The Phenology Handbook-Haggerty Mazer 2008 v1

8/9/2019 The Phenology Handbook-Haggerty Mazer 2008 v1

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The Phenology Handbook

A guide to phenological monitoring for students,

teachers, families, and nature enthusiasts

Brian P Haggerty and Susan J Mazer

University of California, Santa Barbara

2008 Brian P Haggerty and Susan J Mazer


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Acknowledgments

Since the Spring of 2007 it has been our great pleasure to work with a wide variety of students, educators, scien

sts, and nature enthusiasts while developing the Phenology Stewardship Program at the University of California

Santa Barbara. We would like to express our gratude to those who have contributed (and who are currently con-

tribung) me and energy for eld observaons, classroom implementaon, and community outreach. We thank

the Phenology Stewards (UCSB undergraduates) who have helped to collect plant and avian phenological data at

UCSBs Coal Oil Point Natural Reserve and to develop the methodologies and protocols that are presented in thishandbook. Special thanks to the Phenology Stewardship graphic design team who helped develop this handbook

including Christopher Cosner, Karoleen Decastro, Vanessa Zucker, and Megan van den Bergh (illustrator).

We also thank Sco Bull, the UCSB Coastal Fund, and the students of UCSB for providing funding for the devel-

opment and connued operaon of the Phenology Stewardship Program at UCSB.

We would like to thank those in the Santa Barbara region who are implemenng phenology educaon and engag-

ing students to parcipate in Project Budburst, including:

Dr. Jennifer Thorsch and UCSBs Cheadle Center for Biodiversity and Ecological Restoraon (CCBER), as

well as the teachers associated with CCBERs Kids In Nature environmental educaon program;

the teachers in training in the Teacher Educaon Program at UCSBs Gevirtz School of Graduate Educa-

on who work in K-12 schools throughout Santa Barbara; and

docents at natural reserves and botanic gardens, including Coal Oil Point, Sedgwick, Arroyo Hondo, Ran-

cho Santa Ana, Carpenteria Salt Marsh, Santa Barbara Botanic Garden, Lotusland Botanic Garden.

We would also like to express our gratude to the consorum of sciensts and educators associated with the USA

Naonal Phenology Network, and especially to our colleagues in the NPNs Educaon, Cizen Science, and Out -

reach working group. With operaonal support from the University Corporaon for Atmospheric Reserach and

with funding from the U.S. Bureau of Land Management, the Naonal Science Foundaon, ESRI, and the Naona

Fish and Wildlife Foundaon, the ECSO working group launched Project Budburst in the Spring of 2007.

This Phenology Handbook is a work in progress, and will be updated connually through the spring and summer

of 2008. We plan to increase the scope of the handbook by including chapters that address the following topics:

methods for using the herbarium to study historical phenology;

a eld guide to Coal Oil Point Natural Reserve, including species descripons that work in coordinaon

with the Phenophase Quick-Guide (see pages 38 - 40); and

a phenology acvity guide for teachers and students.

Updated versions of the Phenology Handbook will be posted on the UCSB Phenology Stewardship Program

website, which is currently located at: hp://www.uweb.ucsb.edu/~cmcosner/phenology/

Brian P Haggerty, M.S. Susan J Mazer, PhD

Santa Barbara, CaliforniaMarch, 2008


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The Phenology Handbook

A guide to phenological monitoring for students,

teachers, families, and nature enthusiasts

INTRODUCTION

2 What is phenology?

3 Phenologicalvariaonacrossbiologicalandgeographicscales

Quanfyingvariaonwithinandamongindividuals,populaons,andcommunies

Summary&Synthesis

12 Themodernscienceofphenology

20 Phenologynetworks-Agatewaytodetecngenvironmentalchange

Theroleofcizensciensts

TheUSANaonalPhenologyNetwork

ThePhenologyStewardshipProgramattheUniversityofCalifornia,SantaBarbara

PROTOCOLS FOR PHENOLOGICAL MONITORING

28 Introducon

28 Plant phenophases

29 Generalconsideraonsforstarngobservaons

Where,when,andwhichspecies

34 Protocols

Phenophasedenions

ThePhenophaseQuick-Guide

Allcontent,images,anddesignsBrianPHaggertyandSusanJMazer,unlessotherwisenoted


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WHAT IS PHENOLOGY?

PHENOLOGY IS THE OBSERVATION AND MEASUREMENT OF EVENTS IN TIME

ThepassingoftheseasonsisoneofthemostfamiliarphenomenaonEarth.Consider,forexample,theonset

ofspringintemperateclimates.Aswinterends,oursurroundingsburstwithnewlifeforestcanopiesllwith

vibrantgreens,ocksofbirdsmigrateinformaontonorthernbreedinggrounds,andbrilliantwildowersand

theirinsectpollinatorsappearinrapidsuccessionacrosshillsides,roadsides,lakemargins,andelds.Similarly

asautumnapproaches,thedeciduousforestcanopyprogressestowardsacolorfuldemise,birdsnavigatetheir

returntosouthernwinteringgrounds,andmanyplantsripentheirlastfruitsbeforetheonsetofwinter.

Whetherweliveinurbanorruralenvironments,thereareconstantremindersofthechangingoftheseasons

Forexample,manyofusnocewhentherstwildowersappearinthespring,whenourfavoritefruitsareavail

ableinthelocalfarmersmarketsorsupermarket,whentreeschangecolororlosetheirleavesinthefall,when

wildreriskishighestduetothedryingofourforestsfuels,orwhenfrostrstappearsinthefallorwinter.By

studyingtheseasonsingreaterdetailoverthecourseofourlives,wecandeepenourconneconwith,andun

derstandingof,thelandscapesweinhabit.Wealsodevelopourabilitytoobserveandtomeasurethepaceandthemingoftheseasons,theonsetandduraonofwhicharebeginningtoshiwiththechangingclimate.

Scienstsrefertothestudyofthemingofseasonalbiologicalacvi-

esasphenology.Thistermwasrstintroducedin1853bytheBelgian

botanistCharlesMorrenandisderivedfromtheGreekwordsphaino,

meaningtoappear,tocomeintoviewand logos,meaningtostudy.

Phenologyisthesciencethatmeasuresthemingoflifecycleeventsfor

plants,animals,andmicrobes,anddetectshowtheenvironmentinu-

encesthemingofthoseevents.In thecaseofoweringplants,these

lifecycleevents,orphenophases,includeleafbudburst,rstower,last

ower, rst ripe fruit, and leaf shedding, among others. Phenophasescommonly observed in animals include molng, mang, egg-laying or

birthing, edging, emergence from hibernaon, and migraon. Thus,

phenologistsrecordthedatesthattheseeventsoccur,andtheystudy

howenvironmentalcondionssuchastemperatureandprecipitaonaf-

fecttheirming.

Themingofphenologicaleventscanbequitesensivetoenvironmentalcondions.Forexample,inaparcu-

larlywarmanddryspring,leafbudburstandrstowermightoccurweeksearlierthanusual,whereasinan

exceponallycoolandwetspringtheycouldbeequallydelayed.Asaresult,themingofphenophasestends

tovaryamongyearsbasedonpaernsofweather,climateandresourceavailability.Phenologicalobservaons

arethereforeintegravemeasuresofthecondionofthephysical,chemical,andbiologicalenvironment.This

environmentalsensivitymeansthatphenologicalstudiesaresimpleandcost-eecvewaystomeasureenvi

ronmentalchanges,includingclimatechange,overthelong-term.

While the term seasonality is

used to describechangesin the

abiocenvironment,suchasthe

dates of the rstand last frosts

of the winter, or the date that

icemeltsinlakesorstreams,the

term phenology is reserved fordescribing theming ofbiologi-

calacvies.

Weatherisdenedasthenear-termatmosphericcondionsofaregion,suchastemperature,precipita-

on,humidity,wind,andsunshine.The climateofaregion,ontheotherhand,ischaracterizedbythe

generally-prevailingweathercondions.Forexample,SantaBarbara,CaliforniaischaracterizedbyaMed-

iterraneanclimatewarm,drysummersandcool,moistwinters.Thereare,however,dailyandweekly

changesintheweatherthatcanrapidlychangethetemperature,sunshine,andwindcondions.

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PHENOLOGICAL VARIATION

The ming and duraon of phenophases within and among individuals, populaons, and communies

Phenologyisascienceforallseasons,locaons,andspecies.Fromtheleang,owering,andfruingmesof

plantstothemolng,mang,andmigraonmesoftheanimalstheysupport,thephenologicalprogression

ofanindividual,apopulaon,oranenrespeciesmayoccurrapidlyorslowly,andsynchronouslyorasynchro

nouslywithotherorganisms.How,then,canwemeasureandcomparephenologicalpaernsforawidevariety

oforganismsthatexhibitdierentphenophasesandatdierentmescales?Theanswerliesinthescaleofour

observaonsandinthewaysthatwerecordanddescribethem.

