PhotographybyROBERTLLEWELLYN/WrittenbyJOANMALOOFTHE

LIVINGFOREST

AVISUALJOURNEYINTOTHEHEARTOFTHEWOODS

ToJoanMaloofandherdedicationtopreservingourforests.R.L.

•

ToallsupportersoftheOldGrowthForestNetwork;theyenablemetospeakfortheforests.J.M.

TheOldGrowthForestNetworkisthefirstnationalorganizationworkingspecificallytopreserveancientforests

fortheenjoymentofpresentandfuturegenerations.Incountiescapableofsupportingforestgrowthweidentifyatleastoneforestthatwillbeforeverprotectedfromloggingandopentothepublic.Thenwehelpfamiliesconnectwiththeseforests.Theresultwillbeanationalnetworkoftreasuredforestswhere

allgenerationscanexperiencenativebiodiversityandthebeautyofnature.Wealsospeakoutforthreatenedancient

forestseverywhere.Tolearnmore,visitwww.oldgrowthforest.net.

Contents__________

PrefaceTOWARDTHESKYTheCanopyandBirdLife

EYESALLOVERLeavesADEEPBREATHLungsoftheLand

WORLDOFWATERCyclesandCreaturesONTHEGROUNDAnimalsoftheForestFloor

THEMIDDLEREALMTrunksSHEDDINGANDRESTINGAutumnandWinter

BUDDINGWinterintoSpringFLOWERSANDFRUITSSpringintoSummer

CONNECTIONSRootsandFungiTHEONLY-GREENMossandFern

THESPIRITOFTHEFORESTBibliography

AcknowledgmentsIndex

Preface__________

ITISOURINTENTIONthatthisbookbeareminderthatthelivingpartofourplanet,thebiosphere,isjustathinandlovelymembrane.Wehaveharmedmuchofthismembrane,butbecauseitisaliveithastheabilitytoheal.Herewecelebratethepartofthebiosphereknownasthetemperatedeciduousforest,andallthelifeitcontains.Itisourwishthatincreasedunderstandingwillprovidemotivationforallowingtheforesttocontinueunimpededonitswildtrajectory.Althoughthephotographsonthesepagesarestunninglybeautifulandwecanwitnesssuchviews,it’simpossibletoreallyseethewholeofaforest.Aforestisacomplexwebofenergyandmatterthatreachesfarbeyondtheconfinesofidentifiedacreages.Muchofwhataforestdoes,andis,isinvisibleandcanneverbecapturedbythecamera.Atothertimes,theenergyandthematteralignonthescaleofhumanperceptionandwesee:oh,ablossom,asalamander,amushroom,abear!Allamazing,allwonderstobestudiedforeverand,even

then,tobefeltmorethanfullyunderstood.Takeanyoneofthesephotographicimagesandspendsometimewithit,andyouwillunderstandwhatthepoetWilliamBlakemeantaboutseeing“theworldinagrainofsand.”Thesenodesofmattershowusthenever-endingflowofenergyrighthere,rightthismoment.Andtheshutterclicks.

Thepenisslower,butitcanincludeabitmoreoftheintersections.Itcantellofstrandpullingstrandintheweboflife.Itcanhintatthedimensionoftime.Andthen,finally,beyondthecameraandthepenistheenergyandmatterthatyou,reader,bringtotheequation.Forafteryoudouseyourselfintheimagesandthewords,youmustgoastepbeyondandaddthedimensionofexperience.Itisonlythenthatyouwilltrulystartseeingtheforest.Andtheforestwillseeyou,too.Thereareplentyofeyesinthesephotosand,asyouwillreadhere,eventheeyelesstreessenseyourlight.Buthowdoweorganizethisdiscussionofaplacewhereeverythingishappeningatonce?Wechosetostartwiththelargestbones,thetrees—thelivingstructuresthatdefineaforest—andtomovefromthecanopytotheground.Thereismuchlifeinthecanopy,butprimarilyitisasurfaceforcatchinglight—thesourceofenergythatpowerstheentireforest.Treesarecreatorsofrain.Andmanyforestcreaturesdependonthewaterheldinandreleasedbytheforest.Sowelookcloselyatthat.Aswemovethroughtheforest,andthroughthesepages,wewitnessanimals

thatarewhollydependentonthehabitatcreatedbythetoweringtreeseventhoughtheywillneverhavetheexperienceofbeinginthetopofone.Otheranimalsusethetrunksashighways,andtheirmovementsmayreachfromthegroundtothetreetops.Giventhesizeanduprightpostureofhumans,thesightoftreetrunksisthemostcommonwayweexperiencetheforest,too.Wemustalsoconsidertheinvisiblefourthdimensionoftimeinordertotrulyunderstandaforest.Threechapterscapturehowaforestchangesthroughouttheyear:fromfall’sspectacletowinter’ssilence,fromspring’semergencetosummer’sripeness.Next,andperhapsmostimportant,wetakeapeekattheforestpartsthatareintimatelyintertwinedwiththesoil.Thisisthe(mostly)undergroundlifeoftheforest—rootsandfungi.Whilewe’redownthere,wetakeacloselookattheprimitiveplants.Theywillnevermakebudsorflowersorfruit,buttheyarethegreenoneswealwaysassociatewiththeforest:mossandfern,playgroundofthefaeries.Andfinally,nodiscussionoftheforestiscompletewithoutacknowledginghowtheforestmakesusfeel.Fewwordsdojusticetothosefeelings,soLlewellyn’simagesstandaloneattheendtoconveyamostpreciousresourceprovidedbyforests.

Lookingstraightupintothecanopyprovidesanunusualviewofabroad-leavedforest.

TowardtheSkyTheCanopyandBirdLife

WEAREGOINGTOstepintotheforestnow.Weareleavingbehindthecarsandthecomputers,andmaybeeventhepeople.Wearegoingtotakethisjourneytosee,tofeel,anecosystemthathasbeenaroundforaverylongtime—onethatdoesn’tdependonhumans,althoughwedependonit.Thisisarealplace,anditmaynotbefarfromyourhome.It’saplacewhereboththelearningandthemysteriesgoonforever.Let’sstartbylookingupintothecanopy.Toreallyfeelaforestcan-opywemustusedifferentsenses,andoftenthemostusefuloneisthesenseofimagination.Youmightspreadaclothandlieonyourback.Bepatient.Justwait.Soonyou’llnoticetheswayingofthetopsofthetrees.Notethattheydon’tallswaywiththesamemotion.Giveitmoretime.Justwatch.Dothetallertreesswaymorethantheshorterones?Butthey’renotallmovingtogether,right?Themovementseemsmorelikeacircling,likeaneddy.Firstthispartoftheforest,andthenseehowthewavemovesthroughandendsupoverthere?Thisforestisverydifferentfromtheonewewalkthroughthatseemssolidandstill.Listentothehushingmusicthetreesmakeastheymove,amostrelaxingsound.AstheapostleJohnsaidintheBible,“Thewindblowswhereitpleases,andyouhearitssound,butyoudon’tknowwhereitcomesfromorwhereitisgoing.SoitiswitheveryonebornoftheSpirit”(John3:8).Andyourspiritmoves,too,asyoulietherebathedinthesusurrationsofthegreenheights.TheZenmonkShunryuSuzukiputitthisway:“Whenwehearthesoundofthepinetreesonawindyday,perhapsthewindisjustblowing,andthepinetreeisjuststandinginthewind.Thatisallthattheyaredoing.Butthepeoplewholistentothewindinthetreewillwriteapoem,orwillfeelsomethingunusual.Thatis,Ithink,thewayeverythingis.”Ifyouwanderacrossaprairie,mostofthelifeexistsbetweenfootandheadheight,soit’sallveryaccessible,veryvisible.Wanderingthroughaforest,bycontrast,presentsyouwithascenethatseemsattimesveryquietandemptyofcreatures.Lifeisabundantthere,butmuchofitexistsinthecanopy—thathighgreenlayerbetweenearthandsky.Binocularsorclimbingropes,orboth,cangiveusasenseofwhat’shappeningupthere,butwe’restillonlyvisitors.Althoughinourearliestyearswemayfeelaninstincttoclimb,we’reprimarilycreaturesofthegroundnow.Theeasiestlivestoimagine,becausechancesarewe’llhearsomeoftheirsongsalongwiththetreesongs,arethebirds.Theregoesonenow.Whatkind?Too

fasttotell,toodark,toohidden.Onlythosewithearstrainedforsuchthingscanputanametoit.Whenyouwalkthroughaforesttalkingorsinging,thebirdscanhearyou,too,andmaysingback.Andtheyseeyou,probablybeforeyouseethem.Ifit’saforestyouwalkinoften,theymayevenknowyouasyou.Winginginfromtherivernowcomesahugebirdevenchildrenknow.It’sthegreatblueheron,anditsraspycroakingcan’treallybecalledasong.We’reusedtoseeingthesebirdsnearthewater’sedge,butnowthisoneisflyingtowardthetreetopswhereitraisesitsyoungonanaerialraftoflooselywovenbranches.Andnowourimaginationsgetturnedupsidedownasweconsiderthecanopyfromaboveinsteadoffrombelow.Fromabove,theimpressionisoftexture—thehighsandlowsofvarioustreespecies,somerisingfarabovetheothers.Thebirdsknowthislandscapeasintimatelyasweknowthestreetswedriveeveryday.Theywouldmissthattreeifitweregone.

Thecanopyofapineforestgivesaverydifferentfeelingfromthatofabroad-leavedforestasonegazesupward.

Weknow,althoughwecan’talwaysseethem,thatavarietyofbirdsarewingingtheirwayfromonetreetothenextupthere.Manyofthesebirdsaremigrants

thathavelefttheirtropicalwinterhomesfortheabundanceofthenorthernforestduringourall-too-briefwarmseason.Theyreturntothesametreesyearafteryear,sometimesforfifteenyearsormore.Heretheyrestfromtheirjourneys,mate,buildanest,andfeedtheiryoungonesbeforereturningtotheirotherhomes.Otherbirdsarehereyear-round,andalthoughweseethemfrequentlyaroundourhouses,theyalsoneedtheforest.Ifwecouldcrawlthroughthetreetops,wewouldfindevidenceofthesevisitorsthathavethegiftofflight.

Thegreatblueheronhasasix-footwingspanyetweighslessthansixpounds.Itshollowbonesmakeitlightandallowforeasyliftoff.

Fromabird’s-eyeview,theforestcanopyappearstexturedwiththevaryingheightsofdistinctindividualtrees.

Althoughwethinkofheronsaswetlandbirds,theynestintheforestcanopy.Themalebringstwigstothewaitingfemale,whothenweaves

themintoasecuretreetopplatform.

Mostmigratingbirdsreturntothesamearea,andoftenthesametree,yearafteryear.ThebirdsthatreturntoSandyIsland,SouthCarolina,picturedhere,arefortunatethattheirforesthasbeenpreserved.

Raptors,likeeagles,hawks,andosprey,oftenpickthetallesttreeinan

areafortheirnest.Theymayaddtothesamenestyearafteryear.

Howdobirdsknowwhichmaterialstocollectandexactlyhowtoconstructanest?They’rebornknowing—justanotherofnature’s

mysteries.

Easternbluebirdsareverysocial,exceptduringthebreedingseason.Individualsmayliveformorethantenyears.

Butnoteverybirdbuildsanest;somechiselacavity,ortakeadvantageofahollowspaceinadeadordyingtree,orusethespacemadebyapreviouschiseler.Bluebirds,whoweareusedtoseeingnestincylindersorboxesweprovide,oncecommonlyfoundoldwoodpeckernestsorsmallcavitiestocallhome.Thatwasbeforevastdeforestation,competitionfromnonnativehousesparrowsandstarlings,andDDTchangedlifeforbluebirdstothepointwheretheyrelyonourhelp.Bluebirdsarereboundingthankstothisnestingassistance,butunfortunatelythatisnotsoforred-headedwoodpeckers.Onceverycommon,thesewoodpeckersaredeclininginnumberduetomanyofthesameadversitiesthathaveaffectedbluebirds.Whenweseebirdswayupinthecanopy,frequentlytheyarepickinginsectsoffthetrees.Inadditiontoprovidinggreatfoodfortheirbabybirds,it’sabighelptothetrees.Howelsecouldatreeriditselfofthethingstryingtoeatit?Well,actually,anumberofcreaturesinthecanopy,includingotherinsects,helpthetreeriditselfofthesmalleatersandsuckers.Foreveryplant-eatinginsectoutthere,there’sanotherinsectlookingtoeatit.

Maleandfemalered-headedwoodpeckerslookidentical,andbothofthemfeedthenestlings.Sometimesathirdbirdwilljoinindefendingthe

nestandfeedingtheyoung.

Northerncardinalshaveacrestoffeathersontheirheadthattheycanraiseorlowerdependingonmood:calmandquiet,down;agitatedand

aggressive,up.

Thenorthernflickerisamongthefewwoodpeckerscommonlyseenontheforestfloor;they’redownthereeatingtheirfavoritefood—ants.

Crowsareextremelyintelligent.Theyhavealanguagealltheirown,andtheycanrecognizeindividualhumans.

Thepolyphemusmothhaslargespotsthatmimiceyes,onewaytodeterpredators.Thismalewillsearchforunmatedfemalemothsby“smelling”

themwithhislarge,bushyantennae.

Alargeprayingmantiswilleatasmallspider,andalargespiderwilleatasmallprayingmantis.Roundandroundtheygo,farabovetheground.

Thespider’seightlegsareallputtouseinextractingthesilkfromthespinneretsonitsabdomenandcreatingaweb.

Anotherfriendoftreesisthebat.Batsaretheonlymammalsthatfly,andtheyflyfastenoughtocatchnight-flyinginsects,includingmothsthataretryingtolayeggsonthetrees’leaves.Theseinsectsflyatnighttoescapepredationbybirds,butthatjustmakesthemvulnerabletobats.Althoughbatscansee,theireyesdon’tdothemmuchgoodonapitch-blacknight.Sothebathasevolvedanotherwaytofindinsects—itmakesahigh-pitchedsoundthatstrikestheinsectandechoesback.Thebat’slargeearscatchthereturningsoundsanditsbraininterpretstheechoessothebatknowswheretheinsectisandhowfastit’sflying.Butmothsarenothelplessinthissportof“catch-the-thing-in-the-dark.”Althoughhumanscan’thearthehigh-pitchedecholocationcallsofbats,mothscan.Whentheyhearthatsound,theycanquicklydroporchangedirectiontoevadethebat.Somemothsevenproducetheirownsoundstotrytojamthebat’ssonar.Sometimesbatseatinsectsonthefly,butmoreoftentheycatchthemintheirwingsandtakethembacktotheroostforeating.Veryoftenthatroostwillbeinatree,perhapsinahollowspaceorundershaggybark,sothebatprovidesaservicetothetreeandthetreeprovidesahomeforthebat.Besidesrelyingonbirdstoeatbugs,andotherinsectstoeatbugs,andbatstoeatotherinsects,treesalsoproducechemicalstodeterthethingsthatwanttoeat

them.Whenacaterpillaroraslugcrawlsacrossaleaf,thetreeknowsitisbeingtouchedbyaninsect,anditreactsbymakingchemicalsthateithertastebadtotheinsectorinterferewiththeinsect’sgrowthandreproduction.Eveniftherearenoinsectsonourtreebutatreenearbyisbeingeaten,ourtreecansensethecompoundsitsneighborisproducingandstartproducingdeterrentcompoundsofitsown.Tobringthestoryfullcircle,birdshavebeenshowntoincreasetheirhuntingactivityontreesthataregivingoffcaterpillar-deterringchemicals.It’sasifthetreehasemittedasilentcryforhelpandthebirdsrespond.Wehavemuchmoretolearnaboutallofthis.

Aplantleafknowswhat’seatingitbecauseitcansensethechemicalcompoundsonthefeetoftheinsect.Thisforesttentcaterpillarmaynotbewelcomedbythetree,butit’sanimportantfooditemforsomebirds.

Humansaremuchmorecloselyrelatedtobatsthanwearetobirds.Thebonesinabat’swingsareanalogoustohumanfingerbones,andbatshavesensitivecellsontheirwingmembraneslikethesensitivefingertip

cellsinhumans.Anotherlayerofavianlifeintheforestisthebirdsthatdon’tfeedoninsects,

seeds,orberries,butinsteadondeadanimals—thevultures.Thesebirdscleantheforest,liftingthedebrisawayonlovelywingsthatcircleoverheadinlazyspirals.Turkeyvulturesfindcarrionusingtheirextremelysensitivesenseofsmell.Justafewmoleculesofthechemicalgivenoffbythebacteriafoundonrottingmeatwillcauseavulturetocircleandcircleuntilitlocatesthesourceoftheodor.Vultureshavenofeathersontheirheadsbecausefeatherswouldjustholdthesmellandthebacteriafromtherottenmeattheyfeedupon.Butavulture’sbaldheaddoesletusseeanothersenseorganofbirds:ears.Thevulture’searlookslikeadentinitshead.Otherbirdshaveears,too,butwedon’tnoticethembecausethey’recoveredwithfeathers.Owlsmaylookliketheyhaveexternalears,butthoseearliketuftsarejustdisplayfeathers.Therealearsarehiddenunderfeathersonthesideofthehead.Externalearswouldcreateresistanceduringflightandthesoundtheywouldmakecuttingthroughtheair(attheveryhighspeedsanowlsometimesflies)woulddrownoutothersoundstheyneedtohear,likethescurryingofrodentsthroughleaves.Owlsdependontheirearsandlargeeyesforhuntingsmallanimalsatnight.Owlswillsitforhours,listeningandwatching,whilegraspingabranchwiththeirfleshyfeet.Whenpreyisdetected,owlsswoopsilentlydown,feetfirst,andgraspintheirtalonsthemealtheywillbringbacktothenestlings.Birdsofpreythathuntbyday,suchashawksandeagles,relymoreonsightthanonsmellorhearing.Theirvisionismuchbetterthanthatofhumans;theycanseetwiceasclearlyandmuchfarther.Athirdeyelidprotectstheireyesfromcuriouschicksorfromthedustanddryingeffectsofflyingmorethanahundredmilesperhour.Thiseyelidissomewhattranslucent,sotheycanstillseewhenit’sclosed.Raptorsdependontreesforaperchhighabovetheforestfloorwheretheycansurveythehappeningsandspottheirnextmeal—whichmightbeared-headedwoodpecker.Theyalsobuildnestshighinthecanopyforraisingthenextgenerationoftheirkind.Theyoungbirdswilltravelfarintheirlifetimes—seeingforestsyouwillnevergettosee—butiftheycan,they’llreturneachyeartotheforestwheretheyhatchedfromthatdelicateeggfarupinthecanopy.

Largeeyeshelpthisgreathornedowlseeatnight.Theearliketuftsarenotearsatallbutjustfordisplay.Thepresenceofscalesonanowl’sfoot

showsthatitiscloselyrelatedtoreptiles.

Turkeyvulturescircleuntiltheyfindthecarrionthey’vebeensmelling,thencarrythedebrisawayonlovelywings

Blackvulturesdon’thavethekeensenseofsmellthatturkeyvulturesdo.Theywatchwheretheturkeyvulturesgoandfollowthem.Becausethey

havenoheadfeathers,wecaneasilyseetheear.

Birdsofpreyhaveanextraeyelidtheycanseethrough;itprotectstheireyesfromdamageordryingduringhigh-speedflights.

Eaglescanseeclearlyformiles,acapacitydifficultforustoimagine.

Thered-tailedhawkisacreatureoftheforest.Theyellowareaonthe

beakoverthenostrilscanvaryincolor,servingtoadvertisethephysicalconditionandsocialstatusofthebird.

Isthiswhattheworldlooksliketoatree?Weknowthattreessenselight,butwedon’tknowwhatitlooksliketothem.Whatwouldtheworldlook

liketousifourskinwerecoveredwithtinyeyes?

EyesAllOverLeaves

TREESDON’THAVETWOeyeslikewedo,yettheycansee.Theyknowhowmuchlightishittingtheirleaves,andtheyknowthequalityofthatlight,too.Theyknowifit’ssummerorwinterbythelengthoftheday,andtheyknowifit’snoonorafternoonbythewavelengthofthelight.(Thenoonlightisstrongerandmoredirect,whiletheafternoonlightissoftenedbytheadditionaldistanceofatmosphereitmusttravelthroughwhenthesunisatalowerangle.)It’sasthoughthewholeplantiscoveredwithmillionsoftinyeyes—witheachphotosyntheticcellineachleafactingasaminiaturelightreceptor.Bothanimalsandplantssenselightwhenspecificmoleculesaretriggeredbyspecificlightwavelengths.Humanshavesomeofthesamereceptormoleculesthatplantshave.Redwavelengths?Itmustbemorningorafternoon.Bluewavelengths?Itmustbemidday.Plantsrespondtothesewavelengthsandtodaylengths,too,byproducingvarioushormones.Daysaregettingshorter?It’sfall,timetodroptheleaves.Middayinsummer?Ripenthefruit.Plantshaveevolvedthisincrediblesensitivitytolightbecauselightiseverythingtothem.Withoutittheydie.Lightisakeyingredientineverymorseloftheirfood(sugarmadefromsunlight,water,andcarbondioxide).Weareusedtotheideathatplantsneedsunlight,butwhenwethinkaboutthatconceptmoredeeplywefindthatithasmanyimplications.Forinstance,iftheleavesinthetopofatree’scanopyinterceptedallthelight,nobranchesorleaveswouldbeneededlowerdownandourtreeswouldlooklikegreenumbrellas.Insteadtreeshaveevolvedanarchitecturethatallowsthemtosharethelightthroughout.Andnotonlydoesthelightsharinggoonalongabranch,butthebranchesthemselvesarearrangedsotheywon’tcompletelyblockthelightforthebranchesbelow.Alongatrunkthebranchesalternateorgrowoppositeeachother,orperhapsevenspiralaround,butthey’reneverstackedoneimmediatelyabovetheotherlikealadder.Eventually,asatreegrowstaller—indeed,astheentireforestgetstaller—verylittlelightreachestheleavesonthebottombranches.Thetreethengetsonlyasmallenergeticreturnfromtheresourcesitputintomakingthoseleaves.Finallyitmakesnosensetoevenhaveabranchthereatall,sothetreestopssendingresourcestothebranch.Thisprocessmayhappenveryslowly,butfinallythebranchdiesasaresultofthelowlight,andperhapsinthenextwindstormthatbranchwillfall.Manydecadesofthisleadtoatalltreewithmostbranchesonthetopandveryfewlowerdown.Welookattreesliketheseandknowtheyare

“forest-grown”trees.Theyhaveliterallybeenshapedbylight,orthelackofit.

