wilkinson, (1986). SYSTAT. in of edge vegetation Panama. Biotropica Williams-Linera, (1990). Originand earlydevelopment forest G. (in press).

received January 1990) 10 (Received28 July1989; revision