Phenology,likeallenvironmentalsciences,usesquantavemethodstomeasureandtodescribetheoccur-

renceofeventsandpaernsinthenaturalworld.Phenologistsareinterestedinthedatesthatphenophasesoc-

curaswellasinthepaceoftransionsbetweenphenophases,andtheseobservaonscantakeplaceatmulple

biologicalandgeographicalscales.Forexample,aphenologistcanrecordthedatesthataplantopensitsrst

andlastowers(fromwhichtheduraonofoweringcanbecalculated),aswellasthenumberofowersthat

areopeneachdayorweekduringtheoweringperiod.Thiscanbedoneforoneormanyindividualsofthesame

speciesinonepopulaon,formulplepopulaonsofaspeciesthatoccupiesdierenthabitatsorlocaons,andformulplepopulaonsofdierentspeciesthatcoexistinonehabitat.

Levels Of Biological Organizaon

Thebiologicalworldcanbeobservedatavarietyofscalesrangingfromlong-distanceobserva-

onstoclose-upanddetailedviews.Eachscalegenerallyrequiresadierentsetoftools,fromsatel-

litestomagnifyingglassestomicroscopes.Phenologyisoneofthefewsciencesthatrounelymeasure

paernsatmanyofthesescales.Thelevelsofbiologicalobservaonthataremostimportanttothe

globalandlocalstudyofphenologyareshownbelow.

BiomeA group of co-occurring plant, animal, and microbial

communiesthatliveinthesametypeofclimate,share

awell-denedgeographicarea,areadaptedtoaparcu-

larsubstrateandlevelofnutrientcycling,andexhibita

recognizeablesetofdominantlifeformsandhabitats.

Community

Agroupofco-occurringpopulaonsofdierentspecies,

eachofwhichinteractswithsomeproporonoftheoth-

erspecies.

Populaon

Acolleconofindividualsofonespeciesinhabingthe

samegeneralareaorsharingacommonenvironment.

Organism (Individual)

Onememberofapopulaonorspeciesthatmayormay

notdependonothermembersofitspopulaoninorder

tosurvive.

Remote sensing technologies allow

for the detecon of geographically

extensivephenological paerns

Observaonal studies performed

by on-the-ground phenologists pro-

vide site intensive documentaon

of phenological paerns


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Phenology Across Levels Of Biological Organizaon

Individual:Anindividualisonememberofapopulaonorspeciesthatmayormaynotdependon

othermembersofitspopulaontosurvive.Individualsofthesamespecieswithinacommunitybe-

longtothesamepopulaon,andmayexhibitgreatvariaoninthemingoftheirphenophases.The

parcularphenologicalpaernthatanindividualexhibits(e.g.,thedateofgerminaon;theonsetand

duraonofowering;theaveragenumberofowersopenperdayduringtheoweringseason;and

themeofseeddispersal)isusuallyduetobothgenecandenvironmentalinuences.

Theming of an individuals phenophases canhave a profound eecton itsreproducvesuccess,

asitislikelytodeterminewhether:itsurvivestoower;owersarepollinated;seedsareproduced,

ripened,andsuccessfullydispersed;andwhethermostseedswillgerminateandproduceseedlings

thatsurvivetoadulthood.Consequently,themingofphenophasesinplantsasinotheranimalsis

subjecttoevoluonbynaturalselecon.Iftheclimatechangesgradually,andifpopulaonscontain

enoughgenecvariaoninphenologicaltraitssuchasthedateofrstower,theywillbeabletoadapt

asnaturalseleconfavorsthosephenophasesthatperformwellunderthenewenvironmentalcondi-

ons.Evoluonarybiologistsareconcerned,however,thatifclimatechangeprogressestoorapidly,

manypopulaonswillnotbeabletoadaptbecausetheywillnotcontainanygenecvariantsthat

performwellunderthenewcondions.

Populaon: Acolleconofindividualsofonespeciesinhabingthesamegeneralareaorsharinga

commonenvironment.Thephenologicalpaernsofindividualsinapopulaonareimportantbecause

they determine: whether individuals areavailable formangwitheach other; whether individuals

competeforresourcessuchasmoistureinthesoilorfood;whichindividualsareexposedto(orescape

from)ower-andseed-eangpredators;andwhetherindividualscancooperatetodeterherbivoresor

predators.

Species:Speciesisa collecvetermthatusuallyreferseithertooneortoallofthepopulaonsofa

givenspecies.Populaonsofagivenspeciesmaybelargeorsmall,closelyspacedorhighlyisolated,

andcomposedofindividualsthatoccurinclumped,uniform,orrandomdistribuons.Wheremulplepopulaonsofagivenspeciesarephenologicallysynchronizedandnottoofarawayfromeachother,

thepotenalformigraonandmangbetweenpopulaonsismuchhigherthanwhentheyarephe-

nologicallymismatched.Theexchangeofindividualsamongpopulaonscanbeveryimportant,asitis

onemechanismthatcanhelptomaintaingenecdiversitywithinthem.

Community:Acommunityisagroupofco-occuringpopulaonsofdierentspecies,eachofwhich

interactswithsomeproporonoftheotherspeciesinthecommunity.Someoftheinteraconsare

posive,inwhichbothspeciesbenet(examplesincludepollinaonandseeddispersal).Otherinter-

aconsbenetonespeciesbutharmtheother(s),includingpredator-preyinteracons,host-parasite

relaonships,andharmfulmicrobialinfeconsofplantsandanimals.Thephenologyofinteracng

speciesiscrucialtotheirsurvival,asthesurvivalofinteracngmemberswillbestronglyinuencedbywhethertheirphenophasesoverlap.Forexample,ifapollinatoremergesfromwinterdormancyand

itsnectarsourcesarentoweringyet,boththepollinatorandtheplantonwhichitdependswillsuer.

Thepollinatorwillhavenofoodsource,andtheplantwillnotproduceseeds.

Biome:Abiomeisagroupofco-occurringplant,animal,andmicrobialcommuniesthatliveinthe

sametypeofclimate,shareawell-denedgeographicarea,areadaptedtoaparcularsubstrateand

levelofnutrientcycling,andexhibitarecognizeablesetofdominantlifeformsandhabits.Examples

include:deciduousforests,woodlands,coastaldunes,shrublands,tropicalforests,boreal(subarcc)

forests,mangroves,grasslands,anddeserts.

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Asyoucansee,phenologycanbedescribedandmeasuredatmulplelevelsofbiologicalandgeographical

organizaon.Informaonfromeachoftheselevelsprovidesfundamentalknowledgeaboutpaernsand

processesinnature.Phenologicalstudiescaninformusaboutthemingandduraonofresourceavailability

inecologicalcommunies,including:whenpollenandnectarareavailabletopollinators;whenfruitsare

availabletofruit-eanganimals(includinghumans!);whenleavesareavailableforherbivorousinsectsand

mammals;andwhetherplantsmustcompetewitheachotherfortheservicesofpollinatorsandseeddispers-

ers.Phenologicalstudiescanhelptoinformagriculturalplanning,diseaseandpestcontrol,andeco-tourism

industries,as wellasthe ancipaonofallergyseasons.Theycanalsodescribehowpopulaons,speciescommunies,andbiomesarerespondingtochangesintheregionalandglobalclimate.Inshort,phenological

studiesmeasurethepaceofnaturesclock,andindoingsocandetectthechangingpulseoftheplanet.

Therstgoalofthisseconistodemonstratehowvariaoninphenologicalpaernscanbedescribedand

measuredatmulplelevels,including:

withinandamongindividuals;

withinandamongpopulaons;

withinacommunityofcoexisngspecies;and

acrosslargegeographicregionssuchasstates, connents,hemispheres,andeventheenreplanet

The secondgoalistohelpyouunderstandhowthephenologicalprogressionofanindividual,populaon,or

communitycaninuenceitsabilitytosurviveandtoreproduce.

Thisbackgroundinphenologicalvariaonwillhelpyoutodevelopaframeworkforunderstandingthevaria

onthatyouobserveinyourbackyard,schoolyard,ornaturalarea.Itwillalsoprovideyouwithsometermi

nologythatwillallowyoumoreeasilytocommunicateyourobservaonswithotherphenologists.Although

weuseoweringphenologyasourprimaryexample,keepinmindthattheprinciplespresentedinthissec-

oncaneasilybeappliedtootherorganismsthatmayormaynotexhibitsimilarphenophases(e.g.,birthing

mesofmammals,migraonandegg-layingmesofbirds,spawningmesofsh,andpollenproducing

mesofnon-oweringplantslikePinetrees).