Thisforest-growntreeshowsatypicalbranchingpatternwithveryfewbranchesloweronthetree.

Thesebeechtreeleavesarenolongeractivelyphotosynthesizing—it’sfallandtheirgreenchloroplastshavegoneintothetree’srecyclingbininpreparationforleafdrop—butwecanclearlyseethewaytheleaves

sharethelightspace.

Ifyoulieonyourbackintheforest,youcanexperiencethesharingoflightspacefromtheunderside.Warning:thisreveriecouldfillmany

hours.

Eachseasonfewerandfewerleavesareproducedonthelowerbranches.

Ifatreegrowsinanenvironmentwithabundantlightandnoshadingfromothertrees,itmaybeabletokeepitsbranchesallthewaytotheground.Thinkofagrandliveoak.Thisphenomenonoftreeslosinglowerbranches,ornot,dependingonlightlevels,givesuscluestothepast.Whenweseeabigtreeinthemiddleofaforestthathaslargelowerbranches—livingordead—wecansurmisethatthattreeoncegrewintheopen.Mostlikelythetreestoodinapastureandshadedfieldanimals,andperhapsfieldworkers,fromtheharshsun.Intime,forvariousreasons,thepasturewasleftungrazedandunmownandthegentlecreepofsuccessionresultedinaforestspringingupwherenonehadrecentlybeen.Afterafewdecadesthegrandoldpasturetreebecameapartoftheforest,butitslowerbranches,deprivedoflight,begantheirslowdecline.Treepeoplecallthesewolftrees.Perhapsthistermcamefromtheideathatwolvesgobbleupallthefoodaroundthem,andforesterssawthesetreesasgobblinguplotsofspacethatcouldhavebeenthedomainofthetall,thintreesthattheirmillswerebuiltfor.Inthepastitwasthoughtthatthesetrees,liketheanimalstheywerenamedafter,shouldbeeliminated.Butthewolftreeswereoftenunmanageable,andsotheywereleft.ManyoftheseancientoneshavebeenwitnesstoNativeAmericanencampments,tothenow-extinctpassenger

pigeon,andtothebirthoftheUnitedStates.Todaysomeofthesetreesareourlargest,oldest,andmostbeloved.Birdsandotheranimalsarefoundinmuchgreaterabundanceinthesewolftreesthanintheonesreadyforfactoryproductionlines.Unlessyouunderstandtheprimalurgeoftreestogrowtowardthelight,youmightbetemptedtolookataforestcanopyandthinkthattwonearbytreesarepurposelysharingthespace.Butwhenyouseeasatreedoes,yourealizethatoneisavoidingtheothertree’sbranchesbecausethebranchisdark,notlight.Morebudsdeveloponthesunnysideoftheplant,sothat’sthedirectioninwhichitwillgrow.

Thisbeechtree,perhapsthesameageastheoneonpage36,didn’tgrowinaforest.Althoughtheirgeneticsmaybethesame,thisopen-

grownbeechtreehasaverydifferentformfromonethatgrewinaforest.Treesarelight,embodied.

Youngtreesthatdon’tgetenoughlightwillneverreachthecanopy.Butit’snotjustblackorwhite,orevenshadesofthesecolors.Atree’sleavescansensethespecificwavelengthscontainedinthelight.Theydothisthroughchemicalmediatorsthesamewaywedo—certainwavelengthsactivatecertainmoleculesinalivingcell.Inhumansthishappensmostlyintheretina,andtosomeextentinourskin,butinplantsithappensinallthelight-sensitivecells.Shinedaylightthroughaprismandyou’llseeallthecolorsoftherainbowasthelightissplitintovariouswavelengths:red,orange,yellow,green,blue,indigo,violet.Toplantsthisrainbowlooksdifferentbecausetheycan“see”acolorwecan’t:far-red.Theirrainbowis:far-red,red,orange,yellow,green,blue,indigo,violet.Themostimportantwavelengthstoatreearethosethatresultinthecolorsredandblue.Thosearethewavelengthsusedmostinphotosynthesis.Whensunlightshinesdownthroughaforestcanopytheredandbluewavelengthsareabsorbed,andotherwavelengths,suchasfar-redandgreen,passthroughorarereflected.Soaleafknowsit’sintheshadeofaforestwhenredandbluelightarescarcebutgreenandfar-redlightareabundant.Inthiswaytheshadecreatedbylivingplantsdiffersfromtheshadecastbyarockorabuilding.Whenaplantsensesthelightwavelengthspresentinashadyforest,itrespondsbygrowingdifferentlyfromhowitwouldgrowintheopen.It’snotonlybranchesthatwitheranddieduetolackoflight;youngtreesmaymeetthesamefate.Inaleafygreenforestweseemanytreessharingthespaceandlight,andevenlowbranchesstillholdingsomeleaves.Butifwereturndecadesfromnow,someofthetreeswillbemuchlarger,havingsuccessfully

capturedthelight,whileothertrees—losersinthebattleforthelight—willbedead.Althoughithasn’tbeenstudied,humans,too,mayhavebiologicalresponsestoforestlight.Wedon’thavereceptorstosensefar-redlight,asplantsdo,butwhenwewalkintheshadeofaforestwearebeingbathedinfar-redlight.Weknowthattimespentinaforestreducesbloodglucose,bloodpressure,andstresshormonelevels,buttheexactreasoniselusive.Chemicalcompoundsgivenoffbytheleaves?Perhaps.Lightwavelengths?Possible.

Everydifferentcolorrepresentsadifferentwavelength.Plantsrespondtospecificwavelengths.

Imaginethatinsteadofwalkingthroughanenvironmentdefinedbylightanddark,youcouldonlygrowthroughit.Theneachdecisionwouldbe

likethis:light,light,towardthelight.Treesaresensitivenotjusttowherethelightisbutalsotohowlongitlastseach

day.Inspringthetreesleafoutinresponsenotonlytotemperaturechangesbutalsotochangesindaylength.Animalsrespondtochangesindaylengthinverysimilarways:detectionandthenhormonerelease.Whiletheplantresponsetolongerdaysinvolvesleafexpansion,theanimalresponsefrequentlyinvolvesmatingbehavior.Suncyclesaretheprimarydriverofplantresponses,butwhataboutthatbrightmoonreflectingthesun’srays?Experimentsshowthatplantscandetecteventhelowlightlevelsofmoonlight.Full-moonlightpromotesfloweringandrootgrowthinsomeplants(althoughveryfewplantshavebeenstudiedregardingtheirresponsetomoonlight).Inotherplantsmoonlightmayinterferewiththeirtimingofdaylengths,sothey“sleep”bychangingtheangleoftheirleavesatnight—theleavescloseormoveawayfromthemoon.Animalbehaviorintheforestalsochangesinresponsetolight.Manyforestanimals,likebats,ants,andreptiles,aremostactivejustaftersunset.Onfull-moonnights,nocturnalrodentsstayputbecausethey’reeasiertargetsforowlsthen.Night-flyingpollinators,likemoths,aremoreactiveonmoonlitnights.Noonehasstudiedhowforesttreesrespondtomooncycles.Mightnut-bearingtreesbemorelikelytodroptheirnutsonanightwhenrodentswon’tbeouttoeatthem?Onethingwedoknowisthattheforest,asanecosystem,isawareofthemoon’scycles.Soweknowthatplantsandanimalsaresensitivetosunlightandmoonlight.Whataboutstarlight?Birdsaresensitivetostarlightanduseittoaidtheirlong-distancemigrations.Trees?Imagineyouhavelivedforhundredsofyearsinthesamespotwiththesky,thefocusofyouryearnings,constantlyoveryourhead.Youareexceptionallyskilledatsensingandusinglight.Hourafterhour,nightafternight,yearafteryear,thesmallfar-awaysunsrollby.Doyouthinkyoumighthavearelationshipwiththestars?Mostlikely,butitisarelationshipwehumansdon’tyetunderstand.

Moonlightisthesun’slightreflected.Forestplantsandanimalsrespondtoeventheselowlevelsoflight.

Aretreessensitivetostarlight?Wedon’thavetheanswertothatyet.

Rodentsintheforestbehavedifferentlyduringthefullmoon.

Forestshelpedtocreatethissurface-levelcloudand,throughcondensation,theycanalsorecapturesomeofthemoistureitcontains.

ADeepBreathLungsoftheLand

WHOHASN’TBREATHEDONamirrororawindowonachillydayandseenwatervaporcondenseonitssurface?Orwitnessedtheirwarmbreathcreatesteamwhenit’sexhaledonafrigidwinter’sday?Thesearevisibleremindersofanormallyinvisiblephenomenon—watermovingfromourbodiesintotheenvironment.Treesexhale,too;theypassivelyexhalewatervaporthroughmultitudesoftinyporescalledstomatathatcovereachleaf.Indeed,mostofthewatermoleculesthatentertheirrootsandjourneyupthatlong,darkstemarereleasedfromtheleavesunchangedthroughthesepores.Wearestilllearninghowstomatafunction.

Imaginetheforestexhalingwatervaporandoxygen,andinhalingcarbondioxide,forthat’swhatitdoes.

Stomataarevisibleonsomeplantfossils,soweknowthatthesizeandnumberonatypicalplanthavechangedovertime.Manymillionsofyearsago,thefirstplantshadlargerstomatabutfewerofthem.Whencarbonlevelsintheatmospheredropped,someplantsevolvedtohavesmallerbutmoreabundantstomata.Inpart,itwasthischangeinstomatasizeandnumberthatallowedmoreplantspeciestoevolveandinhabitthedrierregionsoftheplanet.Nowthatcarbonlevelsarerisingagain,thenumberofstomataperareaofleafisagaindeclining.Plantsdorespondtotheenvironment,althoughsometimesthatresponseisveryslow.Wearebornwithonemouth,butthenumberoftinypore-mouthsthatwilldeveloponaleafisdeterminedbytheatmospheresurroundingthatplant.How

doesajust-developingleafknowthenumberofstomataitwillneed?Theexistingleavestellit,fortheyhavebeensensingandrespondingtothatatmosphere.Stomatadon’topenandclosesimultaneously.Insteadeachonebehavesasanindividual.It’ssuchaquietprocessthatwearen’tawareofitwhilewalkingthroughtheforest,butagiantmicroscopeprojectinginrealtimewouldfascinateusasitshowedtheopeningsandclosingsofthestomata.Itwouldbelikewatchingfirefliesflashingonahotsummernight—youknowthere’sarhymeandreasonineachflash,butthecomplexityofthecombinedflashescausesyourmindtostoptryingtofigureitout,andfinallyyoujustenjoytheshow.Beyondtheshowofthesepulsingpores,wemustalsoconsiderthehugeglobalinfluencetheyhaveonregulatingatmosphericlevelsofwater,carbondioxide,andoxygen.Foratthesametimethatthestomataarereleasingwatervapor,they’realsoreleasingoxygenandsippinginthecarbondioxidetheplantneedsforphotosynthesis.Everyyearthemajorityofthewatercirculatingthroughtheatmospheremovesthroughstomata.Theoxygenyou’renowbreathing,themoleculethat’skeepingyoualive,hasatsomepointcycledthroughatree.Everyyear40percentofthecarbonintheatmospheremovesthroughstomata.Thesetinyplantporescanchangetheclimateoftheplanetandinfluencethecourseofevolution.Ponderthatasyouwalkthroughtheforest.Thetreesarebreathingoutandyouarebreathingin.Youarebreathingoutandthetreesarebreathingin.

Averycloselookataleafrevealsthetinyveinsthatcarryfluidstoandfromeachcell.Anevencloserlook,withamicroscope,revealstheporesthatallowgasessuchaswatervapor,carbondioxide,andoxygento

passintoandoutofthecells.

Plantssometimesseemsostatic,butinmanywaysthey’remoreresponsivetotheenvironmentthanweare.Thecellscontrollingthepore

openingsareconstantlyadjustingtochangesinlight,humidity,andcarbondioxide.They’reevenawareofwhat’shappeningintherootzone.Whereaswehavemusclestohelpuspumpoxygeninandcarbondioxideandwatervaporout,treesmustdependonamuchlargerandmoreremotepump—thesun.It’sthesun’sradiantenergystrikingthepigmentmoleculesonaleaf’ssurfacethatputsthewholegreenbreathinmotion.Thougheachporeoperatesasanindividual,there’salsoalargerdailyrhythm.Thinkofhumansinacityrisinginthemorning,accomplishingtheirdailytasks,andthenretiringatday’send—alargerpatterntoallthemanyindividualvariationsonthistheme.Imaginethispatternfromaforest’sperspective.Astheplanetslowlyrotateseastward,thelife-poweringorbappears.Onlythetallestofthetreescatchherfirstlight.Asthelightbrushesthetreetops,longshadowsarecast—notontheground,butontheveryair.Theforest’sgiantexhaleofwatervaporhasbecomethecanvasforhershadows.Coolmorningtemperaturescondensethevaporuntilit’svisible,atreetopcloud.Waterdropsthatwererecentlyinthedarkearthhavebecometransformed.They’vemovedintotheroots,squeezedthroughthe

trunk’spassage,escapedthroughtheleafpores,andarenowfloatinginair.Eachofthesetreeswillshepherdtensofthousandsofgallonsofwaterintotheatmospherethisyear.Everydayofthegrowingseason,atreemovesandreleasesenoughwatertofillabathtub.Andthatwaterneverdisappears:itmaybecaughtagainbythetrees,whenvaporcondensesontheleavesorthetrunkandrollsdownintothesoilonceagain;orthewindmaycarrythevaportoadistantforestthatwillbequenchedbytherainfromitsunknownpartner;orthewatermayfinditswaythroughrunoffandriverstotheglassbyyourbedside.Laterinthedaytheairwillwarmandthedropletswilldisperse—ourwhitecanvaswillfadeaway.Thewarmaircolumnwillriselikeaninvisibleballoon.Asthewarmairrises,itcreatesasuctionthatpullsinairfromotherplaces.Thisverylargepatternisliketheearthtakingadeepbreath.Airthat’spulledinfromacrosstheoceancontainsmoisturethatwillfallonthelandasrainorsnow.Thisatmosphericmovementiscriticaltotheexistenceofforestsontheplanet:theprecipitationthatfallsontheforestandenterstheatmosphereaftertravelingupthroughthetreesiscarriedaloftandofftowateranotherforest.Inaway,theforestsarewateringeachotherandthemselves.

Watervaporreleasedfromaleafmayfalltoearthasrainandthenflowintoariver,whereitbecomesourdrinkingwater.

Thesun,nowhighoverhead,castsshadowsonthegroundandprovidestheenergyforourforesttocontinueitsvitalroleofwatermover.Theleaves,likemillionsoflittlefactories,willmakesugarstopowerthetree.Water,oneoftherawingredients,mustbeimportedfromtheground.Carbondioxide,theotherrawingredient,mustbeimportedfromtheair.Oxygenismerelyaby-productof

theprocess,butonethatwearevastlygratefulfor.Anotherwayforestsbreatheisbyplantsdrawingincarbondioxideinthedaytimewhilephotosynthesizingandthenreleasingcarbondioxideatnightasthecellscontinuetofunctionandrespire.Thisisalong,slowbreaththattakestwenty-fourhours.

Water,carbon,andoxygencyclethroughtheforestinalong,slowbreath.

Forestsplayanimportantroleinthehydrologicalcycle.

WorldofWaterCyclesandCreatures

FORESTSANDWATERGOtogether.Forestshelpmakerainbypullingwaterfromthesoilandreleasingitintotheairwhereitrises,cools,andcomesbacktoearth.Butthat’snottheendofthestory.Totrulyknowaforest,youmustunderstandhowitmanagesthewateritreceivesfromtheatmosphere.Someofthewaterthatreachesaforestisstillinvaporform.Fogandmistdriftthroughtheforest,andthevaporcondensesonleavesandneedles.Snowfallsgentlyandcollectsontwigsandneedlesbeforeitreachestheground.Tinyraindropsfallslowly,andthese,too,maygatheronleavesandtwigsbeforeslidingslowlytowardearthalongthetrunk.Themossonthetrunkeagerlyawaitsthismoistureandholdsontoasmuchofitasitcan.Thebark,too,soaksupsomeofthewater.Duringashort,gentlerain,thecanopycatchesanddirectsvirtuallyalloftheprecipitationtothetrunks,andthenthetrunksandthethingsgrowingonthemabsorbeverythingthatreachesthem,leavingnonefortheground.Perhapsyou’veexperiencedbeinginaforestduringarainwhenyoubarelyfeltadrop,althoughyoucouldhearit.It’snotdifficulttoimaginetheforestasaliving,breathingsponge,holdingontothelife-givingdampnessthatcomesfromthesky.Butsometimestherainfallsinbuckets,thelargedropscomingdownhardandfast.Inopenplaces,suchforcefulraincausessoilerosionandcompaction.Runofffromsucharaincarriessoilparticlesthatcloudwaterwayswithsediment.Forestspreventthat.Whenahardrainfallsonaforest,thedropsareslowedbeforetheyreachtheground.Inthesummertheyhitleafafterleafafterleaf,rollingandsplashingfromonetotheotheruntiltheyplopgentlytoearth.Eveninwintermonths,whentheleavesaredown,thefinetwigsbreaktherain’sfall.Andwhentheslowdropsreachtheground,theylandonacushionofdecomposingwoodandleaves.Thisspongylayerholdsontomostofthemoisturefromashortrain.Onlyalongraincansaturatetheorganiclayerandsoakintothemineralsoilbelow.Thisallhappenssogentlyandslowlythatnosoilparticlesarecarriedaway.Mostoftherainwaterstaysrightintheforest.Inthesummeritmaybetakenintothetreeroots,carriedup,andreleasedthroughtheleaves.Inthewintertheexcesswatermaytrickleslowlyintothegroundwaterorintoanearbystream.Oritmayformpoolsintheforest.Thiswaterhasbeenwashedbytheforest.Theforestslowlygivesbackclean,clear

water.Sometimesitseemslikewaterspringsrightfromtheforestsoil,anditdoes,asthespongysoilreleasesitdropbydropfrommanysurroundingacres.Therearefewpleasuresgreaterthanenteringashadyforestonanotherwisehot,drydayandfindingarushingstreamofcool,cleanwater.Andsincesomanyofourremainingancientforestsareinrockyplaceswhereitwasdifficulttobringmachines,thestreamsareoftenfoundrushingoverrockyoutcrops.Yourfirstemotionalresponsemaybeexhilaration,thenjoy,andifyouremainstilllongenough,despitethesoundofrushingwaterasilencewilldescendonyou.Thisisagiftoftheforest,too.

Needlesandleavescancatchmoistureorsmallraindropsfromtheairandtransfertheliquidgentlytotheground.

Rainsoaksintothesoftsoil,andtheforestslowlyreleasesthecleanrainwaterithascaptured.

Cool,flowingwaterinadarkforestisatreatforthesenses.Andthiscleanwaterwillflowfromtheforestouttotheclosestriver,andtherivereventuallytoalakeoranocean,allcleanerforhavingstartedintheforestworld.Eachofthesebodiesofwater,fromsmalltolarge,ishometoplantsandanimals.Allofthem,andallofus,havetheforesttothankforhelpingtokeepthewaterclean.Acrayfishcouldnotliveinitstinyrivuletunlesstheforestcanopykeptthelandcoolandshaded,andpreventedmudfromenteringthestream.Amuddystreamwouldinterferewithfindingprey—andevenwithgettingoxygen.Thedamp,spongyforestsoilreleasesitswaterdropbydropsotheshallowstreamflowsevenindryweather.Noforest,nocleanwater,nocrayfish.Althoughitmightlooklikealobster,acrayfishwillnevergetmorethanafewincheslong.Itwilleatjustaboutanythingitcanfind,includingfrogs,salamanders,dragonflynymphs,andevensmallfishwhenitcancatchthem.Asthestreamsgetdeeper,fishcanliveinthem.Fishspeciesvarywidelyintheirtolerancetotemperatures,oxygenlevels,andwaterclarity.Somespecies,likecarp,canliveinwarmwaterandcometothesurfacetogulpairifoxygenlevelsarelow,butotherspecies,suchasbrooktrout,needcoldandhighlyoxygenatedwaters.Streamsideforestskeepthewatercleanandcold.“Troutgrowontrees”isatrademarkedsloganoftheStroudWaterResearchCenter,aclearreminderofthecloserelationshipbetweenthetwo.Troutdisappearfromastreamafteraslittleas4percentofaforestedwatershedsurfacegetscoveredwithpavingor

buildings.Whensearchingforlifeonotherplanetswesearchfirstforwater,becausewaterequalslife.Asmammalswehaveevolvedtocarryourwaterwithinus.Awomanmaywalktoastream,takeadrink,andthenwalkmilesfromanystandingwater,carryingherunbornbabyinthewateryenvironmentofherwomb.Reptileshaveevolvedasimilartrick,buttheystoretheembryowithwateroutsidetheirbody,inanegg.Totellreptilefromamphibian,lookclosely:ifithasscales,it’sareptile.Askinkmaylookmuchlikeasalamander,butbesidesbeingfour-leggedanimalsintheforestthey’reonlydistantlyrelated.Askink,beingareptile,layseggsthatdon’tneedstandingwater,andtheanimalthathatchesfromthoseeggsisalreadyfullyformed.Themoreprimitiveamphibians,likesalamanders,dependonthewaterinaforesttobetheirnursery.