Phenological variaon within and among individuals - owering curves

Scienstshavedescribedover250,000speciesofoweringplantsintheworld.Althoughsomespeciesower

irregularlyorconnuously(mostlyintropicalbiomeswhereenvironmentalcondionsaremoreorlesscon

stant),mostoweringplantsinthetemperatezoneexhibitconspicuousoweringphenophasesthatstartand

stopatpredictablemesoftheyear.Themingofthesephenophases,andespeciallythepaceofowering

ishighlyvariableamongindividuals,populaons,andspecies.Whilesomespecies(orindividualsorpopula

ons)produceonlyafewowersoverashortperiodofme,othersproducemanyowersoverthecourseof

months.Thesepaernscanbecapturedbygraphsofowering curves,fromwhichitiseasytovisualizethe

datesofkeyphenophasesandtomakecomparisonsamongindividuals(Figure 1).

NoceinFigure1thatthedateofrstowerisApril15,thedateofpeakoweringisMay6,andthedateof

lastower(denedasthedatewhenthelastoweropens)isJune10.Thismeansthattheduraonofow

eringwas8weeks(or56days).IforaldisplaysurveyswereconductedformoreCaliforniaPoppyindividuals

inthesamepopulaon,theresulngoweringcurveswouldrevealvariaoninoweringphenology(Figure

2).

Figure2showsthreehypothecaloweringcurves,eachcurverepresenngadierentindividualofCalifornia

Poppy.First,nocethevariaonamongthethreeindividualsforthedatesofrstower,peakowering,and


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lastower.Theshapesofthethreeoweringcurves

alsodierthebluecurveisthesameshapeasthe

blackcurvebutshiedtwoweekslater,whereasthe

red curve exhibits a dierent shape altogether. Also

nocethateventhoughtheredcurvediersinshape

fromtheblackcurve,thedateofpeakoweringisthe

same.Lastly,nocethattheduraonofoweringis

shortest for the individual plant represented by theblackcurveandlongestfortheindividualrepresented

bytheredcurve.Overall,theoweringcurvesofthese

threeindividualsoverlap,whichmeansthatthesein-

dividualsmightcompeteforpollinaonservice.Alter-

navely, the simultaneous owering of these plants

mighthelptoaractpollinators,astheywillpresenta

moreconspicuousdisplayofowersandprovidemore

nectarthananindividualplant.Theoverlapinower-

ingmesalsomeansthatpollinatorscouldbetrans-

ferringpollenamongtheseplants.

Ifa phenologists goal istoquanfy and to summa-

rize thephenologicalbehaviorof a populaon (such

isoenthecase),asinglegraphthatcontainsthirty

ormoreoweringcurveswouldbeabitoverwhelm-

ing.Instead,themeanandvarianceofaphenophase

ofa populaon canbeexpressedin adierent type

ofgraph.InFigure 3,themeandateofrstoweris

April8for30individualsofthesameCaliforniaPoppy

populaoninSantaBarbara.Theerrorbarsrepresent

thevariaonamongindividualswithinthepopulaon

fordateofrstower;thereisamoderateamountof

variaoninthispopulaon(moreaboutthisshortly).

Ifthepaceandduraonofoweringaretrackedfor

each of these 30 individuals, the same graph could

easilybeconstructedforotherphenophasessuchas

peakoweringandlastower.

Phenological variaon within a populaon what does it mean?

ThevariaoninoweringphenologyamongindividualsinthispopulaonofCaliforniaPoppycouldbeduetodierentsources,including:

en vironmental factorsthemingofanorganismsphenophasescanchangedramacallyasa

directresponsetoenvironmentalcondionssuchastemperatureorthemicrohabitatinwhich

eachindividualplantlives(e.g.,intheshadeofanotherplantvs.directsunlight,inasandysoilvs.

aclaysoil,highlyexposedtocoldwindsvs.protectedfromthem);

genec factors thegeneccomposionofeachindividual(theoweringcurvesofsiblingsor

othercloselyrelatedindividualswilltendtobemoresimilarthantheoweringcurvesofmore

distantlyrelatedindividuals);or

a com binaon of environmental and genec factors.

Figure 2.FloweringcurvesofthreeCaliforniaPoppyindividuals

thatarelocatedatCoalOilPointNaturalReserveinSantaBar-

bara,CA.Datesofrstower,peakowering,andlastowerare

markedbyclosedsymbolsforeachindividual(theblackcurveis

thesameasinFigure1).They-axisshouldbelabeledasnum-

berofopenowers.

Figure 1. Flowering curve for one California Poppy individua

thatislocatedatCoalOilPointNaturalReserveinSantaBarba

ra,CA.Thedatesofrstower,peakowering,andlastowe

aremarkedbyclosedsymbols.They-axisshouldbelabeledas

numberofopenowers.

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While more detailed observaons and experiments

couldrevealtherelaveimportanceofeachofthese

sources of variaon in shaping owering paerns,

wecanalsothinkofthe consequences of variaon in

oweringpaerns.

Paernsofmangandreproducvesuccessinower-

ingplantsandthereforethelong-termsustainabilityofapopulaonarelargelydependentonthem-

ingofowering.Forexample,individualsthatiniate

oweringbeforepollinatorsareacveinthespringare

likelytosuerlowreproducvesuccessduetoalack

ofpollentransfer.Similarly, ifthe beginningandthe

endoftheoweringseasonarecharacterizedbyonly

a few individuals that are owering, the availability

ofpollenduringtheseperiodsmayberelavelylow,

evenifpollinatorsareabundant.Ifindividualplants

cannotaractpollinatorswhenpopulaonsizes are

low(e.g.,atthebeginningandendoftheoweringseason),naturalseleconwillfavorindividualsthatowesynchronously(e.g.,atthemeofapopulaonspeakowering).Inaddion,duringmesoftheseasonwhen

onlyafewindividualsareowering,theprobabilityofmangbetweencloserelavesincreases,andthiscan

reducethegenecqualityoftheseedsproduced.

Ontheotherhand,ifsomanyindividualsofthesamespeciesoweratthesamemethattheycompeteforpol-

linators,thenmanyoftheirowersarelikelytoremainunpollinated.Inthiscase,individualsthatowerslightly

outofsynch(earlierorlater)withthemajorityoftheirlocalpopulaonmaybenet(havehigherpollinaon

success)becausethereislesscompeonforpollinators.Insum,variaoninoweringpaernsamongindividu-

alswithinpopulaonswillaecttheirvisitaonbybothpollinatorsandseeddispersers,andthiswillaectthe

waythatnaturalseleconactstofavorparcularphenophases.

Phenological variaon among populaons

Buildinguponthisunderstandingofphenologicalvariaonwithinpopulaons,wecanconnuetoscaleupou

observaonstocomparephenologicalvariaonamongpopulaons.LetssupposethatstudentsfromSanDiego

SanFrancisco,andArcataeachconductawithin-populaonstudy.Theonsetofspringtypicallyarrivesearliesatsouthernlatudes(SanDiego)andprogressesnorthward(SantaBarbara,thenSanFrancisco,thenArcata),so

thestudentshypothesizethatCaliforniaPoppieswillstartoweringearliestinSanDiego,followedbythenex

northernlocaon(SantaBarbara),andsoon.TheresultsofthishypothecalstudyaregraphedinFigure 4.

TherearetwomajoraspectsofphenologicalvariaonthatareillustratedinFigure4.First,nocethatthestu

dentsdiscoveredalatudinaltrendinoweringmes,justastheyexpected.Thistrendisalsocommonlyob-

servedalongelevaongradients,wherepopulaonsathigherelevaonstendtoowerlaterthanpopulaonsat

lowerelevaons.Thesephenologicalchangeswithlatudeandelevaonarecommonlyobservedinavarietyof

plantsandanimals,andrepresentstrongenvironmentalregulaonofbiologicalacvies,typicallybytempera

Figure 3.Themeandateofrstowerfor30CaliforniaPoppy

individualsintheSantaBarbarapopulaonisApril8.Varia-

onamongindividualsinthepopulaonisrepresentedbythe

heightoftheerrorbarsaboveandbelowthemeanvalue.

Manyplantbiologistsrefertoplantsasphytometersbecausetheirdevelopmentand

reproduconcanprovideexcellentintegraveinformaonaboutambientcondions.


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ture.ThisgeneralprincipleisdescribedbyHopkinsLaw:thedateofrstowering(andotherphenophases)is

delayedbyapproximately4daysperdegreelatudeor120meterselevaon(equivalentto1dayper100fee

elevaon).While these trendsreectthe sensivity ofphenophasesto environmentalfactors,keepinmind

thatgenecfactorsalsoregulatethemingofphenophasesandthatthepaernsweseeinnaturearelargely

theresultofinteraconsbetweenenvironmentaland

genecfactors.

Figure4alsoillustratesdierentlevelsofwithin-pop-ulaonvariaontheSanFranciscopopulaonexhib-

itsthegreatestamountofvariaonamongindividuals

inrstoweringdate,whereastheArcatapopulaon

exhibits theleast amountofvariaon. Anotherway

todescribethispaernisthattheArcatapopulaon

exhibits the greatest amount of synchrony for rst

oweringdate.Aswediscussedearlier,theamount

ofsynchrony withinpopulaonscanhave important

short- and long-term consequences for pollinaon

success,seedproducon,andseedquality.Asyouwill

seeinthenextsecon,phenologicalvariaonamongcoexisngspeciescanhaveimportantconsequences

forcommunitydynamics.

Phenological variaon among coexisng species: community-level phenology

Abriefvisittoanyhabitat,whetheritsabackyard,schoolyard,orpark,canrevealcommunity-levelphenology.

Thecasualobserverislikelytondsomeplantspeciesthatareoweringandotherspeciesthatarenot.Ofthe

speciesthatareowering,somearelikelytobejuststarngtoower(theseplantswillbearalargeproporon

ofclosedowerbudsrelavetotheproporonsofopenowersanddevelopingfruits)andothersarelikelyto

benishingup(theseplantswillexhibitaverylowproporonsofclosedowerbudsandopenowersanda

veryhighproporonofdevelopingfruits).Themoreaenveobserverwillnocethedistribuonofpollinang

insectspeciesamongtheoweringplants,nongwhetherparcularinsectsprefertheowersofcertainplant

species.Similarly,somespeciesofbirdsmighttendtospendmoremevisingcertainplantspecies,foraging

fornectar(hummingbirds)ordispersingfruitsandseeds(manysongbirds).

Arepeatedvisitaweekortwolaterwillrevealchangesthathaveoccurredinthephenologyoftheplantcom-

munity, andprobably in theabundance or assemblage of insect andbird species as well. Community-leve

phenology is quaned as the change over me in the appearance, acvies, interacons, or diversity of the

assemblage of coexisng species in one habitat (Figure 5).

Figure5depictsthephenologyofaporonofahypothecalecologicalcommunity.Foreachofthevespecies

ofoweringplants,thelengthoftheredlinerepresentstheduraonofeachspeciesoweringperiod.Forex

ample,thestarngpointofeachlinerepresentsthedatewhentherstindividualplantinthatpopulaonstarts

owering,andtheendpointrepresentsthedatewhenthelastplantproducesitslastower.Forinsectpollina

Figure 4. ThemeandateofrstowerofCaliforniaPoppy(for30individualsfromeachoffourpopulaons)increaseswith

increasinglatude,fromSanDiegointhesouthtoArcatain

thenorth.TheSantaBarbararesultisthesameasinFigure3.

They-axisshouldbelabeledasdateofrstower.

8

Thereareseveralwaystocalculatevariaonamongindividualsthatcanberepresentedbytherange

barsshowninFigures3and4.Themostcommonesmateofvariaoniscalledthestandarddeviaon.

SeetheAcvityGuideChapterinthisHandbookforinstruconsonhowtocalculatethisandother

measuresofphenologicalvariaon.


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tors(greenlines),insecvorousbirds(insect-eang;orangelines),andfrugivorousbirds(fruit-eang;bluelines),

eachlinerepresentsthestartandendofforagingacvity(pollinang,insect-eang,fruit-eang).Inotherwords,

thelinesshowthestartdateandtheenddateofallmonitoredindividualsinthepopulaon.

Notethatplantsdierfromanimalsinthat,generallyspeaking,itismuchmorelikelythatindividualplantscan

bemonitoredrepeatedlywithinagrowingseason,recordingtherstandlastoweringdateofeachindividua

plant.Withthisinformaon,thestartandenddatesfortheplantspeciesinFigure5could,alternavely,repre

sent the averagedatesofrstandlastower,respecvely.Forexample,wecouldcalculatetheaveragedateof

rstowerifweobservedthisdateinmorethanonedisnctindividualplant.Itsmuchmorediculttodothis

foranimalssimplybecausetheyrehighlymobileanditsdiculttoidenfydierentindividuals.

StarngattheearliestdateinFebruary(youcanimposeavercallineacrosstheenregraphtohelpvisualize

this),onespeciesofinsectpollinatorisacvejustbeforetherstplantspeciesbeginstoowerandbeforethe

rstinsecvorousbirdspeciesbeginstoforage.Movingthedatetomid-Marchwhenfourspeciesofplantsare

owering,therearethreespeciesofinsectpollinators,fourspeciesofinsecvorousbirds,andonespeciesof

frugivorousbirdthatareacvelyforaging.Finallyinlate-Julythereareonlytwospeciesofplantsthatareower-

ing,onespeciesofinsectpollinator,onespeciesofinsecvorousbird,andthreespeciesoffrugivorousbirds.Of

thesethreemeperiods,biodiversityoftheacvecommunityishighestinmid-March.

Figure 5.Graphicalrepresentaonofthephenologicalprogressionofabiologicallydiversecommu-

nity. Each horizontal linerepresents themingand duraonof a parcular kindof phenological

acvityobservedinasinglespecies.Forexample,foroweringplants,eachlinemayrepresentthe

duraonoftheoweringperiodofagivenspecies.Inthiscase,eachlinestartsatthedateofrst

oweramongallmonitoredplantsinthepopulaon,andthelineendsatthedateofthelastower

producedbyanyoftheseplants.Similarly,forthespeciesofinsectpollinators,insecvorousbirds,

andfrugivorousbirds,thelinesrepresenttherstandlastdatesthatanyindividualofthesespecies

isobservedtobeacvelyforaging.Drawingavercallineacrossthegraphatanydaterevealsthediversityandidentyofcoexisng(andpotenallyinteracng)species;notethatdiversityandthe

composionofthecommunitychangesoverthecourseofme.


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Community-level phenology - what does it tell us?

Whatdoesthephenologyofecologicalcommuniestellusbeyonditsapproximatemeschedule?Tostart,it

describestheavailabilityofresourcesinthecommunityandshedslightonwhichspeciesmightbeinteracng

moststronglywithotherspecies.Forexample,thetwospeciesofplantsthatareslloweringinJulywillbe

visitedbyonlyonespeciesofinsectpollinator;ifthispollinatorpreferstoforageforpollenandnectarinone

speciesovertheother,thenfruitandseedproduconoftheneglectedspeciesislikelytosuer.Asamorecom-

plexexample,theabundanceofinsecvorousbirdspeciesishighestinMarchandtheforagingacviesofthesebirdsislikelytodecreasetheabundanceofinsectpollinators.Thismay,inturn,reducepollinaonservicesfo

theplantsthatareinoweratthatme,whichcouldresultinmanyowersbeingleunpollinated.Thisreduc-

oninpollinaoncouldthenleadtoadecreaseintheavailabilityofripefruitsforthethreespeciesoffrugivo

rousbirdsthatappearlaterinthespring.Ifthreespeciesoffrugivorousbirdsareacvelyforagingforalimited

supplyoffruits,thencompeveinteraconsamongthemarelikelytobeintense,andthiscouldpotenally

aectthereproducvesuccessofeitherindividualbirdsoranenrepopulaon.

Overall,ecologicalcommuniesaredynamicassemblagesofcoexisngspeciesthatinteractbothdirectlyand

indirectlyoverme.Thus,thephenologyofcommuniesprovidesameasureoftheirdiversityandproducvity,

thecombinaonofwhichcancontributetothelong-termstabilityofthecommunity.

Phenological variaon among communies (across landscapes, connents, hemispheres, and the globe)

Thedeteconofphenologicalpaernsacrossecologicalcommuniesandacrossbroadgeographicscalesplays

animportantroleinourunderstandingofglobalenvironmentalchanges.Thedevelopmentofnewremotesens-

ingtechnologies(e.g.,satellite-basedobservaons),aswellasnewinformaon-sharingandanalycaltechnolo-

gies,inrecentyearshasallowedfortheintegraonofhistoricallydisparateenvironmentalsciences(moreabout

thislater).Theresulngmuldisciplinaryapproachhasledtothedevelopmentoflargedatabasesthatprovide

layersofinformaononphysical,chemical,andbiologicalprocessesacrossavarietyofgeographicscales.These

Phenologycanalsobeexaminedattheleveloftheowerwithinanindividual.ThisseriesofimagesofCalifornia

Brilebrush(Encelia californica)showsthephenologicalprogressionofonesunower,fromayoungbudonthele

toawitheredowerontheright(awitheredower,alsocalledaspentower,usuallyindicatesthattheowerwas

pollinatedandismaturingitsseeds).Similartothepreviousexamplesthatquanfyanddescribetheamountofvaria-

onofrstoweringdate,therecanbevariaonamongindividuals,populaons,andspeciesfortherateatwhich

individualowersopen,wilt,andmatureseeds.Canyourememberthethree sources of variaonthatarelikelyto

inuencephenologicaltraits?