Crayfishhavemultiplesetsofantennaeandlegs.Theycanvoluntarilyreleaseanappendagetoapredator;avalvethensealsoffthemissingsectionsobodyfluidsdon’tescape.Theappendagewillgrowback.

Abrooktrouthidesneartherocks.Thesurroundingforestkeepsthisstreamclearandcold,conditionsthatarevitalforthetrout.

Ashallowpoolintheforestisimportantreproductivehabitatformanyfrogs,salamanders,andinsectswithaquaticlifestages.

Notallthewaterthatentersaforestleavesitbystreamsandrivers.SometimesthewaterisjustheldthereinlargepuddleswecallbypoeticnamessuchasCarolinabaysorephemeralponds.Theseshallowforestpoolsarealsocalledvernalpoolsbecausetheycontainstandingwaterinthespring.Allfallandwintertherainandsnowhavebeenfalling,andtheforestsoilissaturated.Springwalksrevealwidepuddlesintheheartoftheforest.Manyamphibiansreleasetheirreproductivecellsintothiskindofwateryenvironment,andthefertilizedeggseventuallyhatchintofree-swimminglarvae.Besidescallingyououtfromyourhouse,thewarmingtemperaturescallspottedsalamandersoutfromtheirundergroundburrows.Onarainyspringevening,thousandsofthemmaycongregateatasinglepool.Theretheysocialize,andwhenamalefindsafemalehebeginsanancientchoreography.Firstheswimsupunderneathher,wiggleshistail,andswimshertotheshallowedge.Ifsheshowsinterestbyrubbinghernoseonhim,hewillbegindepositinghisspermpacketsontheshallowunderwatervegetation.Thespermareinanovalcaponatinytowerofgelatinthatheproduces.Thefemalethenpositionsherselfoveroneofthesetowersandsettlesdownontoit.Whensheleaves,theovalcapwiththespermisinsideher,butthegelatinoustowerisstillthere.Shethenlaysherfertileeggsinthepool.Finallytheadultscrawlbacktotheirseparateburrowsin

thesoil,deepunderthefallenleaves.Theyonlycomeouttofeedondampnights,andtheywon’treturntothepooluntilnextyear.Meanwhiletheirembryos,coveredingreenjellylikeballs,areontheirwaytobecominglarvae.Theeggballsaregreenbecausetheycontainalgae.Isthealgaeattractedtothenitrogenreleasedbytheembryo?Doestheembryoneedtheoxygengivenoffbythephotosyntheticalgae?Nooneissure,butwedoknowthatthesalamanderembryosdon’tdowellwithoutthealgaeandthealgaedon’tdowellwithoutthesalamanders.They,likesomanyotherthingsintheforest,needeachother.Aclosemicroscopicexaminationrevealsthatthetinyembryosactuallyhavethealgaethroughouttheirbodytissues.It’snotafarstretchtoimaginethatthesesalamanderembryosarephotosynthesizing.Aftertheembryoshatchfromtheirgreeneggs,thelittlelarvaeswimthroughthepool.Nowtheycanfeedontheirownandnolongerneedthealgae.Theyhavewhatappeartobefeatherywingsspringingfromtheirnecksjustbehindthehead.Theseareexternalgillsthatpickupoxygenfromthewater.Inafewmonthsthelarvawilldeveloplittlelegs,itsexternalgillswillshrink,anditwillbereadytocrawloutofitsnatalpuddle.Salamandershaveavarietyoflifestyles.Theyarein-betweencharactersenjoyinglifeinthewaterandontheland.Red-backedsalamandersspendnopartoftheirlivesinstandingwaterandyetwithoutmoistureintheirenvironmenttheycan’tbreathe,fortheyhavenolungsandmustdependonoxygendiffusingthroughtheirdamp,permeableskin.Others,likethejust-discussedspottedsalamander,mateandlayeggsinthewater,andtheyounglarvaebreathethroughgills,laterdevelopinglungsastheymovetotheland.Themarbledsalamanderfeelstheurgetomatenotinthespringbutinthefall,andmatinghappensonland.Hedisplayshimselftothefemaleandthendepositsaspermpacketontheground.Ifthefemaleisresponsive,sheacceptsthepacketanddrawsitintoherbody.Shethenmakesherwaytoapool.Ifit’sdry,asitislikelytobeinearlyautumn,sheburiesherselfunderthedetritus,layshereggs,andwaits.Thefallleavesdriftdownuponher.Shewillstayundergroundwithhereggsforweeks.Shewon’teatatallduringhervigil;hermerepresencedeterseggpredators.Atlasttherainsbeginandtheleaflesstreesnolongersuckwaterfromtheground.Thepoolfills,andtheeggshatch.Herworkisnowdoneandshecanreturntodrierlandwhilethelarvaeswimintheirnewoceanandsearchforone-celledorganismstoeat.Eventhesmalllarvaeareresponsivetotheirenvironment.Whennightfallsafteraclear,sunnyday,thelarvaealldriftuptothesurfaceofthedarkpooltofeed.Whenthemorningsuncomesout,theysinkbackdownamongthesubmerged

leaves.Onfullmoonnightstheystaylow,becausethemoonlightallowspredatorstospotthem.Butduringaneclipseofthemoontheyrisebackupandresumefeedingforabrieftimeundercoverofdarkness.Afterafewmonthsthey’relargerandhungrier,andsuddenlyamiraculousabundanceappears.Thespringpeepersgatherattheirpoolandlaymassesofdeliciousgelatinouseggsthatarelikemannafromheaven.

Alarvalsalamanderhasexternalgillsthatcollectoxygenfromthewater.Afewspeciesretainthesegillsintoadulthood,butinmostspeciesthe

gillsdisappear.

Thefaceofalarvalsalamanderhasdarkareasbetweentheeyesandthemouththatcandetectchemicalscentsinthewater.Salamanders

havehighlydevelopedsensesofsight,smell,andtouch.

Althoughitlookssimilartoasalamander,thefive-linedskinkhasscalesthatidentifyitasareptile.Femaleslayeggsinmoistdecayinglogsand

remaintoprotectthemuntiltheyhatch.Soontheuneatenfrogeggshatchandlegionsoftadpolesswiminthepool—andnotjustpeepertadpolesbutmanykindsoffrogsandtoads.Afewfrighteningthingsappearinthepool,too,likedragonflylarvae.Thesestrange-lookingthingsjustsitonthebottomofthepoolhidingandthengrabanytinytadpoleorsalamanderlarvaethatswimby.Yellow-spottedsalamanderscometolayeggs,too,andtigersalamanders.Thetigersalamandersarealittlelargerthantheothersandmatealittledifferently,butliketheothers,theyalwaysreturntotheirhomepondtobreed.Salamandershavetinyhomeranges—measuredinfeet.Otherthanbreedingtime,theymayspendtheirwholelivesaroundthesamerottedlogorpileofrocks.Togettotheirbreedingpond,theynavigatebythesunaswellasusingtheirveryacutesensesofsightandsmell.Whentheyarriveatthepond,themaletigersalamanderbumpsintothefemale,andthisishiscuetoherthatsheshouldfollow.Shemaylayathousandfertileeggs.

TheforestsofthesouthernAppalachianMountainsaretheglobalcenterofsalamanderbiodiversityanddensity.Thesespecieswereallfoundin

onedayinonelocationinVirginia.

Springpeepersarethefirstfrogstostartcallingintheveryearlyspring.Asinglefrogsoundslikealoudchickpeeping,butawholechorusof

themsoundslikeasymphonyofsleighbells.

Tadpoleshavenolimbswhentheyhatch;overanumberofweeksfirstthebacklegsform,thenthefrontlegsappear.Afewdaysfromnowthis

littlefrogletwillexchangegillsforlungsandcrawlupontotheland.

Mostofadragonfly’slifeisspentunderwater.Thisnymphisreadytocrawloutofthewater,breakoutofitsoldskin,andbecomeacreatureof

theair.

Theadultbluedasherdragonflywilldefenditsperch.Astheweeksrollbyandtheairgetswarmer,thetreeleavesemergefromtheirbuds.Likestraws,thethirstytreerootsaresuckingthevernalpooldry.Thisisamixedblessing:ifthepooldriesuptooquicklythelarvaemaynotmakeittoadulthood,buttheveryephemeralnatureofthepooliswhyitcontainsnofishthatwouldhappilyeateggsandlarvae.Meanwhilethesalamanderlarvaearegrowinglongerlegs,andtheirfeatherygillsaredisappearing.Thetadpolesaregrowinglegs,too,andtheirtailsareshrinking.Soonallthecreaturesofthepoolclimbout,justbeforethepooldriesupcompletely.Oursalamandershavebecomecreaturesoftheland.Theywillburrowafootortwodownandliveaprimarilysubterraneanlife.Wewalkbyoverheadwithnoclueofwhat’sunderourfeet.Butthetreesarehelpedbythepresenceofthesalamanders.Theburrowsprovidechannelsforoxygenandrainwatertoreachtheroots,andthewaterthatflowsoversalamanderexcrementislikeadoseoffertilizerforthetrees.Initssecondautumntheyoungmarbledsalamanderwillmakeitswaybacktothepondwhereithatched,anditwillreturnformanyyears.Thespottedonemayreturntothesamepondfortwentyorthirtysprings.Whilesalamandersareontheirwaytoandfromthepoolswheretheyhatched,theyfindinsectsorwormsortinyfrogstoeat.Aturtle’spatternisoppositethatoffrogsandsalamanders.Whenmostsalamandersareheadingtowardwatertolayeggsinspring,femaleturtlesareleavingthewaterandheadingtowardlandtolayeggs.Althoughturtlesliveinlargerstreams,rivers,andpondsthatdon’tdryoutinthesummer,theyneedhealthywildlandnearbyfornests.Smallerspecies,likethepaintedturtle,layeggstwoorthreetimesayear,butabigoldsnappingturtlewillonlyhaulherselfaroundtodiganestonceayear.Otheranimals,suchasraccoonsandsnakes,dependonthesenutritiouseggsasanimportantsourceoffood.Asaresult,onlyasmallpercentageofturtleeggslastuntilhatchingtime.

Howmanysalamandersareinthisphoto?Wedon’tnormallyseethembecausetheyspendmostoftheirtimeundergroundwheretheirskincan

staycoolanddamp.

Onewaytigersalamandersprotectthemselvesisbyreleasingtoxicsecretionsfromglandsbeneaththeskin.

Likesalamanders,turtlesofdifferentspeciessharecommonalitiesbutalsohave

variationsintheirlifestyles.Onethingtheyallshareistheirbasicstructure.Foronething,turtleshavenoteeth;insteadtheirjawbonesaresharpandnotcoveredwithfleshasoursare.Andimaginehavingyourbackboneontheoutsideofyourbody—that’swhatlifeislikeforaturtle.Itsshellisactuallyamodificationofspineandribbones,sodespitethecartoonsthatshowturtlescrawlingfromtheirshells,thatcouldneverhappeninreality.Andthisbonyshellneedscare.Algaegrowonturtleshellsaseasilyasonrocks.Onereasonyouseeturtlesbaskinginthesunattheedgeofthewateristodryofftheshellsoalgaewon’tgrowonit.Thebaskingalsowarmstheturtleupsoitsmusclesarereadytoswimandhunt.Itcan’tproduceitsownbodyheat,soit’stotallydependentonthehelpofthesunshineandaplacetocrawlupoutofthewater.Mostoftentheidealbaskingplaceisashorelinetreethathasfallenintothewater.Forestsandturtlesareconnected.Whenapaintedturtleiswarmedup,itwillswimalongtheshallowwaterpokingitsheadintothevegetationandstirringupthingstoeat,liketadpoles,insectlarvae,smallmussels,andcrayfish.Itswebbedfeetareidealswimmingpaddles,anditslongclawshelpitdiginthemudandtearapartitsfood.Onceitsmealhasbeenconsumed,theturtlewillcrawlbackoutintothesunshine,andthewarmthwillhelpmoveitsdigestivemuscles.Asnappingturtle’sbaskingbehaviorisabitdifferent.Imaginetryingtohaulthathulkofabodyupontoalog.Insteaditfloatsonthesurfaceofthewaterandletsthesunwarmitthere.Itssnoutsticksupabovethesurfacebecause,likeallreptiles,itneedsairtobreathe;butwhenit’sreadytogotothebottom,coveritselfwithmud,andwaitforsomethingdelicioustohappenby,itcanholditsbreathforalmostanhour.Althoughsnappingturtleshavealifespanlikethatofhumans,andittakesthemaslongashumanstoreachsexualmaturity,theirspecieshasbeenaroundformuch,much,muchlonger—fourhundredtimeslonger—thanthehumanspecies.Turtleswerearoundduringthetimeofdinosaursandlivedthroughtheextinctionepisodethatkilledthem.Lookagainataturtle’sfaceandthinkaboutthat.

Paintedturtleswillwalkmilesoverlandtofindmates,makingthemvulnerablewhilecrossingroads.Somevolunteergroupsputuptemporaryfencingalongroadwaysandcarrytheturtlesacross.

Mapturtlespreferlargerbodiesofwater.Duringthewarmseasonwhenthey’reactivetheycometothesurfacetobreathe,butinwinterwhenthesurfaceiscoveredbyicetheyslowtheirmetabolismandrestonthebottomwheretheysurvivebyabsorbingoxygenthroughtheirskin.

Thislargeoldsnappingturtle,likeallturtles,ismorecloselyrelatedtobirdsanddinosaursthantolizardsandsnakes,somethingwe’velearnedinthepastfewyearswiththedevelopmentofsophisticatedgenetics

techniques.

Unlikeothereasternturtles,boxturtlesarecreaturesoftheland.

OntheGroundAnimalsoftheForestFloor

WEKNOWTHATTHEforestishometomanyanimals,butit’srareforustoseethem.Humanstendtobeloudandclumsycreatures,givingtheanimalstimetohidebeforeweapproach.Thosewearemostlikelytoseearetheslowones,likeboxturtles.Althoughthey’requiet,harmless,andnotuncommon,westillfeeldelightwhenwefindthesewildcreaturescrawlingacrosstheground.Ourdomesticpetsdependonusfortheirfood,butthewildthingsintheforestmustfindtheirown.Whatdoestheslowturtlefindtoeat?Boxturtleseatjustabouteverything.Thisturtlemightbelookingforhisfavoritefruit,themayapple.Danglingbeneaththelowleaves,thissnackisatjusttherightheightforaturtle.Hewillcrawlawaywithseedsinhisbellyanddeposittheminanotherpartoftheforestwhereperhapstheywillstartanewcolony.Onthewayhemightalsonibbleonfallendogwoodberriesorsycamoreseeds.Maybethisturtle’seyeiscaughtbyacicadanymph;justcrawlingtothesurfaceafteryearsunderground,itwouldmakeafinemeal.Orperhapsheiswatchingthemushroomgrowthatwillbetomorrow’sbreakfast.Arehiseyessharpenoughtoseethetinysporesthatwillpassthroughhisbodyandbedepositedalongwithdigestedorganicmatterthatmakestheperfectcompost?Asnailisfeastingononeofthemushrooms.Turtlesliketoeatbothsnailsandmushrooms.Whichonewillhechoose?Snailsareaveryimportantpartoftheecosystembecauseinadditiontoeatingmushroomstheyeatdeadleaves,afoodfewothermobileorganismsareinterestedin.Thedeadleavescontainsmallamountsofcalcium,andthesnailsstorethiscalciumintheirshells.Otheranimalsthatneedcalciumintheirdiets—liketurtlesandturkeys—eatsnailstosupplythisneed.Turkeysneedthecalciumtoproducestrongeggshells,andresearchshowsthattheyseekoutandeatmoresnailsduringtheegg-layingseason.Turtlesneedcalciumtoensurestrongshells.Possiblytheboxturtleislookingforthepatternedshellofafemale.Eachturtle’sshellisunique,andmaybehewillrecognizetheshellofashe-turtleheencounteredlastspring.Hisfive-toedfrontfootreachesforwardaninchwhilehisfour-toedbackfootpushesoff.Thefemalehe’slookingforwillhavelonger,narrowerfrontnailsandyellowish-browneyes.Perhapsshe’slookingforhim,too,butsinceboxturtlescan’tcalloutinthemannerofbirdsandwolves,theirmeetingmaybestrictlyamatterofchance.

Acolonyofmayappleplantsoffersfoodatjusttherightheightforaturtle.Althoughthefruitsaretoxicwhengreen,boxturtlesreadilyeattheripe

yellowfruitsandspreadtheseedsthroughtheforest.

Whatdoestheworldlooklikethroughtheeyeofamaleboxturtle?Femaleshaveyellowish-browneyes.

Manyboxturtlesliveformorethanahundredyears.Eachonehasdistinctivemarkingsonitsskinandshell.

Turtlesfrequentlyeatwildmushroomsandthushelpdistributethemicroscopicspores.

Snailsareanimportantpartoftheforestecosystembecausetheycancollectcalciumfromdeadleaves,storeitintheirshells,andpassitalong

tootheranimalsthatneedcalciumtoproduceeggs.Theleavesrustleastheturtletakesstepafterstep.Itmightbetimetopauseandnibbleonasmallblackgumfruitthathasfallentotheground,oraworm,oraslugthatlooksback.Orperhapsit’stimeforarest.Haveyoueverstoodstillintheforestandappreciatedthebeautyofthesunlightstreamingthroughtheleaves?Canaturtleappreciateasimilarsight?Anotheranimalyou’relikelytoseeintheforest,becauseit’ssolargeandabundant,isthewhite-taileddeer.Allanimalpopulationsgoincycles,andright

nowdeerareveryabundant,soabundantthatit’shardtobelievethatahundredyearsagotheywererareandendangered.TheywerecompletelygonefromIndiana,andonlytinypopulationsremainedinmosteasternstates.Extremehuntingpressureisthemostlikelyexplanationfortheirformerrarity.Alongwithhuntingdeer,humanswerealsohunting,trapping,andpoisoningwolvesandcougars—predatorsofthedeer.Aftermanyyearsofthis,notmuchlargewildliferemained.Inmanystates,includingIndiana,Illinois,Maryland,Pennsylvania,andTennessee,deerwereeventuallyreintroducedorrestocked.Butthepredatorswerenot.Nowtherearesomanydeerandthey’reeatingsomuchthatweworryaboutthelong-termeffectsontheforest.Ifthedeerhaveeatenalltheseedlings,whatwillhappenyearsfromnowwhenthebigoldtreesdietheirnaturaldeathsandnoyoungonesaregrowingtoreplacethem?Thechainofeffectsisnever-ending.Eventheturtlemaybeaffectedwhentheblackgum,sassafras,persimmon,anddogwoodfruitsnolongerraindowntowherehecangetthem.It’sgettingdark;theturtleseesamousenibblingonsomething.Humansmayspotamouseintheirhome,buttheyrarelyspotoneintheforest.Miceusuallycomeoutafterdusk,andwiththeirexcellentvisionandhearingtheymanagetostayhiddenwhenhumansareabout.Theturtleandthemousesharethebountyontheforestfloorbutworrythatcoyotesorwolvescouldshowupatanymoment.Coyoteseatfruit,too,buttheyprefereatinganimals.Onceboxturtlesgetoldenough,theirshellshardenandtheycanclosethemselvesinsideforprotection;butbeforetheageoftwotheirhingesdon’twork,andbeforetheageofseventheirshellsarestillsoftenoughforacoyotetobitethrough.Ifaturtlelivesbeyondsevenyearsandstaysintheforest—awayfromtheroadsandthemowingmachinery—ithasagoodchanceoflivingfiftymoreyears,andpossiblyahundred.Whatthoseeyeswillsee!

Theforestfloorisaveryfamiliarplaceforaboxturtle.Mostboxturtlesstayinaspecificeight-acrehomerange.Inthewintertheturtlewill

burrowafewinchesunderground.

Scentistheprimarywaythatwhite-taileddeercommunicatewithoneanother.Theirsenseofsmellishundredsoftimesbetterthanthatof

humans,andevenbetterthanadog’s.

Micearehighlysensitivetotheirenvironment.Theirlongwhiskerscontributetotheirkeensenseoftouch,andtheyalsohavewell-

developedsensesofsight,smell,andhearing.Althoughmanymiceinhabittheforest,you’reunlikelytoseethembecausetheyhidewhen

theyfeelandhearyourapproach.Atonetimewolvesandcougarsroamedtheeasternforests.Thesedays,withthosetoppredatorsalmostgone,nearlyeveryanimalintheforest—includinggrayfoxandbobcat—isafraidofcoyotes.Coyoteswillkillandeatthem.Inplaceswithmanycoyotes,veryfewgrayfoxesexist.Climbingtreesisanimportantmeansofescapeforbothfoxesandbobcats.Anyoftheselargeranimalswouldhappilyeatmice.Amousemayescapebyhidinginasmallholeatthebaseofatreeorbetweenrocks.Butitmustchoosecarefully,becauseasnakemightbehidinginanyofthoseplaces.Thetimberrattlesnakelooksthemostfrightening,butthecopperheadorthefriendlylookingblackratsnakewouldeatamouse,too.Inthissituationthecoyoteisafriendtothemouse,becausethecoyotealsoeatssnakes—evenrattlesnakes.Theonlycommonanimalsleftthatarenotafraidofcoyotesareadultbears.Bearsliketoeatberriesandseedsandinsectgrubs—andpeanutbutterwhensomehumaniscareless.