Youmightbesurprisedtoknowthata sunowerisactuallyacolleconofmanysmallowers(calledaninores-

cence)!Eachofthelargeyellowpetalsispartofanindividualrayower,andeachyellowstrandinthemiddleis

partofanindividualdiskower.Noceinthisphenologicalprogressionthatrayowersmaturerst,followedbythe

outermostdiscowers,andnallytheinnermostdiscowers.Canyoudetectthedierencebetweenfreshandwilted

owersonthissunowerinorescence?Thissameinwardprogressioniscommontomanyspeciesofsunowers-take

acloselookthenextmeyouseesomeinyouryard,atthemarket,andattheoristsshop!

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eortshavehelpedtodevelopa strongerunderstandingofhowenvironmentalcondions,especiallythecli

mate system, aectphenologicalpaernsamong ecological communies andbiomes. Somequesons tha

haveemergedfocusondeterminingwhichcommunitytypesaremostsensivetoenvironmentalchange,which

communiesarethemostimperiled,andhowchangesinonecommunitycanaecttheproducvityanddiver-

sityofanothercommunity.

SUMMARY - WHAT IS PHENOLOGY?

From individuals andpopulaonsto communies andbiomes, thephenologicalaributes of living systems

inuencetheirecologicalinteraconsandthereforetheprobabilityofpersistenceinthefuture.Forexample,

ifspeciesthatdependoneachother(mutualists,suchasaplantspeciesanditspollinator)donotthriveatthe

sameme(synchronously),thenbothspeciesmaysuer.Themingofanorganismsphenophasescanchange

dramacallyasadirectresponsetoenvironmentalcondions,andthiscanhavelargereectsinthecommunity

Forexample,warmerweatherattheonsetofspringcancausetheseedsofmanyspeciestogerminateearlier

thantheyordinarilywould.Someindividualsandspeciesaremoremalleable(orplasc)inthisregardthan

others,andasynchronythatarisesasaresultcanhavecascadingeectsthroughthecommunity.Theproducv-

ityandsustainabilityofecologicalcommunies,whetherwildorfarmed,canbegreatlydiminishedwhenthe

phenophasesofinterdependentpopulaonsorspeciesbecomeasynchronous.Withtheseconceptsinmind,wecannowprovideamoreprecisedenionofphenology:Phenology is the study of the ming of recurring bio-

logical events, the interacon of bioc and abioc forces that aect these events, and the interrelaon among

phases of the same or dierent species.

SYNTHESIS - PHENOLOGICAL VARIATION

Withtheintroduconabove,yourenowwellawareofthewaysinwhichthephenologicalscheduleofanindi-

vidual,populaonorspeciescanaectitsabilitytopersistinnaturalcommuniessuchasforests,elds,and

streams,orinmanagedcommuniessuchasfarms,localparks,orrecreaonalareas.

Forexample,whataresomeways inwhichan individualplantsoweringschedulemayresultinlowerfruit

produconorseedqualitycomparedtothatofothermembersofthesamepopulaon?Floweringwhenpol-

linatorsorpotenalmatesarescarceorwhenower-eanginsectsareparcularlyabundantwillreduce

pollinaonandfruitproducon.Alternavely,oweringatthesamemeasmanycloserelavescanleadto

reducedgenecqualityoftheseedsproducedinthesamewaythatmangbetweencloserelavesinhumans

andotheranimalscanincreasetheriskofgenecdiseases.Delayingoweringunllateintheseason,when

theriskofdrought(indesertsandinMediterraneanclimates)orfrost(inseasonaltemperateclimates)startsto

increasemaypreventanysuccessfulseedproduconatall.Therisksassociatedwithoweringatthewrong

mearemanyandsomemesfatal.

Withtheseideasinmind,youshouldhavenotroubleprovidinganswerstothefollowingquesons:

For a given plant species, how may populaons that are spaally separated from each other benet from be -

ing in phenological synchrony with each other?

Unless p opulaonsaresoisolatedfromeachotherthattheynevercomeincontactthroughseed

dispersalorthroughthemovementofpollenbywindorby animal-transfer,populaonsoen

benetfromreceivinggenecallydisnctseedsorpollenfromotherpopulaons.Thisincreases

apopulaonsgenecdiversityandmayincreasetheprobabilitythatitcanadapttoenvironmen-

talchange.


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Within a given community, what are some addional ways in which one or more species are aected by the

ming of the phenophases of other species?

Pollina torsthateatpollen(e.g.,femalebeesandtheirospring)willonlyhaveafoodsourceif

theyareacveduringtheoweringperiodoftheirfavoredplantspecies.Reciprocally,owering

plantsthatrelyonanimalpollinatorswillonlyproducefruits(andseeds)iftheyowerwhentheir

pollinatorsareabundantandsearchingforthem.

Nectar-e angbirds,bees,andantsdependontheoweringmeoftheirprey;plantsthatare

aackedby thesenectar-robberswillsuerreducedfruitandseedproducon,whileplantsthatowerwhensuchthievesarerarewillenjoyhigherreproducon(assumingthattheirpol-

linatorsareavailable!).

Flower-andbu d-eangbeetles,bugs,mothandbuerylarvae,andotherinsectsdependonthe

oweringschedulesoftheirprey.So,notonlydoesthephenologyofplantsaecttheirexposure

tonectar-thieves,butitalsoaectstheirexposuretoower-andbud-eangspecies.

The phe nologyofmothandbuerylarvaenotonlyaecttheplantsthattheyeat,butithas

strongeectsonthesurvivorshipandreproducvesuccessofthebirdsthateatthem.

Pest-controlli nginsectsandbirdseatagriculturalpeststhat,inturn,candestroycrops.Awell-

knownexampleistheladybug,atypeofbeetlethateatsaphids,whichifnotcontrolledwill

aackyoungowers,stems,andleaves.Ifpest-controllinganimals(ladybugs,inthiscase)are

notacveinsynchronywiththeirprey,anenreseasonofacropcanfailenrely.

In sum,thephenologyofplantsinuencestheabundancesoftheinsectsthateatorpollinatethemaswellasof

thebirdsandsmallmammalsthatdependonthemforfruitsandseeds.Thephenologyofplant-eanginsects

andsmallmammals,inturn,aectsthereproduconoftheirplantprey.Thephenologyofpollinanginsects

aectsthereproducvesuccessoftheplantsthattheyvisit.And,thephenologyofinsecvorousbirdsaects

thelarvaeonwhichtheydependandtheplantsonwhichthoselarvaearefound!

Theanswerstotheabovequesonscanhave importantimplicaonsbothforwildspeciesandformanaged

crops.Thelong-termviabilityofmanywildplantpopulaonsdependsonsuccessfulpollinaon,whichcanbe

limitedifplantsowerbeforeoraerpollinatorsareavailable.Iffruitandseedproduconarediminishedbe-causeofplant-pollinatorasynchrony,foodavailabilitytofrugivorous(fruit-eang)birdsandmammalscouldbe

severelyreduced.Itiseasytoimaginehowfoodwebdynamicsinnaturecouldbealteredbyasynchronyinthe

biologicalcommunity.Thisisalsotrueinthecaseofmanagedcrops,wherealmostallofthefarmedfruitstha

humanseatcomefrompollinaonservicesthatareprovidedbyinsects.Thesynchronyofpollinatoravailability

andoweringinfarmedplantsisthereforecricaltoourabilitytoproducefood.

THE MODERN SCIENCE OF PHENOLOGY

The dawn of a new era in seasonal studies

ModernphenologicalstudiesarethoughttohavebeeniniatedinEuropeinthemid-18thcentury.Beginning

in1736,RobertMarshamkeptdetailedrecordsofIndicaonsofSpringonhisfamilyestateinNorfolk,Britain

withthegoalofimprovingmberproduconbylearningmoreaboutthemingofplantandanimallifecycles

Hisobservaonsincludedtherstoccurrencesofleang,oweringandinsectemergence.Hisdescendents

maintainedtheserecordsunl1947,makingitoneofEuropeslongestphenologicalmonitoringrecords.Though

hewasequippedwithonlyanotebookandawringutensil,themodernaspectofMarshamsapproachwas

thesystemacnatureofhisrecordsformanywildspeciesinoneecosystem(27phenophasesfor20plantsand

animals).

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SimilarapproacheshavebeenemployedbypeopleworldwidesinceMarshamsmenotnecessarilytofol-

lowhisleadorbecausetheirlivelihooddependsonit,butbecausetheyenjoyobservingnatureandpredicng

appearancedateseachyear.Phenologicalstudiestodaycanbejustassimple,takingplaceinbackyards,urban

parks,orwildernessareas,ortheycanbeverycomplexandemployadvancedtechnologiesandtoolssuchas

automatedweatherstaons,inconspicuousvideocameras,automatedenvironmentalsensors,andsatellites,

togetherrevealingglobalandlocalseasonalpaernsandtheintricatemechanismsdrivingthem.