Historically,redwolveswerefoundinthebottomlandforestsandswampsofthesoutheasternUnitedStates.Theywerealmostdriventoextinctionbutarenowbeingreintroducedthroughcaptivebreedingprograms.

Grayfoxeshavehookedclawsthatenablethemtoclimbtrees.Thishelpsthemreachtreefruits,escapepredators,andgettodenning

hollowshighupintrees.

Thefemalebobcathasahomerangeofaboutfoursquaremiles.Shemarksherterritorywithscentsandscratches,andraisesheryoung

alone.

Therattlesnakeisadeadlypredatorofmice,butatleastitwarnsofitspresence.

Blackratsnakesareharmlesstohumans,buttheywouldhappilyeatamouse.Inwintertheymayhibernateinthesamedenwithrattlesnakes.Allofthiseating!Andwehavebarelytouchedonafewstrandsofthefoodweb.Theinteractionsareincrediblycomplexandvarybylocation,timeofday,andtimeofyear.Butatthebaseofeveryoneofthesestrandsofthefoodwebyou’llfindtheforestplants,becausetheyaretheonesabletopullenergyfromthesky.Andevenwhenalifehasended,thematterandenergypreviouslyheldinthatformcontinuestobepasseddowntheline—tothebacteriaandthefungi,theinsectsandthesoil,untilallthat’sleftaretheinvisibleatomsandmolecules,andfinallythosereturntothetreesthemselves.Thecyclecontinues,butthestoryendsdifferentlyeverytime.Treesarecapturingenergyandturningitintomatter.Andthematter—leaves,fruits,branches,wholetrunks—continuouslyrainsdownupontheground.Imagineifthousandsofyears’worthofthisorganicmatterstayedwhereitlanded.Noseedscouldreachthesoiltogerminate,itwouldbenearlyimpossibleforanimalstomoveacrosstheland,andfireswouldburnsohotthatnothingcouldsurvivethem.Fortunately,wehaveadifferentsortoffoodwebthatprocessesthedead.Itstartswiththebacteria.Bacteriaareeverywhereallthetime;theycoverourskin,theycovertheleaves,theycovertheground,theyexistinunimaginablediversity,andtheyarechangingallthetime.Whetheryou’reanimalorplant,therearebacteriathatharmandbacteriathathelp.

Primehabitatforblackbearsconsistsofaforestwithmaturehardwoodssuchasbeech,maple,birch,andevergreenspecies.Bearscanboth

climbtreesandswim.

Allstrandsofthefoodwebstartwithphotosyntheticorganisms.Plantsarethegreenfusethatlightsthefireoflifeandhandsenergydownthe

line.Bacterialcoveringsshiftthroughoutthedayandyear.Whensomething(orpart

ofsomething)dies,thebacterialcommunitycoveringitstartsshiftinginresponse.Theprocessesthatpreviouslykeptcertainbacteriaincheckarenolongerfunctioning.Alongwiththebacteria,microscopicfungalthreadsalsostartgrowingonthedetritus.Thedeadmatterbecomesamatrixforthesmalllivingcells.Andother,slightlylargerbutstilltinythingsfeedonthebacteriaandthefungi—likethoseamoebasandciliatesyoumayhaveseenunderthemicroscopeatschool.Andbacteriacanfeedonfungi,andfungionbacteria,andfungionfungi,andsoon.It’satinycomplexworlddowntherethatwehardlyknowanythingabout.Alltheselittlethingsbegintobreakdownthedeadmatterandmakeitmorenutritiousatthesametime.Thenalongcomessomethingthateatsthemallupatonce.Amillipedeismonstrousindimensioncomparedtothemicrobeswe’rediscussing.Amillipedecrawlsacrossthegroundmunchingentireleavesthatcontaincompletetinyecosystems.Andifanythingtriestostopit,themillipedespraysitwithpoisonouscyanide.Itsbrightcolorationiscommontocreatures,likebees,thatareboldenoughtowarnoftheirvenom.Ifyoupickupamillipedeandshakeitlooselybetweenyourhands,you’llsmellitsdefensivescent.Theodorisratherpleasant,butitmightbeagoodideatowashanyway.Millipedeswerethefirstlandanimals,andoneoftheearlyspecieswasmorethansixfeetlong.Liketheturtles,theyhaveseeneventhedinosaurscomeandgo.Today,onlyanimalsliketoadsareinterestedineatingmillipedes.Themillipede’sjobofchewingupfallenleaveshelpsspeeddecompositionalong.Andalthoughitgainssomenutrientsandcaloriesfromthisingestion(mostlyfromthemicrobesontheleaves),themillipede’sre-ingestionofitsownexcrementismoreimportant.Yousee,afterthemillipedeeatstheleaves,iteliminatestheminpelletform.Thepelletsaremoisterandhigherinnutrientsthantheoriginallyingestedleaves,sothey’reanevenbetterplaceforbacteriaandmicrobestoliveandmultiply.Bythetimeamillipedegetsaroundtore-ingestingthepellets,themealhasbecomeevenmorenutritious.Itmaysoundgrossuntilyouconsiderthatwehavebacteriahelpinguswithdigestionandnutrientrelease,too—itjusthappensinsideourlargeintestine.Manyotheranimals,fromruminantstobirds,usemultistagedigestion;itjustdoesn’tincludeanexternalstage.

Deadplantsarealsoapartofthefoodweb.Theslowdecompositionoftreetrunksprovidesimportanthabitatformanyspecies.Eventuallythe

woodbecomessoil.

Asfrighteningasthesecreaturesmayappear,themillipedesinourforestsareharmlesstohumans.

TheFowler’stoadeatsthemanyinsectsthatliveindecayingvegetation.Anothertinycreaturemovingacrosstheforestfloorthatisimportantinbreakingdownallthosefallenbranchesandtrunksisthetermite.Unlikeinourhomes,wheretermitesareanunwelcomepest,intheforesttermitesplayakeyroleinreleasingnutrientsfromwoodbackintothesoil.Butittakesavillagetoaccomplishthisimportanttask.Atermitecanbethoughtofasthevesselandvehicleforthehundredsoftinymicrobesinitsgutthatdotherealworkofwood

digestion.Termitesaresodependentonthesemicroscopicbacteriaandprotozoansthattheywouldstarvewithoutthem.Withinthegutthesetinybeingshavetheirowndivisionoflabor,andthey,too,wouldnotsurvivewithouteachother.Antsaretheenemyofbothmillipedesandtermites,soboththoseorganismshaveevolvedstrategiestodefendagainstthem.Onestudyfoundthatthemillipede’sdefensivechemicalmakesantsactcrazy,crosstheirantennaeabovetheirheads,andthenpassout.Termites,ontheotherhand,haveawholesocialclasswhosemainjobistocrushants.They’resospecializedforthistaskthatthey’renotevencapableoffeedingthemselves.Despitethissoldiercaste,antsarestillthenumberonecauseoftermitemortality.Salamanderscanbeofsomehelphere,sincetheyfrequentlyeatants.

Antsinacolonytendeggsthathaveenteredthepupalstage.InE.O.Wilson’sbookTheAnts,henotesthattheestimatedtenthousandtrillionindividualantsaliveatanyonetimeweighaboutasmuchasallhumanbeingscombined.Theirtightlywovensocialsystemisthekeytotheir

success.

Asix-spottedtigerbeetleroamstheforestfloorbyday,andatnightitreturnstothesameholewhereitsmotherlaidtheeggithatchedfrom.

Wolfspidershaveeighteyesarrangedinthreerows.Ifyoushineaflashlightintheforestatnight,youcanseeglowingreflectionsfromall

thespidereyes.Thelarvalphaseoftheiridescentgreentigerbeetleisanotherallyinthewaronants.Thefastandgracefuladultbeetlerunsacrosstheforestfloor.Afertilefemalemakesindividualholesinthesoilanddepositsoneeggineach.Whenanegghatches,itbecomesanunattractivewhitegrubwithafrighteningdarkhead.Thegrubstaysputinitsnatalholeandwaitsforantstocrawlby;then,springingitsheadfromthehole,itcatchestheminitsjaws.Afterayearofthislifestylethegrubpupatesandthenhatchesintoanadult.Imaginewakinguptobeafast,lovely,iridescentgreencreatureafterayearasanuglystay-at-hometroll!Buttheadultbeetledoesn’tforgetitsroots;everynight,afteradayofforaging,itreturnstothesameholewhereitwasborn.Itwillspendtheentirewinterthere,too.Itcanliveforuptofouryearsifatoadorawolfspiderdoesn’teatitbeforethen.Allofthisishappeningontheforestfloor,andwewalkbyitalmostasifweliveinanotherdimension.Alloftheseinteractions,theselivesintheforest,aredependentonthepresenceofthetreesandthestructure,matter,andenergytheyprovide.

Inbetweenthecanopyandthegroundisthemiddlerealm—thetreetrunksandthecreaturesthatusethemeitherashighwayorhome,or

both.

TheMiddleRealmTrunks

WHILEWALKINGTHROUGHAforestyoumightfocusonthegroundlookingforanimalsandwildflowers;oryoumightfocusonthespreadingbranchesinthecanopyandthebirdsthatmoveamongstthem;butmostoftenaforestexperienceisoneoftrunks,forthatistheforestathumaneyelevel.Youmightfirstregisterthetrunksize:large,medium,small;andafterthatthepatternsofthebark:smooth,striped,shaggy.Takeenoughforestwalksandyoumaystartwonderingwhichspeciesisassociatedwitheachpattern.Manybotanicalguidebooksuseleafshapeforidentification—butthoseleavesaresofarawayfromwhereyoustandthatyoucanbarelytellifthey’repointedorlobed.Thebark,however,isrightwhereyoucantouchitandsmellit.

Athree-trunkedtulippoplarrisesfromthegroundtothesky.Tosome,thestudyofbarkpatternsmightsoundlikeaboringhobby,butonceyougetintoityoudiscoveramazeofgreatcomplexity.Oneavenueofthismazeisthatthebarkpatternsonatreechangeovertime.Theyoungoneslookverydifferentfromtheoldones.Andtheoldertheyget,themoreindividualtheybecome,justlikeyouragingrelatives.Twotreesofthesamespeciesandthesameagesidebysidemaylookverydifferent.

Andbeyondthestudyofbarkasameansofidentificationisthepleasuretobegainedinthesheeraestheticsofbark.Ithasasinglefunction—protectingthegrowinglayerbeneathit—butsomanydifferentwaysofaccomplishingthattask.

Acloselookatbarkrevealsthewondrousdiversityofpatterns.

Sometrees,likethisshagbarkhickory,getshaggierastheygetolder.Thesebarkflapsaregoodplacesforsmallbatstosleepawaythe

daylighthours.

Awholeworldofinteractionsmayoccurbeneathatree’sbark.Thebeetlelarvaewhomadethesetunnelsarelonggone,buttheirwork

remains.Underthisinterestingbrowncoveringyou’llsometimesfindanotherworld—alovely,complexworldcreatedbyinsectsthatlivebyeatingthetenderwoodbeneaththebark.Theinsectsmaybelonggone,butthesignstheyleavebehindindicatethatanintelligentsocietyonceoccupiedthisterritory,muchliketheabandonedcliffdwellingsoftheAnasaziIndians.Thetinycreaturesatworkherearebarkbeetlelarvae.Therearemanydifferentkindsofthesebeetles.Anexaminationofthepatterns,alongwithknowingthetreespeciesthey’refoundon,helpdeterminewhichparticularspeciesmadethedesigns.Insomespecies,themalebeetlewillgnawhiswaythroughthebarkandashortdistanceintothewoodtocreateaspaceforhis“ladyfriends.”Guidedbyhisscent,betweenoneandfivefemaleswillfindtheirwaythroughthepassagehemadeandintohisbachelorpad.Aftertheymatewiththemalebeetle,thefemalesgotheirseparatewaysandbeginchewingtheirelongatednurseries.Inotherspeciesofbarkbeetles,it’sthematedfemaleswhotunnelthroughthebarkandintothetendersapwood.Nomatterhowthefirstpartofthestorygoes,thesecondpartisalwaysthesame:thefemalechewsasmallseparatespaceforeach

ofhereggsandthenlaysthem,atwhichpointherworkisdone.Theeggseventuallyhatchintopalewormlikelarvae.Theselarvaeknowtheworldonlyasadarkplacethatsmellsoffreshwood.Theydon’thavetothinkaboutwhattodonext,forthey’regeneticallyprogrammedtostartchewinginaparticularpattern.SomespecieschewinwanderingS-shapedpatterns,andsomeheadinarightangleawayfromthemainnurserychamber.Asthelarvaslowlyeatsitswayawayfromhome,itgrowslargerandlarger.Justaspencilmarksonafarmhousewallindicatethegrowthofitsyoungresidents,thesizeofanindividualtunnelrevealsthegrowthhistoryofthelarva.Thenonedaythelarvadoesn’tfeellikechewingwoodinalineanymore,asithasbeendoingallyear;insteaditchewsasmallcavityandjustliesthere.Itsskinbeginstogetdryandtight.Itturnsbrown.Ifalarvathinks,itmightthinkit’sdying.Butthenspringarrives,thebrownshellbursts,andithasbecomeawingedinsect.Itstartschewingatarightangleonceagain—butthistimetheturntakesittowardtheouterbark.Afterafinalmouthfulofbarkourcreatureseessunlightforthefirsttime—anditswingsareusedforthefirsttime,too.Inthetreetrunkisasmallholewherethebeetleemerged,andsomewhereintheforestisalittlebeetlelookingforamate.Thetreesaren’tharmedmuchbythesmallin-and-outholes;mostofthedamageisdonebythelarvae-createdtunnelsthatcutacrossthevesselsusedtotransportfluids.Thiscankillatree,althoughthat’snotthelarvae’sobjective.Theemeraldashborerandthesouthernpinebarkbeetlearetwonotoriouskillers.Sometimestheadultbeetlescarryinfungusspores,andthefungusmaydomoreharmthanthelarvae.Intheend,barkbeetlesarebadnewsforindividualtrees,butthey’repartofthegreaterforestecosystem.Bybringingstressedtreestotheground,thebeetlescreatehomesandmealsforamyriadofotherorganisms,includingfungi,birds,reptiles,amphibians,andmammals.Andtheybringlighttothosesmallplantsontheforestfloorawaitingtheirturninthesun.Withallthistalkoftreetrunks,let’snotignoretheanimalswhousethesetrunksasverticalhighways.Awholesuiteofforestorganismsdon’tlivejustontheground,orjustinthecanopy,butinsteadmovebetweenthesetwoworlds—betweenheavenandearth.Squirrelisthefirstonewhocomestomind.

Viewingtheselinesandshapesmadebybeetlelarvaeatamuchlarger

scale,youmightwonderabouttheintelligentbeingswhomadethem.Squirrelsnestupinthecanopyinsmallgroupsofrelatedfemales,orjustsinglemalesontheirown,buttofindandstorefoodasquirrelmustmoveupanddownthetrunk.Squirrelseatjustaboutanything,includingbudsandbark,butseedsthatcanbestoredintheground,suchasacorns,beechnuts,andhickorynuts—allgiftsfromthetrees—aretheirmostimportantfood.Onaharvestingmission,asquirrelwillcollectasmallnumberofseedsandhidethemtogetherunderleavesorintheground.Thenitwillgotocollectmoreseedsandhidethemsomewhereelse,andsoon.Itmaycreatethousandsofthesesmallcaches.Asquirrelrememberswhereithidalloftheseeds,andwithintwodaysitwillbebacktoeattheseedsormovethemtolonger-termstorage.Ifthey’reseedsthatmightgerminatesoon,likethewhiteoakacorn,thesquirrelwillnipouttheembryo.Inthecourseofthisforaging,squirrelsfindotherthingstoeatthatwon’tkeepunderground.Thesetreats,suchasdogwoodberriesorcicadas,theywilleatrightawaywhileperchedinthecanopy.Asquirrelhasbigpadsonthesidesofitsthumbs,anditholdsitsfoodbetweenthesepadswhileiteats.Thisgivestheimpressionthatit’spraying.Thinkinglikeatree,itmayatfirstseemthatthesquirrelisapredator—eatingthetree’soffspring.Butifsomethingweretohappentothesquirrel(coyote!),thenithasdonethetreeafavorbytuckingseedsintothegroundoutfromunderthecanopy.Soasquirrelisneitherallgoodnorallbadforatree.Whileallofthiscollectingandburyingisgoingon,anotherdramaisunfolding,forsquirrelsliveinaworldwithadominancehierarchy.Theoldermalesarethemostdominant,andwhenayoungermaleapproacheswithouttheproperattitudeofsubmissionthedominantsquirrelgetsaggressive.Thisisallcommunicatedwiththetail.Thetailcanbecontrolledtobendjustatthetip,tobendnearthemiddle,ortobeheldstraight(eitherupbythebackoroutflat).Atailfoldedoverinthemiddleorlowerindicatessubordination,andthissquirrelisunlikelytobechased;butifoursquirrelholdshistailstraightup,thedominantmalewillchasehim.Femalesquirrels,ontheotherhand,havetheirownsocialhierarchies.Andinbothsexes,friendlinessexpressedasgroomingorplayingisjustasimportantasaggression.Afemaleisfertileonlytwiceayear,forlessthanadayeachtime,anddozensofmalesmayapproachheronthatday.Matingusuallyoccurshighinthecanopy.Adominantmalewillaggressivelydefendhisaccesstoher,butsheoftenrunsfromhim;asaresultoneoftheyoungmaleswaitinglowerinthecanopymaygetanunexpectedopportunitytomatewithherinstead.

Dog-daycicadasdonoteatatallasadults,buttheymaybeeatenbysquirrels.Thenymphofthisspeciesspendsonlyayearunderground,not

thirteenorseventeenyearslikethered-eyedperiodicalcicadas.

Graysquirrelscancommunicatewiththepositionandactionoftheirtails.Asquirrelalsousesitstailinresponsetothreatsfrompredators.Anapproachingcatwillelicitawildwavingofthetail.Interestingly,anaerialpredator,likeahawk,willnotcausetailwavingbutwillinsteadcauseoursquirreltocallout“quaa,”andalltheothersquirrelswillstopeatingorburyingfoodandlookup.Consideringhowcloselysquirrelslivewithhumans,it’ssurprisingthatscientistslearnedthesethingsaboutsquirrelsjustafewdecadesago.Ithasbeenobservedthatwhensquirrelpopulationsgetsuperabundant,generallyintheearlyfall,thesquirrelsmayswarm—largegroupsmovelongdistancesacrossthelandscapetogetherandevenswimacrossbodiesofwater.These

swarmingeventsweremorecommoninthepastthantheyarenow.Anumberofotheranimalssharethishighwayofbarkwithsquirrels.Oneisapredatorthatapproachesslowlyandsilently:theredcornsnake.Thesnakemightmakedinnerofanestofbabysquirrels.Thismayseemunfairuntilyouconsiderthatadultsquirrelssometimeseatbabybirds,andakingsnakewilleatacornsnake.Roundandroundgoeslifeintheforest.Andwestepbackandseethatsomealwayssurvive—somesquirrels,somebirds,sometrees,somesnakes,someturtles.Besidesthebarkborers,multitudesofotherinsectscallthetrunkhomeanddon’tkillthetrees.Manyoftheseinsectsareeatenbytreefrogs,whospendmostoftheirtimeinthecanopyandthenmovedownthetrunkwhenit’stimetobreed.Treefrogsareprimarilycanopydwellers,buttheyneedapoolofwaterfortheireggs.Matingandegglayingaredoneinthesamepoolswherethesalamandersbreed.Sincetreefrogscan’tfly,orevenleapthatfar,theymustmaketheirwaydownthetrunkandbackup.They’revulnerabletopredatorsduringthistime,butsomeprotectionisofferedbythefactthattheirskincanchangecolorstotakeonthehueandpatternofthesurfacetheysitupon.Howanimalschangecolorstomatchthesurroundingenvironmentisstillbeingresearched.Itprimarilyhappensthroughtheactionofspecializedstar-shapedcellsintheskinthatcontainsmallbubblesofpigment.Whenthebubblesallmovetothesamespotinthecenterofthecell,theskinappearsonecolor;whenthebubblesspreadthroughoutthecell,theskinchangestoadifferentcolor.Infrogs,thesecellsaretriggeredtomovethebubblesbyhormonalchemicalscirculatingthroughthebloodstream.Butwhatcuesthefrogtomakethesehormones?Lightreflectancereceivedthroughtheeyes?Therearestillmanymysteriesintheforest.

Althoughmanyhumanshaveafearofsnakes,theredcornsnakeisaverygentlecreature.

Thegraytreefrogcanchangecolortomatchitssurroundings.

SheddingandRestingAutumnandWinter

ALTHOUGHWETENDTOthinkofindividualtreesasunchangingovertime,considerthattheleaf-droppingspeciesgetanewcoveringeveryyear.Itwouldbelikeallyourhairandskindroppingoffeachyearandthengrowingbackmonthslater.Soundsextreme,doesn’tit?Whywouldtreesdothat?Thereareanumberofreasons;oneisthatitcanbeawayofsheddinginsectsanddiseasesandmakingafreshstart.Somenaggingproblemsarebettertojustridoneselfofinsteadoftryingtofix.Imagineyouhadacontagiousskindiseaseandaninsectinfestationbeforeyour“bigdrop.”Youmightbefreeoftheseproblemsafterthegrowingback—atleastuntilyouwereexposedagain.Thissheddingcouldbenefitevergreentrees,too,sowhydon’ttheyshedtheirleaves?Actuallytheydoshed,justnotallatonceinsuchadramaticway.Pineskeeptheirneedlesforanumberofyearsbeforetheyshed,sosomethinggreenisalwaysleftonthetreewhentheoldneedlesaredropping.Wetendtonoticethegreenonthetreemorethanthebrownneedlesontheground.Thenumberofyearspinesholdtheirneedlesbeforedroppingthemvariesbyspecies:whitepinesfortwoorthreeyears,somewesternbristleconepinesformorethanthirtyyears.

Americanbeechisoneofthefewdeciduousspeciesthatholdsomeleavesuntilspring,asillustratedbythisbeechforestinwinter.This

featuremaydeterdeerfrombrowsingthetwigs.Anotherreasonbroad-leavedtreesshedisthattheleavesarethemostdelicatepartofthetree,andthey’resusceptibletobeingdestroyedbyahardfreeze.Leafcellscouldswellandburst,ordryout,especiallyifthegroundisfrozenandtherootsareunabletotakeinwater.Andwhenitsnows,leavesonatreecollectthesnow,andthecombinedweightofsomanysnow-ladenleavescouldcausemajorbranchestocrackandfall.Bettertodropyourleavesthanyourbranches!Ifleavesheldonthroughthecoldmonths,theywouldactlikelittlesailswhenwinterwindswhippedthrough.Baretreesletthewindssliprightby.Windandhailmightalsotattertheleaves.Whenthespringsunshinereturned,theleaveswouldbealmostuseless.No,bettertoshedthembeforewinterandstartfreshnextyear.Pinesandmanyotherevergreenspecieshavedevelopedleavesspecializedfordealingwiththosewinterchallenges.Highwinds,hail,andsnow?Verynarrowleaves,likeneedles,allowthewindtosliprightbyandthesnowtoslideright

off.Frozenground?Awaxylayerovertheleafkeepsitfromdryingout.Munchinginsectsandanimals?Sharp,toughleavesdeterthem.Buttherearealwayscompromises,andthesesmall,protectedneedlescan’tcollectasmuchsunlightasthebroadleavesduringthewarmgrowingseason.Theymakeupforthatlackbybeingabletocapturesunlightyear-round.Alookacrosstheglobetellsusthattheevergreentechniqueworksbetterwherewinterislonger(thinkborealforest)andthebroadleaftechniquewherewinteriswarmer(thinktropicalforest).Ourtemperateforestliesbetweenthesetwoextremes,andtreessuccessfullyexhibitbothlifestyles.Andsotheleavesfall.Thereasonssoundutilitarian,buttheresultissoverybeautiful.Wecouldimaginethatthecreationofbeautyistherealreasonforleavesturninganddropping—becausetheleavesdon’tjustdieanddropbutinsteadarepartofacarefullycontrolledprocess,anactiveprocess.Thetreelooksatthesun’sclock,feelsachillintheair,andsensesit’stimetokisstheleafgoodbye.It’salonggoodbye,lastinganaverageoftwentydaysfromthefirstsignsofcoloruntiltheleaffalls.Thetreeaccomplishesthiswithchemicalmessagesthatdirecttheleafcellstostopmakingthegreenpigmentchlorophyll,andtoshredandrecycleanyoldmoleculesofthissubstance.Asaresult,leavesgofromgreentoyellow.Eventuallytheseleavesgofromyellowtobrown,sincethey’renolongerphotosynthesizingandmakingtheenergytheyneedtorepairagingcells.Sometreespecies,likesycamore,beech,redbud,andtulippoplar,neverhavereddishfallcolor.Theyturnyellow,andthat’sit.Thismightseemlikeanunfortunatelylimitedpaletteuntilthedayyouseethosegoldentreesabsolutelyglowingintheafternoonlight.It’sasifthesunwereshiningbackupfromtheground.

Agedhumansgetblotchy,andsodoagedleaves.Insometreespecies,redpigmentisproducedinthefallbyhealthycells.Theblackandgreen

cellsherearediseased.Thisleafwillsoonbeshed.

Wecanwonderwhichinsectsorbacteriaorfungusgottothisleaf,orwecanjustadmiretheartisticresultstheyhavecreated.

Anoakleafhasjustfallenfromthetree.Somepeopleconsideritgoodluckifyoucancatchaleafonitswaytotheground.

Thenthereareotherspecies,likeblackgum,thatneverhaveyellowleaves.Theygofromgreentoadeepburgundyred,andleavesdropfromthetreewhilestillred.Theredcolorationinfallleavesisnotsomethingthatwasthereallalongandjustcoveredupbythegreenpigments,liketheyellow;no,theredpigmentsareactivelyproducedbyleavesduringthecold,crispdaysofautumn.The

pigmentsareakindofself-manufacturedsunshade.Treesneedthesunshadebecauseoncold,brightdaystheirleavesarestillproducingoxygenthroughphotosynthesis,butthecell’snormalprocessfordealingwiththeveryreactiveoxygenmoleculeissloweddownduetothechillytemperatures.Theredsunshadepreventstoomuchoxygenfrombeingproduced—likeabeachumbrellaprotectingfromsunburn.Ourfavorites,ofcourse,arethosetreeswhoseleavesshoweveryimaginableshadeofgreen,yellow,orange,andred.Likesnowflakes,notwoarealikeandeachoneseemsmoreremarkablethantheonebefore.Themaplesandthesweetgumsaretheverybestatthissortofshow.Themoresunaleafgets,themoreitwillneedtheredpigments.Sotreesgrowninfullsunhaveleaveswiththemostremarkablehues.

Eachindividualcellinthiscolorfuloakleafmakesitsowndecisionsaboutwhetherthegreenchlorophyllgoesorstaysandwhetherthered

pigmentsshouldbeproduced.

Differenttreespeciesturndifferentcolorsinthefall.Somespeciesturnonlyyellowwhileotherspeciesturnonlyred.Others,likeredmapleand

sweetgum,turnamultitudeofcolors.

Adiversityofcolorsindicatesadiversityofspecies.Thedeepgreencolorisfromconiferssuchaspineandhemlock.

PeoplefromotherregionstraveltotheeasternUnitedStatesjusttoseethisshowofautumncolor.Itmightbelistedasoneofthewondersofthe

worldwereitnotsoephemeral.Becauseofthesedifferencesinfallcolorationfromspeciestospecies,itispossibletotellfromadistancethegeneralcompositionofaforest—evenwhiledrivingbyorviewingfromanoverlook.Assoiltypeschangeacrossthelandscape,sodospecies,andsodothecolors.Onehuegracesthenorthsideandanotherthesouthsideoftheslope.Differentspecies,differentcolors,atthebottomofthedrainagecomparedtothetop—anamazingever-changingartpieceofferedforfree.Onlyforestscontainingbroad-leavedtreesinplaceswithcoldwintersandhotsummerswillproduceaspectacularautumnshow.Thesearethetemperatedeciduousforests,andtheyoccuronlyineasternNorthAmerica,partsofEurope,andpartsofeasternAsia.Ofthesethreeplaces,theshowintheeasternUnitedStatesisconsideredthebestontheplanet.Consideryourselfluckyifyou’reintherightplaceattherighttimetowitnessit.Pausetoadmirenature’sbeautyandhowitresonatesdeeplywithinyou.Anddonotmakethemistakeofjustwitnessingfromafar.Enterthisworldoftangiblerainbowsandfeelthecolorsaround,above,andbeneathyou.Breathetherainbowair.Smellthefungifeastingonthefreshleaves.Thebiologicalprocessesthatcauseleafcolorchangesaredifferentfromtheprocessthatcausesleavestofall.Toinitiatetheactualfalling,aspeciallayerofcellsatthebaseoftheleaf’sstemmakesalayerofscartissueonthetwigsideanddissolvesthecellsontheleafside.Abitofwindmaymakethedifferenceinwhenanindividualleaffalls,butevenoncompletelywindlessdaysinautumnonecansitandenjoytherainofleavesthatmayseemcompletelyrandombutcontainsitsownimpulse.AlbertEinsteinsaid,“WhatIseeinNatureisamagnificentstructurethatwecancomprehendonlyveryimperfectly,andthatmustfillathinkingpersonwithafeelingofhumility.Thisisagenuinelyreligiousfeelingthathasnothingtodowithmysticism.”Leaf-dropreligion?Whynot.Whentheairhasachill,andthesunnyskyisaperfectblue,andthecoloroftheleavesagainstitmakesyourheartbeatalittledifferently,youmayreadilyconvert.Fallenleavesmeanlife,notdeath,toamyriadoforganisms.Thelivesoftinysoilinsects,andbacteriabeyondcount,dependonthisannualbounty.Eventhelargercreaturesthatdon’tfeedontheleavesdependonthismoistfoliarblankettoprotecttheirdelicateskinandhidethemfrompredators.Ifallthiswerenotenoughofatreasure,inafewyearstheleaveswillbecomesoil—thesubstancethatisourtruestwealth.

Whenyoupayattentiontotheseseasonalchangesintrees,you’llnoticethatsomeoftheleavesdon’tfallinfall.Instead,somebrownandhangonthroughthewinter,droppingonlywhenthebudsofspringswellandpushthemaside.Insomespecies,suchasbeech,oak,andhornbeam,thisisanormaloccurrence,butwereallydon’tknowwhy.Afewscientistshavespeculatedthatbrowsinganimalsdon’tlikeeatingdeadleavessothey’llnibblebarebranchesinstead,givingthefitnessadvantagetobranchesthatholdtheirleaves,butthere’sverylittledatatosupportthis.Althoughevolutioncreatedallformsandprocesses,notallformsandprocesseshaveanevolutionaryreason.Somethingsarejustby-productsofevolutionaryhistory.Wecanappreciatetheirbeautyevenwithoutknowingthereasonforit.Onoccasion,leavesthatnormallydropoffinthefallmaywitheranddiebeforetheyhavebuiltthespeciallayerofcellsthatwillseparatethemfromthetwig.Atemporaryproblemsuchasanearlyhardfrostmaycausethis,oritmaybeasignthattheentiretreeisdying.It’sinterestinghowwefeelthe“notrightness”ofleaveswhosenormalprocesshasbeeninterruptedwhencomparedtoleavesthathavebeenthroughtheirnormalseasonalprocess.

Thisleaf,inmidfall,isonitswaytobecomingsoil.

Thesmellofdampfallleavescomesfromdecaythat’salreadybeginning.Formanyofus,thatsmellevokespleasantmemories.

Redmapleleavescreateabeautifulbutephemeralcarpetontheforestfloor.Formanysmallorganisms,theyarebothalayerofprotectionanda

sourceoffood.

Inthemountainsinwinter,thegroundiscoveredwithalayerofsnow,andthebaretreesemergefromitlikefur.

Allthatcolor!Butthatdoesn’ttellthefullstory,becauseforagoodportionoftheyeartheforestisn’tgreenandbrownbutwhiteandbrown.Thetreeflowershavecomeandgone,theseedshavecomeandgone,theleaveshavecomeandgone,nextyear’sbudshavebeenmade,andnowit’stimetowait.Lifegoesonintheforest,butit’stuckedaway,sloweddown,suspended.Thetenderplantsontheforestfloorhavediedbacktotheirroots.Treesstandwaitingforthetriggersthattelltheirbudstoopen.Turtlesandfrogsareburieddowninthemud.Batsandbearsandskunksaresleeping.Thedelicateinsectshavebeenkilledbythecold,buteggsforthenextgenerationarewaiting.Noonehasresearchedthephysicalandpsychologicaleffectsofhavingourworldblazewithcolorforashorttimeeachyear,orgoallwhitewithsnowovernightinthewinter,butwecelebratetheseeventsanyway.Andwaitforspringtoarriveagain.

Sometimeswhentheairishumidandthetemperatureisbelowfreezing,themoistureintheairwillfreezeonsurfacesandcreatecrystalsofhoarfrost.Whenalltheseclearcrystalsdiffractthelightindifferent

directions,theresultisaworldofwhite.

Aspecialquietnessinhabitstheforestnow.Notonlyhavethetrees

suspendedtheirnormalactivities,plantsandanimalsofallkindshavealsopaused.Mostofthebirdshaveleft.

Whenthesuncomesoutafterasnowfall,theworldfeelstransformed.Asitmelts,thesnowwillseepintotheforestsoilandprovidewaterfortrees,

streams,andvernalpools.

Redmaplesproducetwotypesofflowers:maleandfemale.Thefemaleflowersstayonthetreeandproducethewingedseedsweareallfamiliarwith;themaleflowersdroptothegroundaftertheirjobofpollenrelease

isdone.

BuddingWinterintoSpring

INMANYPARTSOFtheworld,theforestseasonsaremeasuredbymoisture:wetordry.Inthetemperateforest,however,seasonsaremeasuredbytemperature:warmorcold.Duringthewarmseason,theleavesareoutandactivelyphotosynthesizing;thesugarsproducedinthisprocesspoweractivecelldivisionandgrowth.Thetreesandshrubstakeadvantageofthiswarm-seasongrowthspurtbypreparingfornextyear’sflowersandleaves,evenasthisyear’sleavesarestillgrowing.Ifyoulookcarefullyatthebaseofaleaf,you’llseeabumpjustaboveitonthetwigwhereit’sattached;that’sthebudfornextyear.Whiletheleavesareoutthesebudsarenotverynoticeable,butinthewinteraftertheleavesfalltheybecomemoreobvious.Imaginethatyou’reinanairportwatchingtheluggagegoaroundonthebaggageclaimcarousel.Althoughthepiecesofluggagedifferinsizeandcolor,theyalsohavesimilarities.Manyofthepiecesofluggageareroughlyrectangularsuitcases,whileothersarecylinder-shapedduffelbags.Youhavenowayofknowingforsurewhat’sineachbag,butyoucanguessthatmostofthemcontainclothesandtoiletries.It’sthesamewaywithtreebuds:similaritiesanddifferences.Thesizeandshapeandhairinessofthesebudscanhelpyouidentifythetreespeciesevenwithoutleaves.

Wavesofpastelcolorsspreadacrossthelandscapeinearlyspring.Thecolorsarefromtreeflowersandnewleavesjustemergingfrombuds.

Eachspecieswilladdadistinctivehue.Justastherearetwocommonkindsofluggage,suitcasesandduffelbags,soaretheretwocommontypesofbuds:thosecoveredinbudscalesarrangedlikeshinglesonaroofandthosewithbudscalesarrangedinpairsclaspingeachother.Cherry,oak,maple,andsweetgumallhavetheshingletypeofbuds.Ifyoucutoneofthosebudsopeninthefallorwinterandhadagoodmicroscopetoexamineit,youwouldseethatnextyear’sleaves,andsometimesflowers,are

alreadytherewaitingfortheseasontochange.Inspringtime,thelongerdaysandwarmertemperaturesinitiatebudbreak.Thebudscalesarepushedopenbythetinyleavescontainedwithin.Fromadistance,abarelydiscernibledifferenceincolorcomesovertheforestcanopy.Thegray-brownturnstosomethingelse,somethingwhitishorpinkish,somethingthatdefinitivelysaysspringtothosewhoeagerlyawaitsuchsigns.Andwhenyougowalking,althoughtheairisstillcool,youseetinyleavesunfurledandreadyfortheirturninthesun.

Budsarevisibleatthebaseofeachleaf.Theyarelikesuitcasespackedforajourney.

Ifyoulookcarefullyatthissweetgumtwig,youcanseetheovalscarwherelastyear’sleafwasattached.Insidetheovalscararethreesmallerroundbumpswherethevesselscarryingfoodandwaterwentfromthetwigtotheleaf.Theshapeoftheleafscarscanhelpidentifythetree

species.

Thebudsareapparentonatreeaftertheleaveshavefallen—somepointy,somerounded,somehairy,somenot.Packedtightlyinsideeach

budareleavesfornextseason.

Thebudsfordogwood’sspringflowersarealreadynoticeableinthefall.Thebudsthatcontainnextyear’sflowersoftenlookdifferentfromthosecontainingleaves.Thedogwoodisastrikingexampleofthis.Thefat,roundbudsattheendofatwigaregoingtoopentorevealnextyear’sflowers.Theleafbudsaremuchsmaller.Ifyou’reabettingperson,youmightbetsomeoneinmidwinterthatyouknowhowmanyflowersaregoingtobeonthatdogwoodtreeovertherecomespring.Countingthefatbudsisrelativelyeasy,andeachfatbudcorrespondstooneflower,right?Well,maybenot.Whenspringfinallycomestheclaspingbudscalesbreakapart,butinsteadofdroppingtotheground,astheydoinmostotherspecies,thedogwoodbudscalesopenandenlargeandturnpinkorwhite(dependingonthevariety).Insidetheseexpandedbudscalesisaclusteroftinyflowers—aboutthirtyperbud.Soifyoubetthatonebranchwiththreefatbudswasgoingtoproducethreeflowers,youwouldlose.Therewouldactuallybeninetyflowers.Andeachoftheseflowersmightbepollinatedandbecometheredberriesweassociatewithdogwoodtrees.Squirrelsconsiderthemadelicacy.

Whiteoakleavesunfurlafterbeingpackedinthebudallwinter.The

hairinessoftheyoungleavescanmakethetreeslooksilverywhiteatsomeangles.

Frombudtofruit,thedogwoodmakesitsannualjourneythroughthegrowingseason.Thefruitsareeatenbysquirrelsandreportedlytaste

badtohumans.

Tulippoplarbudshavedelicatepairedbudcoverings.Likehandsinprayer,eachpairopenstorevealatinyfoldedleaf.Atlast,inthecenter,

istheflowerbud.Tulippoplarsareamongourmostancienttreespecies,andtheyhaveretainedalovelyyetunusualwayofexposingtheirnewleavesinthespring.Theclaspingbudscalespeelawaytorevealtwotenderleaflikestructuresatthebaseofeachnewleaf.Thesemini-leaves,calledstipules,openandfoldbackasthedelicatetrueleafemerges.Firstoneleaf,thentwo,thenfour,andthestipulesfanoutandfinallyexposeaflowerbudinthecenter.Andassoonastheseleavesarefullyexpandedandfeedingthetree,theystartbuildingbudsattheirbasefornextyear’sshow.Anyonewhohasspentmuchtimelookingatforestsknowsthatthisprogressionfollowsacertainorder.Budbreakisprimarilytriggeredbyanaccumulationofdaysoveracertaintemperature,butnotalltreesopentheirbudsatthesametime.IntheeasternUnitedStates,themaplesarethefirst,floweringinFebruaryorMarch.Fromadistanceyoumaynoticethatthetopsofthetreesaresuddenlytingedpink,evenifyoudon’tknowthatthereasonistheflowersoftheredmaple.Orperhapswhereyoulivetheglowismoreofachartreusecolor—thatwouldbethesugarmaplesortulippoplars.Itdoesn’tmatterwhatthecalendar

says,biologistscallthatspring.

Thelargepinkstipulesonthishickorytreepeelbackasentirenewbranchesemerge.Thestipuleseventuallydroptotheforestfloorandcancreatepuzzlementinforestvisitorsfarbelow.Whyaresmallpinkleaves

droppinginthespringtime?

Budbreakisthefirstblushofspring.Redmapleflowersgivethetreetopsapinkhuewhileotherspeciesaddyellowsandgreens.Theoakleaves

won’temergeuntillater.

Springcomeslatertothehighaltitudes.Thefarthersouthyouaretheearlierthishappens,butthetimingalsodependsuponaltitude.Low-elevationtreesleafoutearlier.Springisdelayedthreedaysforevery330-footincreaseinelevation.Infalltheprocessisreversed:high-

elevationtreesshowcolorandlosetheirleavesfirst.Soifyouliveinmountainousterrain,youcanwatchtheseasonssweepupanddownthehillsidesinwaves.Andwithinthewavesarethesmallerripplescreatedbythenextspeciestobreakbud,orthenextspeciestocolor.Summerislongerinthelowelevations.Butsummerisgettinglongereverywhere.IntheeasternUnitedStates,springnowcomesoneweekearlierthanitdidin1976.Youdon’thavetoaskwhy;youalreadyknow.Budbreakisadvancingbyoneortwodaysadecade,whichdoesn’tsoundlikemuchuntilyoucountyourownpersonaldecadesandmultiply.Sospringisgettingalittleearliereveryyear,astheresearchshows,andfallishappeningalittlelater.Summerisgettinglonger,andthetreesaregrowingfaster.Thisisnotspeculation.Thisishappening.

InthiswildbluebellmeadowinVirginia,treeleavesarejustemerging;whentheleavesfinishexpandingandshadetheforestfloor,thebluebell

plantswilldisappearuntilnextspring.

FlowersandFruitsSpringintoSummer

SOILITSELFMAYNOTappearbeautifultous,butspringingfromthesoilandwhollydependentonitaremanythingswelabelasbeautiful.Themostobviousarethewildflowers.Thesesmallplantshaveneverevolvedthestifftrunksthatenabletreestogrowtallandcompeteforsunlight.Thehumblewildflowersmuststayclosetothegroundandhopeforthebest.Andthebestdoescome,whentemperatureswarmintheearlyspringbutthetreeleaveshaven’texpandedtotheirfullsize.Itisthenthatlife-givingsunlightreachesallthewaytotheforestfloor,andthesmallestplantsgettheirshare.Theflowersassociatedwithspringcomeandgojustasquicklyastheseason,sowecallthemephemerals.Someindividualflowerslastonlyoneday,andanentireplantmaycompleteitsreproductivechoresinjustamonthortwo—thoughthememoryofseeingthoseephemeralbeautiesmaystaywithusforever.