Todays phenological studies: integrang scienc disciplines

Someofthemostrigorousphenologicalstudiesperformedtodayarecarriedoutbyenvironmentalsciensts

represenngawiderangeofapproaches,includingpopulaonbiologists,communityecologists,climatologists

hydrologists,andspecialistsinsatellite-drivenremote-sensing.The integraonof sciencdisciplinesmakes

forparcularlypowerfulstudiesbecausethe site intensivenatureofonetool(e.g.,botanicalinventoriesand

detailedphenologicalstudies)cancomplementthe geographically extensiveinformaonprovidedbyanother

(e.g.,satellites).Letsconsiderthisinmoredetailinthefollowingexample.

Attheconnentalscale,sensorsinNASAsMODISsatellitesinspacemeasuretheamountofsunlightreected

fromtheearthssurface.Leavesreectlightthatisparcularlyrichinwavelengthsinthenear-infraredporonofthelightspectrum,whichthehumaneyecannotseebutthatsatellitesensorscanmeasure.Thereectance

oflightfromleavesisgreatestinmid-summerwhenleavesaremostabundant,andlowestinwinterwhen

grasslands,shrublands,andforesttreesaremostlybare.Thus,satellitesdetecttheonsetofspringbydetecng

arapidincreaseinthereectanceofinfraredwavelengthsthisphenomenoniscalledgreen-up.Alternavely

thesenescenceofleavesintheautumnleadstoadecreaseinreectance,andthisiscalled brown-down.By

collecngtheseremotely-senseddataonaregularbasisandona largegeographicscale,scienstscanaccu-

ratelymeasuretheonsetofspringgrowthacrosstheenreNorthernorSouthernHemisphere.

Atthesamemeasgreen-upisdetectedaroundtheworldbysatellites,plant-watchersonEarthcanobserve

thenedetailsofthisprocess.Families,students,natureenthusiasts,andprofessionalbotanistswhoregularly

Phenology is an ancient science

Phenology is oneoftheearliesteldsofscience,studiedbyhumansformillenniatopredicttheavailability

offoodthroughthecomingsandgoingsofseasons.Earlyhumansdependedlargelyontheirabilitytolo-

cate,idenfy,andprotectedibleplantsduringallmesofthegrowingseason.Thismeansthattheyhadto

befamiliarwithboththevegetave(stems,leaves,androots)andthereproducve(owers,fruits,seeds)

unitsoftheplantsintheirecosystemsothattheycouldidenfythemduringallseasons.Thesamewas

trueforagrariansociees,whosemembersalsocollectedseedsandlearnedtomaximizecropproducon

byopmizingthemingofplanngandharvesng.AnadagecommontosomeNaveAmericantribes

ofeasternNorthAmericastatesthatcornseedsshouldbeplantedinthespringsoonaertheleavesof

nearbyoaktreeshavereachedaparcularsize(i.e.,thesizeofamousesearoraredsquirrelsfoot!*).

Thispracce,basedongeneraonsoftrialanderror,predictstheearliestplanngmesothatthecorn

plantshaveenoughmetofullymaturebeforetheendofthegrowingseason;ifcornseedsareplanted

beforethismetheyruntheriskofrongintheground.Althoughfarmingmethodologiesvarygreatly

today,thecraofmaximizingcropproduconisnolessimportant.Withwidespreadlossofagricultural

landstourbandevelopmentandaglobalhumanpopulaonsizeexceeding6.6billion,themingofplant-

ing,harvesng,andcroprotaonsiscricaltoourabilitytosupplyfoodforlocalandglobaldemands.

* Russell, Howard S. 1980. Indian New England Before the Mayower. University Press of New England. 284 pp. ISBN

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visita naturallandscapecantrackgreen-uponadailyorweeklybasis.

Byidenfyingwhichhabitatsandplantspeciesareleangout,wecan

idenfythephenologicaleventsandspeciesthatcontributemosttothe

infrared reectancevaluesobservedfromspace(thisprocessis called

ground-truthing).

Atanevennerspaalscale(e.g.,fromsquaremilestosquaremeters),

biogeochemicalsensorscanmeasuredailyandseasonaluctuaonsintemperature,precipitaon,atmosphericgasses,sunlight,soilnutrients,

streamow, andother components of theabioc environment upon

whichplantsandanimalsrequireforgrowth.Withthesedataonhand,

scienstscandeterminetheinuenceofenvironmentalfactorsonphe-

nologicalpaerns.

SUMMARY

Overthelong-term,themingandduraonofgreen-upcanbecomparedfromyeartoyearandfromdecadeto

decadetodetectwhetherclimacvariaonaectstheproducvityorgeographicdistribuonofnaturalplant

communies.Sciencdisciplinescanbecombinedtoinvesgatephenologicalpaernsacrossbothbiologica

andgeographicallevelsoforganizaon.Thisintegratedphenologicalresearchcanprovidelayersofinformaon

spanningbothspaceandmethatrevealwhenandhowdierentcommuniesandbiomeschangeasafunc-

onofclimatechangeandotherenvironmentalcondions.Althoughremotely-senseddatacanprovidebroad

measuresofglobalphenologicalpaerns,groundtruthingandbiogeochemicalmonitoring arecricalto ou

abilitytoidenfythefactorsthatdrivephenologicalpaernswithinandamongcommuniesandbiomes.The

synthesisofinformaonfrommulplelevelsofbiologicalandgeographicalscalesisthebestmethodbywhich

wecandiscoverbroadecologicalpaernsandtheprocessesthatdrivethem.

Inthefollowingseconsofthishandbook,wewillhelpyoutodevelopthebotanicaltoolsandrecord-keepingmethodsneededtobecomeanindependentPhenologyStewardwhocancontributetotheNaonalPhenology

Networksgrowingcommunityofground-truthers.

Green-upreferstotheprogressiveincreaseinplantgrowthatthebeginningofspring.Springmeincreases

inplantgrowthcanbetrackedwithgreatsensivitybysatellites,detecngandmeasuringtherateatwhich

theearthsvegetaonproducesitsannualushofleaves.Fromaweeklyseriesofsatelliteimagescovering

theNorthernHemisphereduringthespringmonths,scienstscanseethatspringstartsearlieratsoutherly

latudesandprogressesnorthwards,andspringstartsearlieratlowerelevaonsandprogressesupwardin

elevaon.IntuivelythisshouldmakesensetomanypeoplespringcomesearlierinSouthernCaliforniathanitdoesinNorthernCalifornia,andspringcomesearlierinthevalleysthanitdoesinthemountains.Of

coursetheintricatedetailsofthispaerndependonthetopographyofthelandaswellasmanyphysical,

chemical,andbiologicalfactors,butthebroadgeographicpaernsaresimplyremarkable:whenreviewing

theseriesofsatelliteimages,itappearstoscienstsasthoughagreenwaveisprogressingnorthwardin

latudeandupwardinelevaon!(IntheSouthernHemisphere,thegreenwaveprogressessouthwardin

latudeandupwardinelevaon).

Thelongestknownphenological

monitoring record comes from

theRoyalCourtofKyotoinJapan,

wheretheonsetofoweringfor

cherry trees has been recorded

since705AD.Somewine-produc-

ingregionsinEuropealsoboastrecordsofbloomingandharvest

datesthatextendbackhundreds

ofyears.

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Althoughmanypeoplearefamiliarwithfourseasons(spring,summer,autumn,winter),somere-gionsoftheworldexperienceadierentnumberofseasons.InAustralia,forexample,aboriginals

whohave inhabited theconnent for over50,000 years have developed calendars based onthe

seasonalacviesofplantsandanimalsinrelaontochangesintemperature,wind,andrainpat-

terns.Thesecalendarsactuallynotebetween5and10disnctseasonsdependingonwhichregion

ofAustraliaisbeingdescribed.

Keatley,M.andFletcherT.2003.Phenologicaldata,networksandresearch:Australia.Pp.22-44,InPhenology:AnInte-

graveEnvironmentalScience.Ed.M.Schwartz.KluwerAcademicPublishers.563pp.ISBN1402015801

Who parcipates in ground-truthing phenology?

botan istsexploringforests,elds,streamsides,andcoastlinestomonitorournaturalbiodiver-

sity;

far mers recordingthegrowthoftheircrops(ortheappearanceofweeds!);

park rang ers and land managerswhomonitorthehealthofourforests;

livestock managerswhomovecowsandsheepamongpasturessothattheanimalsgrazeonplantsofaparcularseasonalage;

pa rk visitors, hikers, and backpackersinterestedinspyingthespringmeacviesofplantsand

theanimalsthatdependonthem;

birdw atchersobservingthemigraonsoftheirfavoritebirdsandkeepingwatchonthefoods

thattheyeat;

agriculturalists concernedabouttheappearanceofinsectpeststhateatourcrops;

maple sugar harvesters planningwhentotapmapletreesfortheirsapinthefall;

promote rs of ecotourismwhoneedtopredictthemingofopmumannualwildowerand

autumnleaf-turningdisplays;

ecologi stswhomonitorrodentsandothermammalsthatcarrydiseasessuchasLymeDisease

andHantavirus,andwhosepopulaonsgrowandshrinkwiththeavailabilityoffresh,edible

plantmaterial;

re g htersconcernedaboutthebuild-upoffuel(allgrowingplantmaterial)thataectssum-

mermewildrerisk;

physicians interestedinpredicngtheintensityofallergyseasons;

plant naturalists interested in collecng wild species for herbs, spices, dyes, and medicinal

uses;

home gardeners aimingtopredictthebestmetosowvegetableseedsortotransplantseed-

lings;and

students trackingthephenologyofschoolyardgardenstostarttheirownPhenologyDiaries.

Inshort,withalilebitofbotanicalknow-how,families,students,sciensts,andanyindividualwith

anaeconforplantscanparcipateinphenologicalmonitoringintheirneighborhoods,backyards,

andnaturalareas.


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PHENOLOGYANDCLIMATECHANGE

A (VERY) BRIEF OVERVIEW OF CLIMATE CHANGE

ItiswidelyknownthattheEarthsclimatesystemischangingrapidly.Althoughthereisarangeofopinionson

howbesttomigatetheconsequencesofclimatechange,thereisstrongagreementonhowandwhyitisoccur-

ring.Herearejustafewfactsaboutchangesintemperature,fromthe2007AssessmentReportoftheIntergov-

ernmentalPanelonClimateChange(IPCC):

Global averagesurfacetemperatureshaveincreasedby0.74Cduringthelast100years;thisis

thelargestandfastestwarmingtrendinthelastmillennium.

T he rate ofwarminghasincreasedinrecentdecades.Themostrecent50yearshavewarmedat

nearlytwicetherateseenintheprevious50years,and11ofthelast12yearshavebeenthehot-

testinrecordedhistory(since1850).

Landsurf aceshavewarmedslightlyfasterthanoceansurfaces,andthegreatestratesofsurface

warmingareconnuallyfoundinmid-tohigh-latudeconnentalregionsoftheNorthernHemi-

sphere.

These changes,andmanyothersthatarenotdescribedhere,arelargelyaributabletotheemissionofgreenhousegassesintotheatmosphereand their connuedaccumulaon,includingcarbondioxide(CO2),meth

ane(CH4),nitrousoxide(N2O),anduorinatedgasses(e.g.,hydrouorocarbons,peruorocarbons,andsulfu

hexauoride). There is no doubt in thescienccommunity that human acviessuch as industrializaon

burningoffossilfuels,anddeforestaonhavecontributedtoincreasedconcentraonsofgreenhousegassesand

thereforetoglobalclimatechange.

CLIMATE CHANGE ON A SEASONAL TIME SCALE THE CHANGING GROWING SEASON

Inordertodiscusstheconneconsbetweenclimatechangeandphenology,werstneedtounderstandhow

climatechangeisaecngthemingandintensityoftheseasons.Thetermglobalwarmingoenevokesim

agesofunbearablyhotsummers.Whileitistruethatsummermehightemperaturesarerisingworldwide,itoensurprisespeopletolearnthatinseasonalenvironmentsaroundtheworldthosecharacterizedbycoolor

coldwintersandwarmorhotsummerswinteristhemostrapidlywarmingseason.Infact,bymanymeasures

wintersarewarmingatnearlytwicetherateofsummers.Wherethisoccurs,wintersarenotonlywarmer,they

are also shorter,andthischangecanbeseeninseveralways:

Theamountof methatlakes,rivers,andpolaroceansarecoveredwithiceduringthewinter

isdecreasing.IntheNorthernHemisphere,observaonsofearly-winterfreezingdatesforrivers

andlakesoverthelast150yearshasfoundthatfreezingisbecomingdelayedatarateofnarly

6dayspercentury.Icebreakup,inturn,istakingplaceearlier,atarateof~6.5daysper100

years.

Therearemulplewaystodenethe growing season ofaregion,butmostdenionspointtotheme

ofyearthatisfavorabletoplantgrowth.Thismetriccanchangedependingonhowcold-hardytheplants

inagivenhabitatare,butacommondenionisthemeperiodbetweenthebeginningofspringandthe

occurrenceof therstkillingfrostsof autumn,otherwiseknownasthe frost-freeseason.Theexamples

aboveshowtheeectsofclimatechangeonthephysicalworld(e.g.,snowpack,duraonoflackandriverice,andtheareaofthearccicesheet),butclimatechangehassimilarlystronginuencesonthebiological

world.Onemajordierenceisthat,whilehighlycomplexmathemacalmodelshavebeenquitesuccessful

inpredicngtheeectsofclimatechangeonphysicalglobalacvies(suchasicemelt),theeectsonthe

biologicalworldareimmeasurablydiculttomodeland,asaresult,arehighlyunpredictable.

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Addionally,currentandpredictedratesofwarmingmightoutpacethedispersalratesofmanyorganisms,es

peciallyforthethousandsofplantspeciesforwhichseeddispersalrequiresparcularspeciesofanimalstoeat

theirfruits.Iftheirseeddispersersdonotcarrytheseedstootherhabitats,thenplantswillnotbeabletocolo

nizenewandpotenallysuitablehabitats.

Lastly,foodwebswithinecologicalcommuniesarelikelytochangewhennovelspeciescolonizeahabitat,po

tenallydisrupngresourceavailabilityformanymembersofthecommunity.Forexample,asthenumberand

abundancesofdierentspeciesinanareachangeasspeciesmoveinandoutinresponsetoclimatechangethenumberandtypesofspeciescompengforthesamefoodscanincrease.Whenthishappens,compeon

fora rarefoodresource(suchasripefruitsornewlyemergedinsectlarvae)canbecomeparcularlyintense

Underthesecondions,theriskofexnconislikelytoincreaseformanyspecies(recallourearlierexampleo

frugivorousbirdscompengforripeningfruitsinFigure5).

Temporal responses to climate change

Onatemporalscale,livingorganismsrespondtotheexpandingor shiinggrowingseasonbychangingthei

phenologicalschedules.Thousandsofbiologicalrecordsspanningboththeglobeandthe20thcenturyindicate

anearlyubiquitousshiinspringphenologytowardearliercalendardates.Tosummarize,organismsaretrackingan earlierstarttothegrowingseasonby emergingfromdormancyearlier(mammals, insects),migrang

earlier(birds,insects,mammals),developingnewleavesandstemsearlier(plants),andbreedingearlier(mam-

mals,insects,birds,plants).Notsurprisingly,similardelaysinautumnphenologyareoccurringduetoapersist-

ingsummerdeciduoustreesaresheddingtheirleaveslaterthantheyusedto,andbirdsandmammalsare

migranglaterintheautumnthantheyusedto.

Asyoucanimagine,theconsequencesofphenologicalshiswilldependonthelevelofbiologicalorganizaon

thatweinvesgate.Foralong-livedindividualtree,forexample,leangoutseveraldaysearliereachspringmay

allowittogetarapidheadstartinitsannualgrowth,especiallyifneighboringtreesremainleaess,whichwould

allowitsleavestoreceiveagreatdealofsunlight.Ontheotherhand,ifitproducesitsleavesearlierthananyof

theothermembersofitspopulaon,itmaybeparcularlyvulnerabletoleaf-eanginsects(e.g.,mothandbut-

terylarvae,leafminers,andleaf-cungants)simplybecauseitwillbearareandlimitedresourceforthem.

The Intergovernmental Panel on Climate Change (IPCC)

Aerdecadesofsciencdiscussionaboutthepossibilityofglobalwarming,in1988theWorldMeteo-

rologicalOrganizaonandtheUnitedNaonsEnvironmentalProgrammecreatedtheIntergovernmental

PanelonClimateChange(IPCC).TheIPCCisamuldisciplinarysciencbodythatregularlybringstogether

hundredsofexpertsfromallregionsoftheworldtoassesstheglobalclimatesystem.Theireortsare

broadlyfocusedontwotopics:theassessmentandinterpretaonofsciencinformaon,andtheformu-

laonofrealiscresponsestrategiesatnaonalandgloballevels.Theirndingshavebeenpublishedina

seriesofground-breakingAssessmentReports(1990,1995,2001,2007)aswellasinahandfulofSpecial

Reportsexaminingtopicsingreaterdetail.