Virginiabluebellflowerschangecolorastheyage,sendingasignaltotheirbutterflypollinatorsabouthowmuchnectartheymightcontain.

Wildgeranium,aperennialwildflower,spreadsbyundergroundrhizomes.Eachflowercanproduceamaximumofonlyfiveseeds.

Springwalksintheforestcandelightvisitorswithsplashesofcolorfromperennialwildflowers,plantedandtendedbynoone.

Bluets,lady’sslipper,purplefringedorchid,bloodroot,showyorchid,trillium

Althoughflowernectarisanimportantfoodsourceforhummingbirds,themajorityoftheirnutritioncomesfromeatinginsects,suchastinyspiders.

Polleniscarriedfromplanttoplantonbeakandheadfeathers.

Tinyflowersembeddedinthecentralspatheofthejack-in-the-pulpithaveanodorthatattractssmallfliestopollinatethem.

Outofthedarkground,whichseemedfrozenandlifelessjustafewweeksago,greenshootsandflowerbudsappear.Therootsandrhizomesofperennialplants

werewaiting,alive,underground,forjusttherightcombinationofcuesfromtheenvironmenttosignalsprouting.Allindividualsofthesamespeciesrespondtothesamecues,soseeminglyallatoncetheirflowersappearinnumerousplacesacrossthelandscape.Nooneplantedortendedthesewildflowersthatmayhaveundergroundpartsasoldasthetreesnearby.Anexceptiontoallofthiscolorandsweetscentistheinterestingjack-in-the-pulpitflower.Smallfliesareitspollinators,andtheyareattractedtothenot-so-pleasantodorofthetinyflowersembeddedinthecentralspathe.Therootsofanindividualplantmayliveformorethantwenty-fiveyears.Flowerhuesarebeyondanythingtheleavesarecapableof.Thesearenotthecolorsofdegenerationorprotectionthatweseeinfallleaves;thesearethecolorsofattraction,forifflowerswanttomatewitheachotherintheheadyspringwarmththeymustenlistapollinator.Andpollinatorslovecolorandscentandsweetness,sotheflowersofferallofthat.Weloveflowersforthesamereasonspollinatorsdo.Afterthespringwildflowers,thefloweringshrubstaketheirturn.Theywantthepollinatorstobeabletofindthemeasily,too.Azalea,rhododendron,andmountainlaurelalladdcolortotheforest.Andtheireyelashlikepollenwandsarejustwaitingfortherightvisitor.Sometimestherightoneisahummingbird,sometimestherightoneisabumblebee,andsometimestherightoneisabutterflywhosewingsbrushdelicatelyandcarrythepollenaloft.Thesmalleasternredbudtreealsojoinsinthiscelebrationofspringandpollination.Hundredsofscienceexperiments,andacolorthatcontrastswithgreen,tellsusthataflowermustbeseenforittoattractapollinator.Inthedarkunderstoryofaforest,whereredbudlives,it’sdifficultforflowerstobeseeninmidsummer.Soredbudbloomsveryearly,thesametimeasthespringwildflowers,andtakesadvantageofthesolarspotlightforabrieftime.Thenectarinredbudflowersisveryimportantforearly-emergingnativebees.Althoughweoftenappreciateredbudflowersfromadistance,theydeserveaclose-uplookforthefulleffectofthecontrastbetweenthedelicateflowersandthewoodylimbtheyemergefrom.

Thezebraswallowtailbutterflyfeedsonlyonflowernectarandlaysitseggsonlyonpawpawleaves.Becausethecaterpillarlarvaewouldeat

eachother,thefemalelaysonlyoneeggperleaf.

Pawpawleavesproduceachemicalthatinducesthezebraswallowtailtolayeggs.Zebraswallowtaillarvaecanretainthechemicalintotheiradultphase,whenthechemicalmakesthebutterfliestoxictobirdpredators.

Brightcolorswarnthebirds.

WildrhododendronscoverthousandsofacresinthesouthernAppalachianMountains.Theseevergreenshrubsprovidehabitatfortendifferentliverwortspecies.Rhododendronsrequiredeep,moist,organicsoils,andtheirshedleavescontributetomaintainingthisrichsoillayer.

Anindividualleafstaysontheplantforsixyearsbeforeitfalls.

Theyellowmarksonwildrhododendronflowersarenectarguidesthatattractpollinators.Theflowerslookverydifferenttohummingbirdsand

butterflies,whoseecolorswecannot.

Theflowersofmountainlaurelhaveaninterestingpollendispersal

strategy.Justbeforethebudopens,theelongatingstamenspushthepollen-containinganthersupwardintotinypocketsinthesidesoftheflower.Asthefloweropensandexpands,thestamensbendbackwardundertension.Whensomethingtouchesthebentstamen,theanther

springsoutofthepocketandflingsitspollen.

Theredbudtreeaddsasplashofcolorandadelicatebeautytothespringforest.

NativeAmericansateboththeflowersandtheroastedseedsoftheredbudtree.Theflowerscontainbeneficialantioxidantsandarealsoan

importantsourceofpollenandnectarfornativebees.Bythetimetheredbudfinishesblooming,thetipsofthetallertreesaredustedwithhuesofgreenandred.Onmanyplantsthemaleandfemalepartsexisttogether,withinthesameflower;butonmaple,andanumberofothers,flowersareeitheroneortheother—maleorfemale.Acloselookatthespringgroundrevealsthemaleredmapleflowersthathavedroppedfromfarabove,perhapsbouncingoffafewbranchesontheirwaydown.Theseflowerswillbecomesoil,buttodaytheyaddbrightsparkstolastyear’sfallenbrownleavesandthedark,muddyplaces.Ifyoucanfindalowbranch,orperhapsoneonthegroundthatasquirrelhasnippedoff,youcanseetheperfectdetailsofthemaleflowersbeforetheymaketheirlongjourneytotheground.Thefemaleflowersthatwerepollinatedremainaffixedtothetree.Insidetheseflowers,ovariesareswellingandgrowingintothewingedseedsweassociatewithmaples.Whenripe,these,too,willfalltotheground,butinaslowerspinningmotion,opentothewind’ssuggestion.Someoftheseseedsmightgerminateandrootinthesoilandgoontoliveforhundredsofyears.Theotherswillbecomesoil—andsobecomehomeforfuturegenerationsofseeds.Thespringwaveofleaf-outandtheautumnwaveofleaf-turningeachoccurinlessthanamonth’stime,butthewaveoftreefloweringislongerandlesseasyforhumanstosense.Intropicalclimatesthepatterniscomplex,withsometreespeciesfloweringmanytimesayearandothersgoingmanyyearsbetweenflowerings.Thinkofthepatternscreatedwhenwaveshitabeachandthenheadbackouttoseawhileotherwavesareyetrollingin.

Thereddishtintisfromtheflowersofredmapletrees.TheymaybloomasearlyasFebruary.

Thetwoantennae-likethingsarethestickypollen-capturingstructuresofthefemalemapleflower.Theywillinterceptwindblownpollen,andthe

fertilizedovuleswillformtheseedswe’refamiliarwith.

Thesearetheimmaturepollen-producingstructures(anthers)ofamaleredmapleflower.Theywillsplitopenoncetheyripen,andthewindwill

carrytheirpollentothefemaleflowers.

RedmapletreesareamongthemostcommonandwidespreadtreesineasternU.S.forests.Evenveryyoungtreesproduceseeds.Aone-foot-

diametertreecanproduceamillionwindblownseeds.

Manypeoplewhoarefamiliarwithmagnoliatreesdon’trealizethatthetulippoplarisamagnolia,too.Acloselookatthecentralconeandthe

abundantpollen-producingantherswillbringtherecognitionthatthis,too,belongstothefamilyofplantsthatwerethefirstontheearthtobeinsect

pollinated.

Blacklocustflowersprovideabundantspringnectarfornativebees.Thetreeprefersfullsun,soitismostlikelytogrowasapioneerspeciesin

recoveringforestsandisnotnormallyfoundinancientforests.

Inatemperateforest,however,floweringisonapredicableannualschedule:earlybloomers,midseasonbloomers,andlatebloomers.Redbud,early.Maple,early.Tulippoplar,midseason.Blackgum,late.Theearlyonesbloombeforetheirleavesareout.Themidseasonbloomershaveyoungleavesjustfullyexpandedwhentheygetflowers,andthelatebloomershavefullyformedleaveswhentheirflowersfinallyappear.Themoreyearsyouwatch,themoreyougetusedtothepattern.Justasbird-watchershaveasenseofwhentoexpectthefirstwarblerstoreturnfromtheirmigration,tree-watchersknowwhentoexpecttheblacklocustflowersandtheiraccompanyingsweetbreezesandbuzzingsounds.Speakingofthebirdsandthebees,thesightofcertainnativeflowershasbeenshowntocuemigratorybirdstopauseandrefuel.Wherethereareflowersthereareinsects,andinsectsareanimportantfoodforbirdsthatareflyinglongdistances.Insectsareahigh-qualityfood—muchhigherinfatandproteinthanberriesare.Evenpollinatingbeesareoccasionallyeatenbybirds.Differencesbetweenspeciesintree-floweringtimeshavetodowithtreeswantingthebeesalltothemselves.Ifyouneedahelpertomate,it’sbettertohaveyourhelpergodirectlyfromyoutooneofyourkind(orfromoneofyourkindtoyou).Ifalltreesbloomedatonce,beesmightbestoppingtosamplealltypesofdeliciousnectar,andyourpollenmightgetlostalongtheway.Orthebeescouldbringyoupollenthatdidyounogood.Formatingpurposes,it’sbettertobetheonlythingblooming.Sofindyournicheandstickwiththat.Butmanytreesdon’tneedpollinators;they’refertilizedbythewindcarryingpollenfrommaleflowerstofemaleflowersofthesamespecies.Theseflowershaveallthesexualorgans,buttheydon’tneedtheshowy,colorfulpetalsorthesweet-tastingnectar.Thosethingsareonlyneededforattractingpollinators.Whatthewind-pollinatedplantsneedinsteadislotsandlotsofpollen.That’swheretheyputtheirenergy.Theyneedtoreleaseenoughpollenintothewindthatatinydullfemaleflowerisalmostguaranteedtobefertilizedjustbychance.Thepollenyouseeinyellowlayersonpuddles,onyourcar,andonthepatiofurnitureisalmostallfromwind-pollinatedtrees.Manykindsofpollenaredriftingintheair,butforawhitepinetreeonlywhitepinepollenwilldo.Thefemaleflowerstructuresarecoveredinpinkscalesthatprotectovulesjustwaitingforawindblowngrainofwhitepinepollen.Ittakesayearandahalffortheconestoproduceseedreadyfordispersal.Duringwetweatherthescalesstayclosedandholdthematureseedswithin,butduringdryweatherthescalesopen,allowingtheseedswiththeirpaperywingstocatchthewind.Mostconesareproducedhighinthetreetoassurethefarthestjourneypossible.

Noticethattheshowy,sweet-smelling,pollinator-needingflowerstendtohavebothmaleandfemalepartsinthesameflower.Butthesmall,plain,wind-pollinatedflowersareoftensegregatedintomaleorfemale.Ifamaleflowerreleasesitspollentooearlyandnomaturefemaleflowersareopenyet,that’stheendofthatgeneticline.Thereverseisalsotrue:ifafemalefloweropenstooearlyortoolate,itwillreceivenopollenanditsovarieswillwither.Thisishownaturenarrowsthefloweringwindowuntilit’sopenjustacrack.Afterthemaleflowershavereleasedalltheirpollen,theywilldroptotheforestfloorontopoftheredmapleflowers,whichbynowarehardlyrecognizable.Someofthemostabundantmaleflowersfoundontheforestfloorinlatespringarefromoakandpine.Highabovearethetinyfemaleflowernubbinsthatwillbecomeacornsorpinecones.Alongdropawaitsthem,too,butnotjustyet,asitcouldtakethemtwoyearstocompletetheirjourneytomaturity.Thepointofallthisenergycollectedandthenspentbytreeflowersistoproduceavesselthatwillcarrythegenesoftheorganismforwardintime—notjustsothegenescansurvive,butideallysotheywillmultiplyandspreadacrosstheglobe.Alllivingthingsdie—whetherwearetalkingaboutweedsorwarblers—andallspeciesshareacommonimpulsetoreproduce.Wecansaythiswithcertaintybecauseifaspeciesdidn’tcarrythisimpulse,itwouldnotstillbehereforustocelebrateorcurse.Inmammalsthegenesarecarrieddeepwithinthereproductivecells,butintreestheyaredangledoffthebrancheswaitingforapuffofwindorawanderingcreaturetocarrythemaway.Theyareinpollen;theyareinseed.

Ontheleftisthefemale“flower”ofawhitepinetree.Underthesepinkscalesareovuleswaitingforawindblowngrainofwhitepinepollen.Thematureseedsarescatteredbythewindwhentheconesopenindry

weather.

Amaleeasternredcedardevelopsaslightorangecolorintheearlyspringfromthousandsoftinypollen-producingconesliketheonesshown

here.Afemaleproducesbluish“berries.”

Muchlargerthanlife,thesearethetinyflowersofthewhiteoaktreethatwillcatchthewindblownpollenanddevelopintoacorns.Thetreethatsproutsfromoneoftheseacornsmayliveforhundredsofyears.

Whenaredoaktreeisinfullbloom,themale,pollen-producing,flowersdangledowntocatchthewindwhiletheearlyspringleavesare

expanding.Theseedsneedhelpgettingaround,andtheymightalsoneedalittleprotection.

Manyseedsaredeliciousandnutritious,butiftheygeteateninsteadofplanted,theregothegenesforthenextgeneration.Nature’ssolutiontothisistocreateanotherstructure,afruit,tofulfillthecommonassignmentsofdispersalandprotection.Justashumanbabiesarepreparedforlifeinawombthatgrowsastheydo,plantseedsarepreparedforlifeinsideafruitthatgrowsastheydo.Thedifferenceisthatwhenababyisdeliveredtotheworld,thewombstayswiththemother.Inplantstheprotectivewomblikestructurethatwecallthefruitisreleasedalongwiththe“baby.”Incommonusagewethinkoffruitsassomethingsweetandfleshy:blueberries,apples,oranges.Wearelessusedtothinkingofdogwoodberries,orthepaperycoversofredbudandelmseeds,orthespinyballsofthesweetgum,asfruits.Butallthesestructuresareindeedfruits,withtheseedscontainingthegenestuckeddeepinside.Thesweet,brightfruitssay,“Pleasecomeeatme.”Andseedsgetspread.Thepaperyfruitssay,“Wind,comecarryme.”Andseedsgetspread.Seedswiththickhusksrolldownthehillandgetspread.Inanaturalforest,nooneneedstoplantanything.Seedswillbeproducedandwillspreadgenerationaftergenerationaftergeneration.Someseedswillnotliveoneday,andotherseedswillproduceaplantthatwillliveforcenturies.

Whentheredbudtreeshavefinishedblooming,theirfertilizedflowersproducebrownseedpodsthatrattleinthewind.Thepodswilleventuallyfallfromthetreeandbeblowntootherpartsoftheforest.Ifthepodsremindyouofpeas,it’sbecausethey’reinthesameplantfamily.

Sweetgumtreesalsohaveseparatemaleandfemaleflowers;thefemaleflowersproducethesespinyballs.Sweetgumsareamongourmost

colorfulfalltrees,andtheyprovidefoodformanycreatures,frombutterflycaterpillarstosongbirds.

Sycamoretreesnaturallygrowalongrivervalleys,wheretheymayreachastoundingsize.Eachofthelittlebumpsontheseedballisanindividualseedsurroundedbylittlehairsatitsbase.Theseedballshangontothetreeallwinter,breakingapartgraduallyinthespring.Theseedsmaytaketotheairorfloatalongwiththespringfloods,andideallywillreacha

distantfineplacetogerminate.

Windblowntulippoplarseedsmaytraveladistancemorethanfourtimestheheightofthetreeandremainviableformorethanfouryears.

Thewhirligigdownward-spinningflightofaredmapleseedisadelightforallagestowatch.

Lookingattheseseeds,ofpecan,beech,hickory,andoak,onemight

wonderwhattypeofdispersalmechanismtheyuse.Rollingdownahill,perhaps?Actually,squirrels,bluejays,andmicearethepredominantdispersers.Alltheseseedscontainalargestarchymealthat’softenhoardedforlaterconsumption.Unconsumedseedsmaysproutwhere

they’vebeenburied.

Seedsthatfalldownintothedeepleaflitterhavedifficultygerminatingandsurviving.Thetopofarottinglogisamuchbetterplacetoland.Manyofourforesttreesstartedlifeonanurselogsuchasthis.

Therandomnessandcomplexityofnatureareamonghergreatestbeauties.Wedon’tknowenoughaboutforeststoknowhowtoaccomplishthat.Forestsarecommunitiesthattakegenerationstodevelop.Theyarecertainlymorethanjustplantedtrees.TryingtorecreateaforestafterithasbeendestroyedislikesomeonefromanotherplanettryingtorecreateNewYorkCity.Theymayrebuildthemajorbuildingsandevenpopulatethecitywithhumans,butitwillneverbethesamecitythatdevelopednaturallyoveraperiodoffivehundredyears.AsthepoetW.S.Merwinsays,“Onlyaforestcangrowaforest.”

Atfirstitmightseemthatthebestplaceforaseedtolandwouldbedowninthesoftest,deepestplace.Butthatisnotidealforaseed.Downthereaseedmightlandonsuchathicklayerofleavesthatitsrootswouldneverreachthesoil,andevenifitgerminateditwoulddryanddieduringthesummerdrought.Aseedinsuchaplacecouldalsogetcoveredbysomanyleavesthatitsshootwouldnevermakeituptothelight.Aseedinadeep,richplaceisalsomorelikelytobeeatenbythecreaturesoftheforestfloorwhoarealwaysscoutingforfood.Surprisingly,thebestplaceforaseedtolandmaybeuponarottedlogoramossyboulder.Startinglifeinoneoftheseplacescreateschallengesforarootsystemthatmusteventuallyfinditswayintothesoilandstabilizeapotentiallyhugetree.Thesechallenges,andthewaytheyaresolved,createbeautifulforestart.

Thetopofamossyrockisagoodplacetogerminatebecauseitprovidesanescapefromthenibblingmouthsofhungrydeer;however,ayoungtreewillsoonoutgrowthisshallowlayerofsoil,anditsrootsmustfinda

waydowntothetrueforestfloor.

Thepatternofswellingsatthebaseofatreelikethisold-growthwhitepinegivesanindicationofhowmanymajorlateral(horizontal)rootsthe

treepossesses.

ConnectionsRootsandFungi

THEFIRSTSIGNOFlifefromaseedistheemergenceofatinywhiteroot.Theseedcoatcracksandthereitcomes,graspingforconnectionwithasoftfuzziness.Italreadyknowsupfromdown.Thisdelicatewhiteorganhascapabilitieswearejustbecomingawareof.Anelephant’strunkhasbeendescribedbynaturewriterCarlSafinaasacombinationofeyes,nose,hands,andmachinery.Onemightimagineatreerootinmuchthesameway.Iftheseedisfromaspecieswithataproot,likeoakorpine,thatfirstrootwilldivedeeplyintothesoilandsobecomeacriticalanchorforthetreeforaslongasthattreeshalllive.Thedepthoftherootislimitedbytheavailabilityofoxygen.Toodeepandthereisnotenoughairfortheroottolive.Other,lateralrootsgrowclosertothesurface,spreadingoutinalldirections(unlesstheyencounteraphysicalobstacle).Betweenfourandelevenofthesemajorlateralrootsemergenearthesoilline.Itisthisinterfaceoftrunk,roots,andsoilthatJoyceKilmerwasreferringtointhesecondstanzaofhisfamouspoem“Trees.”

AtreewhosehungrymouthisprestAgainsttheearth’ssweetflowingbreast

Theroottipisnotapassivestructurepushingblindlyintotheearth.Root

tipsmovethroughthesoilsniffing,feeling,seeking,andreactingtotherichundergroundworld.

Agerminatingmapleseedshowsthenewlydevelopingrootsystem.Thesehorizontalrootswillmaketheirwayaroundothertreesandgrowfarbeyondthem.Theywillcrossovereachother,eachrootonitsownindividualmission.Therootsdon’tstopwherethetree’sbranchesstopbutmaycontinueonforseventimesthatdistance—fartherthanthedistanceequalingtheheightofthetree.Whentheyencounteranobstacle,theyknowifthey’rerunningintoarockoranotherroot.Andtheyreactdifferentlyifit’stheirownrootorarootfromanotherspecies.Buttherealworkofwaterandnutrientabsorptionisnotdonebythesebigwoodyroots;therealworkisdonebythethinwhiterootsthatfanoffofthem.Weoftenthinkoftreerootsasdeepdown,butthevastmajorityofthemareclosetothesurface—99percentareintheupperthreefeetofsoil.Theserootsstayclosetowherebothoxygenandnutrientsareabundant.Whentheyfindanutrient-richpatch,morerootswillgrowinthatlocation.Androotscansenseandgrowtowardmoisture.Thesoftfeederrootsareadelicacytomanysmallsoilorganisms,soatreemustcontinuallygrownewonestoreplacethosethathavebeeneaten.Justasleavescomeandgoonatree,soalsodorootscomeandgo.Whenyoulookattheabovegroundpartsofatreeandimaginethebelowgroundparts,keepinmind

thatthesurfaceareaoftherootsystemismuchgreaterthanthesurfaceareaofalltheabovegroundparts,includingtheleaves.Atypicaltreehashundredsofthousandsofroottips,eachconnectedbyvesselstothetopofthetree.Andmostofthesedelicaterootsareaccompaniedbyafungusthatwrapsaroundthemandincreasestheirsurfaceareaevenmore.Imaginewrappingyourselfinalargefleeceblanketthatcontinuallydeliversfoodandwatertoyouinexchangeforalittleofyourbodyheat.Astrangeanalogy,perhaps,butitdoesworksomethinglikethat:inexchangeforalittlesugar(madefromsunlightenergy)fromthetree,thefungussupplieswaterandnutrientstothefeederroots.Mycelia(themassoffilamentsthatformsmostofafungus)collectnutrientsfromallovertheforestanddeliverthemtothetreeroots,andthetreerootsaretiedtoeachotherwiththisundergroundweb.Manyplantswoulddiewithouttheirfungalhelpers,butallweseeofthefungiarethereproductiveorgansthatoccasionallypokefromsoilorstemtoreleasespores.It’sabeautiful,complexsystemwe’rejustlearningabout.Andthefungiconnectedtothoserootsthatextendsofarfromthetree,extendevenfarther.Eachtreecanthusbeawareofwhat’shappeningtomany,manyacresofforest.Androotsofindividualtreesofthesamespeciescanformgraftswitheachother,sharingresourcesandspreadingtheirreachevenfarther.Thefungalnetworkcanevenconnectindividualsofdifferentspecies.Notreeisanisland,indeed.

Rootsmayencounterobstacles,andhowtheydealwiththeobstaclesprovidesarecordoftheorganicimprovisationofnature.Thisrockisnow

ananchorforthetree,lendingitsupport.Treerootsrangeinsizefromthedelicatefeederrootstorootsalmostaslargeandhardasthetrunk.Theselargeoldrootsevenformringsastheygrowlargereveryyear.Oneofthedelightsofwalkingthroughaforestisappreciatingtherootsthatweareabletosee—theirshapesareevenmoreinterestingandunexpectedthanthoseofthebranches.Alargerootmaysnakeacrossarockseekingthenextpocketofsoil,orreachforananchortobalancethetopgrowththatgetsheaviereachyear.Sometimestherootscannotholdatreeupright:perhapsthewindsweretoostrongduringthatlaststorm.Thenweseethetreelyingdownwiththelargerrootsrippedfromthegroundbutstillattachedtothetrunk.Atfirstsoilisstillheldbytheseroots,buteventuallythesoilgetswashedawaybyrepeatedrainstorms.Itisthenthatwewitnessnature’sabstractrootart,freeforalltoenjoy.Andwhatbecomesofthedownedtrees?Entirelydifferenttypesoffungiworktodecomposethem.Fungiaremasterrecyclers,turningdeadplantsandanimalsandevenstonesintonutrientsforthelivinggreenlayer.Somefungispecializeinlivingplants,whetherparasitizingorhelpingthem;otherfungispecializeindead

anddyingplantparts,recyclingthembackintosoilandnutrients.Andwithinthosetwospecialtiesarefungithatspecializeinparticularspeciesofplants—whetherdeadoraliveorsomewhereinbetween.Themushroomsthatpokeoutfromtreestemsareoftencalledshelffungus,andyoucanimaginehowaforestgnomemightrestsomethingontopofone.Thefungus’srelationshipwiththetreestartedlongagowhenasmallwoundfromaborerbeetleorafallenbranchgaveanairbornesporeaplacetolandandgrow.Thesporefedoffthelivingwoodcells,replacingtheircontentswithitsownthreadlikestrands.Wherethefungushadfinishedfeeding,thewoodfeltwetandlookedwhitish.Asthefunguscontinuedtogrow,cracksinthebarkweresometimesfilledwithitswhitecottonymass.Eventuallythefungusgotsostrongitwasabletoformitsspore-makingstructure—thepartwecalltheshelffungus.Asmallwhitebulgepokedoutfromthebarkandgotlargerbytheday.Thetoptookonashinydarkreddishcolorandtheunderneathbecamecoveredwithtinyholeswheresporeswereformedandreleased.NowahumanwalkingbycouldtellthatthetreewasinfectedwithGano-derma—afungaldiseasethatcouldeventuallykillthetree.Ifthehumanalsohappenedtobeahealer,hemightcallthisfungusreishiandcollectsome,sinceithasbeenshowntobeeffectiveasawoundhealerandacancertumorreducer.

Deadtreeswithrootsstillattachedaddbeautyandcomplexitytotheforestecosystem.

Abeechtreesendsrootsdownaboulderface.Largerootssuchasthesearenottakinginwaterandnutrients;theyprimarilyprovidestructural

supportandgetthethinfeedingrootsintoabeneficiallocation.Butletussupposethatourtreeisdeepintheforestwherenohumanseverpassby.Inthiscase,thehemlockvarnishshelffungusisstillusefultoanotherorganism.Alargeorange-and-blackbeetlewiththesweetnameofpleasingfungusbeetleisverypleasedtoseethisfungus.Shewilllandonit,eatit,andlayhereggsonit.Whentheeggshatchintowormlikelarvae,theywillfeedonit,too.Withinjustafewweekstheywillpupateandbecomeadultbeetles—andcontinuetofeedonthefungusaslongasitremainsingoodcondition.Someshelffungiinhabitandthenkillatree;othersgrowonatreeonlyafterit’salreadydead.Thedeadtreemightstillbestanding.Chancesareifyouwalkpastatreecoveredinmanysmall,thinshelffungi,whenyoulooktothetopofthattreeyou’llseethatit’sdead.Sometimesasinglespeciesoffungushascolonizedatreeandallthe“shelves”lookthesame.Inotherplaces,avarietyoffungusspeciescanbefoundgrowingtogether,andit’snotuncommontoseelichensormossgrowingamongthem.Whentheshelffungifirstemergefromthewood,theyaredelicateandbrightlycolored.Somehaveconcentricringsthatremindoneofafanned-outbird’stail,andthesegobythenameturkeytail.Whenthefungihavereleasedtheirsporesandgrownoldanddry,theybecomeconvenientplatformsforgreenalgaetogrowon.Thefungalorganismsthathaveneverphotosynthesizedbecomethestageforthatprocess.Layeruponlayer.Otherfungalspeciesresembleananimalhoofmorethanashelf,andthey’recalledconks.Althoughmanyofthesoil-growingmushroomsareephemeral—lastingonlyafewdays—thefruitingbodyofthetinderconkmaylivefordecades,addinganewspore-producinglayeronthebottomeveryyear.Ifyouweretopeelbackthebarkfromanyoneofthesecolonizedtrees,youwouldobservewhitefungalmycelia.Thestrandsgrowingthroughoutthedeadtreeappeardelicateandweak,yettheycontainchemicalenzymespowerfulenoughtodissolvewoodfibers.Thedecompositiontheyperformisveryimportanttotheforestecosystem.Soonthesetreeswillbelyingontheground,becomingsoil.

Thefalseturkeytailshelffungusisapathogenofhardwoodtrees.Ithasbeenshowntobeaneffectiveantibacterialfortreatingstaphinfections.

Varnishshelffungusisaspeciesthatgrowsonlyonhemlocktrees.Connectedtothisfungusisavastwebofmyceliainsidethewood.

Thistypeoffungusconkgrowsallovertheglobeandhasbeenusedbyhumanssinceprehistorictimes.Oneofitsmainuseswasforstarting

fires;henceitbecameknownastinderconk.

Behindeverymushroomisavastnetworkofmyceliacollectingwaterandnutrients.Somekindsfeedonlyonlivingmatter,andotherkindsfeedon

deadorganisms,extractingtheleft-behindenergyandnutrients.Othertypesoffungidonotgrowdirectlyoutoftreetrunks;insteadtheyspringupfromthebaseoftreesorfromthesoil.Itmaynotbereadilyapparentthatthesemushroomshaveanythingtodowithatree,butchancesareverygoodthatthey’reconnectedtotreerootsorsomeotherfoodsourcethroughtheirmycelialnetworks.Oneeasy-to-identifyfungusthat’ssometimesassociatedwithtreerootsisthecoralfungus.Insteadofshelvesorconks,thisfungusformsnarrowfingerscoveredwithspores.Onetypeoffungusthatcangrowonlyonlivingplantsiscedar-applerust.Infact,itneedstwokindsoflivingplantstocompleteitslifecycle.Itfirstinfectsajuniperorcedartreeandproducesabrowngallthatlooksnothinglikeafungustomostpeople.Whenthatgallismature,initssecondyear,strange-lookingorangehornsemergefromitpracticallyovernight.Ifyouweretoexaminetheseorangestructuresunderamicroscope,youwouldseethesporesthatwillbecarriedonthewindtoappletrees.Therethefunguslivesinspotsontheleavesandproducesadifferenttypeofsporethatwilleventuallyblowbacktothecedartreeandkeepthecyclegoing.Hundredsoffungispeciesinhabitatypicalforest,andthefruitingbodiescome

andgounpredictably;asaresult,veryfewhumanscannamemorethantendifferentkinds.Ifyou’reinterestedinnamingthemushroomyou’relookingat,you’llhavetoconsiderthecolorandtextureofthecap,whetherithasgillsorpores,whetherthegillsareattachedtothestalkorseparatefromit,andthecolorofthespores.Anotherfeatureusedforidentificationistheremainsofthetissuethatcoveredthegillsbeforethecappushedfromthegroundandexpanded.Thisringoftissueispresentinsomeofthemorepoisonousspecies.Fungiplayeveryroleimaginableintheforest,fromhelpertoharmer,andrangeinsizefrommicroscopictoanorganismcoveringundergroundacresandsendingupplate-sizefruitingbodies.Westillhavemuchtolearnaboutthefungalkingdom.Wehavemuchtolearnaboutforests,too.Keeplooking.

Notallfungigrowoutofthesoiloroutofatrunk.Theunusualcedar-

applerustfungusgrowsontheleavesandtwigsofredcedartrees.Duringspringrains,orangegelatinoushornsemergefromthebrownball-shapedfungusgalltoreleasetheirspores.Thesporescanonlygrowonaplantintheapplefamily;inturntheappleplantwillproducesporesthat

canonlygrowontheredcedar.

Sometypesofcoralfungusformbeneficialrelationshipswithtreeroots,althoughthemushroomsmayappeartobegrowingrightfromtheforest

floor.

Oncethenetworkofmyceliaisstrongenoughtoproduceamushroom,itcandosoveryquickly.Acapsuchasthiscanspringupovernight.This

onegrewsoquicklyithasstretchmarks.

Somemushroomsemergefromastructurethatresemblesadelicatewhiteundergroundegg.Asthemushroompushesthroughtheenclosure,bitsofitmaysticktothecap,resultinginthewartsweseehere.Thepresenceofwartscanbeakeytoidentification.Manyofthepoisonous

mushroomshavewarts.

Anotherkeytoidentifyingmushroomsisthewaythegillsareattachedtothestalk.

Theshapeandstructureofamushroom’sspore-bearingsurfacescanalsohelpwithidentification.Hereweseetheforkedfalsegillstypicalof

theediblechanterellemushroom.

Thepoisonous“destroyingangel”mushroomcanbeidentifiedbythepresenceofaringoftissueleftoverfromwhentheimmaturegillswere

stillcovered.Alsonotethewartsonthecap.

TheOnly-GreenMossandFern

SOMEFORESTPLANTSWILLnevermakecolorfulflowers—theyareonlygreen.Moss,clubmoss,andferninhabitedthisplanetbeforetherewassuchathingasaflower.Andsomeofthesespecieshavepersisteduntilthepresentera,sowecanstillappreciatetheirearly-evolvedadaptationsandtheiruniquebeauty.Whenweentertheforest,oureyesfeastuponthebrilliantglowinggreenofmossaseagerlyastheydouponthecolorsofwildflowers.Thesetinyplants,too,aresoakingintheraresunshinethatreachesaforestflooronlycertaintimesofthedayoryear.Somethingaboutmossdrawsmorethanjustoureyes.Itdrawsourhands,too.Wewanttotouchmoss.Wewanttostrokeit.Andwe’renotdisappointedwhenwedo.Themossitselfismarvelouslysoft,butwhenithassporecapsulesourstrokesarerewardedwithanunusualspringingmotion.Onceisnotenough.Wemustdoitagain.Andbydoingsowebecometheanimalswhoarespreadingtinyspores.

Isitpossiblethatmosshasevolvedtoattracttouch?Whentheforestoffersaplacetolie,it’softenamossybed.

Thesestalkedsporecapsulesarenotjustanextensionofthegreenmoss.Thecapsulesaretheirowngeneticallydistinctorganism,simplybeingsupportedbytheparentplantbelow.(Thismaysoundfamiliartothosewithgrownchildrenathome.)MosswasoneoftheearliestplantsonEarth.Itwasherewhendinosaursroamed.Itwasherewhentherewerenotrees,backwhensunshinewasabundant

atgroundlevel.Butnowmosshaslearnedtolivewiththetrees.Treeskeepthesoilmoistandcool,somosscannowliveinmoreplacesthanitdidbefore.Ithasfoundwaystogetthesunlightitneeds.Onewayistosoakitupinthespringbeforethetreesgetitall.Anotherwayistogetclosertothesunshinebygrowingonatreetrunk.Themossdoesn’thurtthelivetreeatall.Onadeadtree,themossholdsmoistureandprotectsthefungalfilamentsthatarehelpingtodecomposethewood.This,inturn,helpsthebeetlesthatinhabitsoftenedwood,whichinturnhelpsalltheforestcreaturesthateatbeetles—fromsalamanderstoskunks.Rocks,too,aregoodplacesformosstogrowupandawayfromthesmotheringautumnleaves.Mosshasanadvantagehereoverwildflowersandothersmallplantsbecausemossdoesn’tneedsoil;ithasnoroots,justsmallappendagesthathelpitclingtoasurface.Rootsandvesselsandseedsallcamelaterinevolution.Mosshasnoneofthosethingsanddoesjustfinewithoutthem.Thenextstepinthemarchofevolutionwasthetypeofplantwesometimescallclubmoss.Theseplants,likethemosses,reproducebysporesinsteadofseeds;buttheyhavevesselsthatcantransportwater,sotheycangettallerthanmosses—tallenoughtopokeoutfromamongthefallenleaves.Theyoung,andyoungatheart,amongussometimesmistaketheselittleplantsforbabytrees,butyoucouldwaitfordecadesandtheywouldnevergetanytaller.AtonetimeinEarth’shistoryclubmossspeciesdidgetastallastrees,butthosespeciesdiedoutwhentheclimategotcolder.Theirremainsbecamethefossilsinthefossilfuelswe’restillburningtokeepwarm.

Thissideviewofmossplantsshowsthegreen,parental,generationandtheyellowsporecapsulesthatareasecondgenerationsupportedbythe

parents.

Mossplantsaretheidealhabitatformanymicroscopiccreaturessuchaswaterbearsandnematodes.

Bothstandinglivetreesandfallendeadtreesaregoodplacesformosstoescapethedownedautumnleavesthatwouldburyitandblockthe

light.

Sporecapsulescomeinmanyshapesandcolors.

Mosscangrowonrocksandcapturethemoistureitneedsfromtheair.

Clubmosswasthefirstplanttohavewater-conductingvessels.Asa

result,itwasoncethetallestplantonEarth.

Themanyspeciesofclubmosshavedifferentcommonnames.Thisonemightbecalledrunningcedarorcrowsfootbysome.

Fernscamenext,allthosemanymillionsofyearsago.Theyspreadbyspores,too.Fernsloveforestsforthesamereasonsmossandclubmossdo—dampness.Seedscanlandinadryspotandstaythereforyearswaitingforraintoreleasethem,butsporesaremuchmoredelicate.Mossandclubmossandfernaregreenalloverbecausetheyhavenoreasontoattractpollinatinginsects.Theycantakecareoftheirsexualreproductionneedswithoutanintermediary.Justgivetheseprimitiveplantsalittledampness,andthemalecellscanswimtheirwaytotheeggs.Many,manydifferentspeciesofferns—morethantenthousand—exist,andalmostallofthememergeinthesamedistinctivemanner,likearopeunwindingfromacoil.Oneachfrondthebottomleafletunfurlsfirst,andthenthenextandthenextandthenext,allthewaytothetoptip.Theenergyproducedbythephotosynthesizingbottomleafletshelpstopowerthisexpansion.Thiscoillookssomuchlikethescrollusedtodecoratethetopofaviolinneckthatithascometobecalledafiddlehead.Themorescientificterm,circinatevernation,isbarelyused(circinatemeaningcoiled,andvernationmeaningleafexpansion).Whilemossandclubmosssporesarebarelynoticeable,mostofusrememberseeingfernspores.Theytypicallyappearinclustersonthebackofthefrond,andtheshapeandpatternoftheseclustershelpusidentifythefernspecies.Insomespecies,clustersdon’tuniformlydotthebacksofgreenfrondsbutinsteadareentirelyabsentfromsomeareasandentirelycoverothers.Thesearrangements,too,helpusidentifywhichspecieswe’relookingat.Somefernspecies,liketheroyalfern,haveremainedexactlythesameforthepast180millionyears.Todayveryfewanimalseatferns,butatonetimefernshadavoraciouspredator,oneofthelargestanimalsevertowalktheplanet.The

dinosaurDiplodocuswouldripoutfernsbyhugemouthfuls.Butfernshaveoutlivedthedinosaurs,andnowweseethemmostlyleftalonebylargeanimals.Apparentlydeerfindthemdistasteful.Thesearetheglorydaysformanyfernspecies,likehayscentedfern,whichnowcarpetsmanydeer-impactedforests.

Themiddleleafletsoftheinterruptedfernareentirelycoveredbysporecapsules.Aclose-upofthesecapsulesshowsthempoppingopento

releasetinygreenspores.

Manyferncoloniesspreadbyundergroundrhizomes.Thehairsonacaterpillarcandeterpredatorsandinsulatefromtemperatureextremes,

andthehairsonafernmaydothesame.

Morethantenthousanddifferentfernspeciesliveonthisplanet,andeachhasitsownfavoredhabitat.

Fernsfrondsuncoilfromadistinctivefiddleheadastheyenlarge.

Ofallthegreenground-levelvegetationintheforest,fernisamongtheleastfavoritefoodsofdeer.Sothetrilliumsgeteaten,theshrubsget

eaten,thetreeseedlingsgeteaten,butthefernsremain.

Unlikefungi,lichensliveonlyonthesurfaceoftreesanddonotdecomposethewood.

Mostlichensaresomewhatflat,buttheso-calledreindeerlichensareshrubby.Theseorganismspreferwell-drainedsitesinsparseconiferous

forests;theretheymayliveforahundredyears.

Likemoss,manylichenspeciesgrowbestonrocks.Rocksprovideasurfacefewotherplantscanuse,andasaresultthereislittlecompetition

forspace.Lichensareanotherinterestingmemberoftheforest.They’reauniquecombinationoffungiandalgae.Thiscombinationallowsthemtoliveanywheretheycanfindalittlesunshine.Thealgalpartnerphotosynthesizesandmakessugarthatitshareswiththefungalpartner,andthefungalpartnerprovidesprotection,moisture,andmicronutrients.Togethertheyformaminiatureecosystem.Namingthesetiny,uniqueecosystemshasbecomequiteachallenge.They’retypicallynamedafterthefungalpartner,butonetypeoffungusmaypairwithdifferentkindsofalgaeorcyanobacteria,resultingindifferent-lookingorganismsthatgobythesamename.Thenamingoflichenswilllikelychangeinfutureyears.Justasthevariousshapesandcolorsoffungiarekeystoidentification,soitis

withthelichens.Somemaybespiky,almostlikeminiatureshrubsgrowingfromtheground,whileothersarepressedflattorocks.Somehaveblackhairsthatlooklikeeyelashes,whileothershavetinyreddotsorhornlikeappendages.Althoughthey’regroupedherewiththeonly-greenbecausetheydon’tmakeflowers,lichensalsoexistthataremoreyellowandorangethangreen.Amagnifyingglassishandyforappreciatinglichens.Anysurfacecanprovidehabitatforlichens.Rocksandtreesarelikelysites.Ifyouseelichensgrowingonatree,it’snotasignthatthetreeisinpoorhealth.Unlikethefungithatmayhavethreadlikemyceliainfiltratinganddecomposingthewood,thelichensliveonlyonthesurface,andtreesaremerelytheirscaffold.Liketherootsthatformuniqueandbeautifulpatterns,lichensaddanotherlayerofbeautyandmysterytotheforest.

Thepoetryofsycamoreboughs

TheSpiritoftheForestSOMANYTHINGSTOseeintheforest!Somanystrandsintheweboflife.Somanyquestionsabouthowitallfitstogether.It’sstunningtorealizehowlittleweknow.MariaMitchell,thefirstfemaleastronomerintheUnitedStates(1818–1889),said,“Weespeciallyneedimaginationinscience.Itisnotallmathematics,noralllogic,butissomewhatbeautyandpoetry.”Thissamethinkingcanbeextendedtoourforests,fortheforestisnotonlysomethingtobeunderstood,itisalsosomethingtobefelt.Theforestoffersbeautyandpoetrytothosewhoareopentoit,perhapswaitinginsilenceforittoappear.Itfeelslikeashiftoftheheart,likefallinginlove.Theforestisworthyofbeingloved,andofbeingmournedwhenitisgone.Itismorethantreesandtimberdollarsandhabitat.Itisaspirit-filledplace.Haveyoufeltit?

Thelightfromabove

Theslowembraceoflimbs

Thegreenofspring

Reflectionsoflight

Themagicofnight

Thefeathersofanowl

Theskinofatoad

Thesparkleofice

Anuncommonsight

Soundsthatsoothe

Thefadingofday

Thesensingofspirit

Thelivingforest

Bibliography__________

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Acknowledgments__________

FROMR.L.IstartwithgreatappreciationforJoanMaloofandherpassionfortheforest.SheisagreatteacherandhastransformedwhatIknowabouttheforest.IamforeverchangedwhenIaminaforestandnowseemanythingsthatpreviouslywerehidinginplainsight.Joansentmeasuggestedlistofpossiblephotographs.Furtheralongshesentspecificanduniqueitemstofind.Formethisprojectthenturnedintoacompellingtreasurehunt.Iliveinthemiddleof384millionacresofforest,soIneededhelp.Themosthelpfindingthesetreasures(likea“kissingtree,”wheretwodifferenttreesmeltintoeachother)wasfrommasternaturalistEmilyLuebke,whotookonthelistasachallengeandfoundandguidedmetomanyoftheforesttreasures.MasternaturalistJohnHoldenguidedandcarriedequipmentformeonasalamanderandfrogtriptoMaplesFlats,Virginia,anditsfamouslimestonevernalpools.HewasassistedbytheamazingsalamanderfindersEvaandNorahFloyd.ThanksalsotosalamanderexpertKatyCain.HelpwithotheranimalscamefromCarlaMurray,BuzBireline,andAnakaNazarethoftheRobbinsNatureCenteratMaymontinRichmond,Virginia.HelpwithbirdscamefromRitaMcMahonandherWildBirdFundinNewYorkCity.Someanimalswerephotographedincaptivity.OtherhelpcamefrombirdersThomasJones,AlbertConnette,andWalterCatlett.IwouldalsoliketothankMarkGardnerforhelpwiththeold-growthforestattheWilliamCullenBryantHomesteadinMassachusetts;BarbaraCorcoranandJessicaArcateSchulerwiththeThainFamilyold-growthforestattheNewYorkBotanicalGardens;andAdamDowning,whoguidedmethroughtheold-growthforestatMontpelier,Virginia.OtherassistancecamefromJulieShurenoftheShenandoahNationalParkTrust;andWillNeuman,whoguidedtripsintotheBludeRidge.ComputerandequipmentassistancecamefromJonGolden.ThankstophotographerNickNicholsforguidingmeupthemountaininsearchofwildforestorchids.Thedescentwasinatorrentialmountainthunderstorm.Theforestdoeswhatitwants.IalsoappreciatemyfriendandfellowphotographerScottBarrow,whomakesphotographsonlyinfirstlightandlastlight,forgettingmeupatzerodarkhundredtomakephotographsinfirstlightforthisbook.Thankyoutomybrother,Boxley,andhiswife,Jane,for

transportingustothemostexquisitespecimenofbeechrootsintheworld.IfondlyhonorKathyandTroostParkeralongwithBillandCarolynAchenbach,whotookonthisprojectandguidedandassistedmeonjourneysthroughthemoss-coveredforestfloorintheCheatMountainrainforestandotherold-growthforestsofWestVirginia.Ihavegreatappreciationformypublisher,TimberPress,andforTomFischer,editor-in-chief;PatrickBarber,creativedirector;andAdriannaSutton,bookdesigner,fortheirsupportandthetransformationofthisprojectintoagreatbook.IamgratefulforthephotographysupportoftheSouthernEnvironmentalLawCenteranditsworkpreservingroadlessforestsintheGeorgeWashingtonNationalForestinVirginia,savingold-growthforestsfromtimberinginthePisgahNationalForestinNorthCarolina,andsavingtheuniqueforestofSandyIsland,SouthCarolina,fromdestruction.Andfinally,andmosttreasured,Ihaveenormousloveandappreciationformywife,Bobbi,wholovestowalkintheforestandwouldaccompanymeonmostofmyadventures.ShewouldoftenwalkinlargecirclesaroundmeasImadephotographsorasIwaitedforthelight.

FROMJ.M.MydeepestthankstoRobertLlewellynforinvitingmetoputwordstohiswonderfulimages.Ithasbeenadelightfuljourneyfrombeginningtoend.

Index__________

A �B �C �D �E �F �G �H �I �J �K �L �M �N �O �P �Q �R �S �T �V �W �Y �Z �

Aacorns,116,174,176algae,71–72,82,198,227Americanbeech,124amphibiansvs.reptiles,68animalsbehaviorresponsestolight,46,49,72colorchanges,118,119dead,asfoodsource,28primitive,82,85,102,220asseeddispersers,185assporedistributors,88,90,114,209duringwinter,83,92,99,107,135,138Seealsospecificanimals

anthers,166,170,173ants,104,107AppalachianMountains,74,165applefamilyplants,202azalea,163

Bbacteriacyanobacteria,227inthefoodweb,99humandigestionand,102onrottingmeat,28

barkbatroostsunder,26,113chewingpatternson,113,114fungaldiseasesof,114,195,198patternsandtexture,110–111,232

barkbeetle,113–114batroosts,26,113bats,26,113,135bears,95,101,135beechnuts,116beechtreeleaves,37,124,132beechtrees,40,101,124beesaspollinators,163,167,173beetlesbarkbeetle,113–114intheforestecosystem,114fungusbeetle,198southernpinebarkbeetle,114tigerbeetle,107

birchtrees,101birdscanopyashabitat,14,16,19–22,28ears,28,30foodsources,21human-birdinteraction,14,22migratory,16,19,46,173nestsandnestingbehavior,16,19,21,22,28asseeddispersers,185useofsenses,28,30Seealsospecificbirds

birdsofprey,28,30blackbears,101.Seealsobearsblackgumfruitasfoodsource,92asalatebloomer,173leafcolor,126

blacklocustflowers,173

blackratsnake,95,99blackvultures,30bloomtimesoftrees,168,173bluebirds,21bluedasherdragonfly,79bluejays,185bobcat,97borealforest,124boxturtle,87,88,92branchingpatterns,36,39–40,233,243breathingofforests,51–52,54–55,57,60bristleconepines,120broad-leavedforest/trees,14,124,131brooktrout,68budbreak,143,150,155budscherrytree,142commontypes,142description,141–147dogwood,147maple,142–143intreeidentification,142,145tulippoplar,150–151

butterflies,158,163,164

Ccalciuminanimaldiets,88,90canopy,viewsfrombelow,14,231carbondioxide,52,54–55,57,60cardinals,21,22Carolinabays,71carp,68caterpillars,26cedar-applerust,202cedartrees,176,202chanterellemushroom,205cherrytreebuds,142chlorophyll,124cicadanymph,88cicadas,116circinatevernation,defined,220clubmoss,209,212,219,220colorinattractingpollinators,158,166changingbyanimals,118,119only-green,209,227wavelengths,44

colordiversityaltitudeand,155diversityofspeciesand,129,131seasonalcontrasts,135inspring,143,150

cones,173coniferousforests,227conifers,129copperhead,95coralfungus,202cougars,92,95coyotes,92,95crayfish,68,82crows,22crowsfoot,219cyanide,102

Ddampconditions,64,68,71,133,220daylength,46deadanimalsasfoodsource,28deadmatter,99,102deadplantsinthefoodweb,102decomposition,102,195,198,212,227deerdeterrencesto,124,189effectsontheforest,92fernsasdeer-resistant,220,225population,92senseofsmell,95

dinosaur-ageplantsandanimals.Seeprimitiveanimals;primitiveplantsDiplodocus,220dog-daycicadas,116dogwoodberries,87,116dogwoodbuds,147dogwoodtrees,148dragonflies,79dragonflylarvae,74dragonflynymphs,68,79

Eeagles,19,28,30easternbluebirds,21easternredbudtreeflowers,163easternredcedar,176easternUnitedStatesfall,131forests,168,171spring,150,155wildlife,97

eggsandegg-layingamphibians,71ants,104barkbeetles,113–114calciumneedsforproducingeggs,88,90frogs,72,74fungusbeetles,198reptiles,68salamanders,71–72,74skinks,68,74tigerbeetles,107treefrogs,118turtles,80zebraswallowtailbutterfly,164

eggsasafoodsource,80embryoshuman,68reptile,68salamander,71–72seed,116

emeraldashborer,114ephemeralponds,71evergreenshrubs,165evergreenspeciesashabitat,165leavesandwinterchallenges,124sheddingrate,120

Ffalseturkeytailshelffungus,198far-redlight,44fernsasdeer-resistant,220,225fiddleheads,220,223identifyingfeatures,220asprimitiveplants,209,220rhizomatousspreadingof,223species,220,223sporecapsules,220

fishspecies,68flicker,22fliesaspollinators,163floweringshrubs,163flowerreproductivepartsanthers,166,170female,168,169,170,174,180male,168,169,174,177,180maleandfemaleinthesameflower,168,174ovules,166,170,174stamens,166

flowerscolorinattractingpollinators,158,163,166ascuesformigratorybirds,173edible,167ephemerals,157odorinattractingpollinators,163wildflowers,157–160Seealsobuds;treeflowers“forest-grown”trees,36

forestpools.Seevernalpoolsforesttentcaterpillar,26Fowler’stoad,104frogscolorchanges,118,119eggs,72,74asprey,68,80reproductivehabitat,71,72,74springpeepers,72,74,78tadpoles,74,78,80,82treefrogs,118,119inwinter,135

fruits,defined,178fungaldisease,195fungicedar-applerust,202conks,198,201coralfungus,202decompositionfunctionof,195,198,212,227inthefoodweb,102galls,202medicinalproperties,195,198mycelia,193,198–199,201,202,204odor,131recyclingfunctionof,195spores,193,195,198,202Seealsomushroomsfungusbeetle,198

GGanoderma,195grayfoxes,95,97graysquirrels,116graytreefrog,119greatblueheron,14,16greathornedowl,28

Hhairinessofbuds,142,147asidentifyingfeature,142,227oflichens,227forprotection,223

hardwoodtrees,101,198hawks,19,28hayscentedfern,220hemlocktrees,129,199hemlockvarnishshelffungus,198heron,14hickorynuts,116hickorytrees,151hoarfrost,137homerangesofbobcats,97ofsalamanders,74ofturtles,92

hornbeam,132hornedowl,28housesparrows,21humansdigestionandbacteria,102embryonicenvironment,68interactionwithbirds,14,22responsestoforestlight,44sensingofwavelengths,35,44

hummingbirds,162,163,166

Iinsectsbarkashabitat,113,118canopyashabitat,24–26effectonleaves,124asfoodsource,21,22,26,116,162,173night-flying,26aspollinators,163,167,173poolsasreproductivehabitat,71treeproductionofdeterrentcompounds,26inwinter,135

interruptedfern,220

Jjack-in-the-pulpit,163junipers,202

Kkingsnake,118

Llarvaechewingpatterns,113,114insect,74,82,113–114,164,198salamander,72–73,74,80

leaf-droppingprocess,120,124leafingout,timingof,46,155,168leafscarsintreeidentification,145leaf-turning,timingof,168,173leavesdeciduousspeciesthathold,124,132fallcolor,124,126,129,131growth,141aslightreceptors,35microscopically,55duringrain,64redpigment,124,126stipules,150,151stomata,51,54Seealsoleaves,fallenleaves,fallenasfood,88,102,132,133leaf-droppingprocess,120,124asprotection,132,133

lichens,226–227life,impulseof,174,232lightaestheticaspects,231,236,240,246animalresponsesto,46,49,72effectonbranchingpatterns,36,39–40far-red,44humanresponsesto,35,44moonlight,46,49,72plantresponsesto,45,46,57sharingbytrees,36–37,39starlight,46,49sunlightinforestexhalation,57,60wavelengths,35,44

liveoak,40liverwortspecieshabitat,165

Mmagnoliatrees,173mapletreeasanearlybloomer,173budbreak,150buds,142–143ineasternUSforests,168fallcolor,126flowers,140,150,155,168–170asahardwood,101leaves,133seeds,168,171,184,193

mapturtle,83marbledsalamander,72,80mayapples,87,88miceasprey,95,99seeddispersalby,185senses,92,95

migratorybirds,16,19,46,173millipedes,102–103,104moonlight,46,49,72mossgrowingconditions,212ashabitat,213planthistory,212onrocks,187,212,217sporecapsules,209,212,215spores,209astouchable,209,212ontrees,63,215

moths,24,26,46mountainlaurel,163,166mushroomschanterelle,205coralfungus,202“destroyingangel”,207asafoodsource,88,90,205identifyingfactors,202,204–205naming,202poisonous,202,204,207

shelffungus,195mussels,82mycelia,193,198–199,201,202,204

Nnectarcollectionbybees,163,167,173,176bybutterflies,158,164byhummingbirds,162,163,166

nectarguidesonflowers,166nematodes,213nestsandnestingspacesbluebird,21constructionof,19greatblueheron,16raptor,19,28turtle,80woodpecker,21,22

night,238night-flyinginsects,26,46nocturnalanimals,28,46northerncardinals,21,22northernflicker,22nurselog,187nutsasafoodsource,116

Ooakflowers,174oakleaves,124,126,155oaktrees,132,142,191odor.Seescent;smell,senseofosprey,19owls,28,239oxygenabsorptionbyturtles,83availabilityandrootdepth,191,193channelstoreachroots,80collectionbysalamanders,72fishspeciesandlevelsof,68inforestexhalation,52,54–55,57,60inleafcellprocesses,54–55,57,126

Ppaintedturtle,80,82pawpawleaves,164photosynthesiscarbondioxideand,54,60leafdropandstoppingof,37,124lichenformationand,227lightwavelengthsin,44

photosyntheticorganismsinthefoodweb,101pinebarkbeetle,114pinecones,174pines,120,124,129,174,191–192pinevs.broad-leavedforest,14pollendispersalstrategyofmountainlaurel,166pollenwands,163pollination,wind,170,173,174,176,177pollinatorsbees,163,167,173butterflies,163flies,163flowercharacteristicsforattracting,158,163,166hummingbirds,162,163,166moths,46night-flying,46

polyphemusmoth,24pools.Seevernalpoolsprimitiveanimals,82,85,102,220primitiveplants,209,212,220puddles.SeevernalpoolsQquotations,14,132,187,191,229

Rraccoons,80rain,60,63–65rainbows,44,131raptors,19,28rattlesnakes,95,98red-backedsalamanders,72redbudasanearlybloomer,163,173fallcolor,124flowers,163,167seeds,167,178

redcedartrees,176,202redcornsnake,118,119red-eyedperiodicalcicadas,116red-headedwoodpeckers,21.22,28redmapletree.Seemapletreeredoakflowers,177red-tailedhawk,33redwolves,97reindeerlichens,227reishi,195reptilesasair-breathing,82amphibiansvs.,68scalesasidentifyingfeatures,28,68,74timewhenmostactive,46Seealsospecificreptiles

rhizomes,159,160,223rhododendron,163rocksasagrowingsurface,189,212,217,227asobstaclesforroots,193,197

rootsofdownedtrees,195,197encounteringobstacles,191,193,197extentandsize,193,195lateral,191–193forstructuralsupport,191,194taproots,191tipsof,192waterandnutrientabsorption,193

runningcedar,219

Ssalamandersburrows,80embryos,71–72highlydevelopedsenses,73,74homerange,74larvae,72–73,74,80lifestylesandtypes,72,74,80matingbehavior,71,72,74waterasreproductivehabitat,68,71–72

SandyIsland,SouthCarolina,19scentinattractingpollinators,163offallenleaves,133ofmillipedes,102Seealsosmell,senseofseasonscolorcontrasts,135elevationandlengthof,155inthetemperateforest,141Seealsospecificseasons

seedsdispersalof,178,180,183,185,187asfoodsource,116fruitsasprotectivestructure,178germination,placesof,187,189seedballs,180seedpods,178

shagbarkhickory,113shelffungus,195,198–199shrubs,evergreen,165shrubs,flowering,163sight,animalswithwell-developed,73,74,95skink,68,74slugs,26smell,senseofanimalswithwell-developed,73,74,95inlocatingprey,28,30inmatingbehavior,24,113Seealsoscentsnails,importancetoecosystem,88,90

snakes,80,95,98,99,118,119snappingturtle,80,82,85snow,124,135,138

soildecomposedtreesasacomponent,198depthandtreeroots,193erosion,64,195fallenflowersasacomponent,168fallenleavesasacomponent,132,133,165Seealsofungi;rootssoundsbats,26moths,26soothing,245springpeepers,78wind,14

SouthCarolina,19southernAppalachianMountains,74,165southernpinebarkbeetle,114spiders,24spirit,sensingof,14,229,249sporecapsulesferns,220moss,209,212,215

sporesdistributionbyanimals,88,90,114,209fern,220releasebyfungi,193,195,198,202winddispersal,202

spottedsalamander,71,72,80springcolordiversity,143,150,155,234ineasternUnitedStates,150,155timingofonset,155

springpeepers,72,74,78squirrelscommunication,meansof,116,118dominancehierarchy,116food,116,118,147,148matingbehavior,116responsetopredators,118seeddispersingby,185swarming,118trunksashighwaysfor,114,116

starlings,21stipules,150,151stomata,51,54streams,64,68,80StroudWaterResearchCenter,68

sugarmaple,150summer,altitudeandlengthof,155sweetgumtreeflowers,180sweetgumtrees,126,142,145sycamoreseeds,87,180sycamoretrees,124,180,229

Ttaproots,191temperateforestfallcolor,131leavesduringwinter,124measurementofseasons,141timingofflowering,173

termites,104tigerbeetle,107tigersalamanders,74,81timberrattlesnake,95tinderconk,201toads,102,104,239touch,senseof,73,95tree-climbingtoescapepredators,95,97treeflowersblacklocust,173colordiversity,143,150,155easternredbud,163flowering,timingof,168,173oak,174redbud,163redmaple,150,155,168–170redoak,177sweetgum,180whiteoak,176Seealsobuds;flowerreproductivepartstreefrogs,118

treeidentification,142,145trilliums,225tropicalforest,124trout,68trunksclimbingtoescapepredators,95,97descriptivefeatures,109uses,110asverticalhighways,110,114,118

tulippoplarbuds,150–151fallcolor,124asamidseasonbloomer,173seeds,183

three-trunked,110turkeys,88turkeytailfungi,198turkeyvultures,28,30turtlesanimalrelatives,85baskingbehavior,82boxturtles,87,88food,87–88,90,92homerange,92lifespan,88,92mapturtles,83matingbehavior,82,88nests,80paintedturtles,80,82physicalstructure,82asprey,92snappingturtles,80,82,85inwinter,83,92,135

Vvarnishshelffungus,199vernalpools,71,80,138Virginia(state),74,158Virginiabluebells,158vultures,28

Wwatercleaningof,64–65,68livingconditionsforfishspecies,68movementandreleaseof,55,57,60,65,68rain,60,63–65asreproductivehabitat,68,71–72sensoryappreciationof,66streams,64,68,80vapor,51–52,54–55,57,60,63vernalpools,71,80,138

waterbears,213whiteoakacorns,116whiteoakflowers,176whiteoakleaves,148whitepines,120,174,192white-taileddeer.Seedeerwildflowers,157–160wildgeranium,159wildrhododendrons,165,166winddispersalofseeds,178,183dispersalofspores,202effectsonleaves,124pollination,170–171,173,174,176–177soundof,14

wingedseeds,168,169winteranimalsduring,83,92,99,107,135,138challengestoleaves,124contrastwithfall,135

wolfspiders,107wolftrees,40wolves,92,95,97woodpeckers,21,22

Yyellow-spottedsalamanders,74

Zzebraswallowtailbutterfly,164

AbouttheAuthors__________

RobertLlewellynRobertLlewellyn’sphotographshavebeenfeaturedinmajorartexhibits,andmorethanthirtybookscurrentlyinprint.Hisbook,EmpiresoftheForest:JamestownandtheBeginningofAmerica,wonfivenationalawardsinnonfictionandphotography,andWashington:TheCapitalwasanofficialdiplomaticgiftoftheWhiteHouseandStateDepartment.

JoanMaloofJoanMaloofisascientist,writer,andthefounderanddirectoroftheOld-

GrowthForestnetwork,anonprofitorganizationcreatinganetworkofforestsacrosstheU.S.thatwillremainforeverunloggedandopentothepublic.ShestudiedplantscienceattheUniversityofDelaware,environmentalscienceattheUniversityofMarylandEasternShore,andecologyattheUniversityofMarylandCollegePark.SheistheauthorofTeachingtheTreesandAmongtheAncients.

Textcopyright©2017byJoanMaloof.Photographscopyright©2017byRobertLlewellyn.Allrightsreserved.Publishedin2017byTimberPress,Inc.TheHaseltineBuilding133S.W.SecondAvenue,Suite450Portland,Oregon97204-3527timberpress.com

TextdesignandcoverdesignbyAdriannaSuttonLibraryofCongressCataloging-in-PublicationDataNames:Maloof,Joan,1956–author.|Llewellyn,RobertJ.,photographer.Title:Thelivingforest:avisualjourneyintotheheartofthewoods/photographybyRobertLlewellyn;writtenbyJoanMaloof.

Description:Portland,Oregon:TimberPress,2017.|Includesbibliographicalreferencesandindex.

Identifiers:LCCN2016057868|ISBN9781604697124(hardcover)Subjects:LCSH:Forestsandforestry—Pictorialworks.

Classification:LCCSD385.M352017|DDC634.9022/2—dc23LCrecordavailableathttps://lccn.loc.gov/2016057868

AcatalogrecordforthisbookisalsoavailablefromtheBritishLibrary.