TheIPCCsFirstAssessmentReportin1990playedadecisiveroleintheestablishmentofaninternaonal

treatyforclimatechangemigaonin1992,theUnitedNaonsFrameworkConvenononClimateChange

(UNFCCC).TheirSecondReportprovidedkeyinputforthenegoaonsoftheKyotoProtocolin1997.The

IPCCreceivedthe2007NobelPeacePrizeaerthepublicaonofitsFourthAssessmentReport,whichcon-

tainedthestrongestlanguagetodateconcludingthatwarmingoftheclimatesystemisunequivocaland

thatthereisveryhighcondencethattheneteectofhumanacviessince1750hasbeenoneofwarm-

ing.YoucanlearnmoreabouttheIPCConlineatwww.ipcc.ch

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Theeectsofaphenologicalshionagivenpopulaonofaspecieswillalsodependonwhethertheotherspe

ciesonwhichitreliesforfood,pollinaon,orseeddispersalchangewithit.Forexample,ifpopulaons

thatbenetfromeachotherresponddierentlytoclimatechange(e.g.,ifaplantpopulaonowersonaverage

vedaysearlierbutitspollinatorshaventarrived),thenoneorbothofthemmayexhibitpopulaondeclines

Ontheotherhand,ifapopulaonowersearlyenoughtoescapeoneormoreofitsnaturalenemies(e.g.

owerbud-eangbeetlelarvae),itmayreapthebenetsofhighowerandfruitproducon.

Atthecommunitylevel,well,prediconsarehighlyelusive.Wevelearnedfromradio-trackingandbird-bandingeortsthatbirdsarereturningtospringandsummerbreedinggroundsearlierandearlier,buttheeectsof

thesealteredmigraonpaernsonboththebirdsandthespecieswithwhichtheyinteract(aspredatorsoras

seeddispersers)arelikelytobehighlyspecies-andcommunity-specic.Nevertheless,thereareseveralruleso

thumbthatprobablyapply.

First,themorespecializedtherelaonshipbetweenspecies,themorevulnerableeachofthemarelikelytobe

tothephenologicaleectsofclimatechange.Forexample,plantspeciesthatarepollinatedbyonlyonespecies

ofpollinatoraremorevulnerabletolosingthesepollinatorservicesthanarespeciesthatarevisitedbyawide

rangeofpollinatorspecies.Similarly,insectsthatrelyonasingleplantspeciesastheirfoodsourcemayexperi

enceseverepopulaondeclinesiftheiremergencemesaremismatchedwiththegrowingoroweringseason

oftheirhostplant.

Second,animalsthathabituallymigratebetweengeographicallydistantbiomestocompletedisnctporonsof

theirlifecycles(e.g.,growthandsexualmaturaonvs.breeding)areparcularlyvulnerabletothephenologica

eectsofclimatechangesimplybecausetherearemoreinter-specicinteraconsthatmaybedisrupted.Such

speciesmustconnuetoexhibitaphenologicalschedulethatisviablegiventheco-occuringspeciesandthe

climateineachoftheirhomehabitats.Forexample,speciesofbirdsthatmigrateoverlongdistancesmustco-

occurwiththeirfoodsources(whileavoidingtheirenemies)inthebiomeinwhichtheygrowandreachsexua

maturity,andthen,followingmigraontotheirbreedinggrounds,theymustbeinsynchronywiththefood

sourcesthattheyfeedtotheirnewborns.Migratoryanimalsmustmeetthesedemandswhilealsoavoidingthe

naturalenemiesthatdestroythem.

The ngerprint of climate change

These shis ingeographic rangeandphenology arewhat sciensts call thengerprint of climatechange.

Whatatrstappearstobeasituaonwherelifeisaccommodangtoclimatechangebyalteringthepaceand

duraonofitsannualcycleisactuallyquiteworrisomephenologicalschedulesarechangingrapidlyandat

dierentratesforco-dependentspeciessuchasplantsandpollinators.Thereisanincreasingnumberofreports

inrecentyearsofpopulaoncrashesinthewildduetosuchasynchronousresponsestoclimatechangeamong

co-dependentspecies,butweareonlybeginningtolearnhowwidespreadtheyareandinhowmanydieren

kindsoforganismstheyoccur.

A note about the worlds oceans - data from IPCCs 2007 reports

Sea levels are ontheriseworldwide,butnotjustbecauseglaciersandseaicearemelng.Infact,melnggla-

ciersandoceaniceaccountforonly28%and15%oftheriseinsealevel,respecvely.Themostcricalfactorin

risingsealevelsisactuallythewarmingoftheoceanfromsolarradiaonandthegreenhouseeect.Theoceans

areabsorbingover80%oftheheataddedtotheclimatesystem,whichleadstoheatbeingdispersedvercally

throughthedepthsoftheoceans.Asaresult,theaveragetemperatureoftheglobaloceanhaswarmedto

over3000mdeep,raisingscienstsconcernsoverhowoceanicprocessessuchascirculaonpaerns(e.g.,the

NorthAtlancOscillaon)willoperateinthefuture.Notonlythis,butwhenwaterheatsup,itexpandsthis

thermalexpansionoftheoceansaccountsfor57%ofthetotalriseinsealevel.


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PHENOLOGY NETWORKS

A GATEWAY TO DETECTING ENVIRONMENTAL CHANGE

ShortlyaerRobertMarshaminiatedmodernphenologicalsurveysonhislandinBritain,theSwedishbotanist

CarolusLinnaeus(1707-1778)revoluonizedtheeldofphenologybyconducngastudysimultaneouslyin

manylocaons.In1750,Linnaeus,thescienstwhoinventedtheclassicaonsystemsllusedtodaytoname

andtodescribeallplantsandanimals,selectedmulplesitesinSwedenwhereherecordedannuallythedates

ofleang,owering,fruing,andleaffallformanyspecies,alongwiththeweathercondionsatowering.He

describedhismethodologyinhiswork Philosophia Botanicasothatotherscouldreplicatethestudy.Thiswas

therstblueprintforthecreaonofaphenology network,whichmanyscienstsandcizensaroundtheworld

thenimplementedatdierentgeographicscalesoverthefollowingdecadesandcenturies.

Thegoalof aphenologynetworkistocoordinateagroupof peopleto monitorsimultaneouslyphenologica

eventsacrossawidegeographicareaandtosharetheirinformaon.Observersacrossacountryorconnent

relyonacommonsetofdenionsandprotocolsinordertostandardizetheirobservaonsacrossregions

andformanydierentspecies.Thesestandardizedstudiescanhavemanyuseshistoricallythemostcommon

goalhasbeentounderstandandtopredictvariaoninthelifecyclesofplantsandanimalsforagriculturalap

plicaons.Apraccalknowledgeofphenologicalpaernshasalsoallowedhunterstopredictgameavailability

enabledcityandlandmanagerstoplanforresourceuse,andpermiedsocieestoplanculturaleventsaround

naturalphenomena.

One consequence of asynchronous phenological responses to climate change among interacng species

TheDutchPiedFlycatcher(Ficedula hypoleuca)isasmallpasserinebirdthatmigratesfromWesternAfrica

toEuropeeachspring.Uponitsarrivalatitsnorthernbreedinggroundsitconsumescaterpillars.Thecat-

erpillarshavebeenrespondingtowarmerspringtemperaturesbyemergingasadultsearlierinthespring,

butsomeFlycatcherpopulaonshavenotmigratedearliertokeeppace.ThesemismedFlycatcherpopu-

laonshavebeeninsteepdeclineinrecentyearsbecauseoftheasynchronyoftheirmigraonphenology

withthephenologyoftheirfoodsourceatitsbreedinggrounds.

Photo freely available from Aelwyn and the Wikimedia Project.

CC-BY-SA terms of usage available at www.wikimedia.org

Visser,ME,C.Both,andMMLambrechts.2004.Globalclimatechangeleadstomismedavianreproducon.AdvancesinEco-

logicalResearch35:89-110.

Durant,J.M.,D.OHjermann,G.Oersen,andNCStenseth.2007.Climateandthematchormismatchbetweenpredatorre-

quirementsandresourceavailability.ClimateResearch33:271-283.

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PhenologynetworksblinkedonandothroughoutEuropeandtheworldduringthelate-19thand20thcentu

ries.Manyofthemwerephasedoutduetowars,ageneraldecreaseintheirapplicability,oralackofgovernment

support(oracombinaonofthesefactors).Somesuccessfulnetworks,suchastheCanadianeortthatwasini-

atedasearlyas1890,provedtheirworthinthebotanicalorclimatologicalsciencesandthenwereexpandedby

eitherindividualscienstsornaonalmeteorologicalagencies.Thus,thedominanttrendwastowardnaonally-

coordinatedphenologicalobservaons,buttherewasanunfortunatelackofcoordinaonamongnetworksa

theinternaonallevel,especiallyacrossEurope.Asaresult,monitoringprotocolshavevariedamongnetworks

andcountries,anddatasetshavebeendiculttomerge,therebylimingthescopeoftheirstudies.

TherstnotableexceponstothislackofintegraonweretheInternaonalPhenologicalGardens(IPGs).The

IPGswereestablishedinseveralEuropeancountriesinthelate1950sbasedonthegrowingneedtopredictagri

culturalseasons.Recognizingthatgenecallybaseddierencesinphenologicaleventsamongplantpopulaons

collectedfromthewildcanoverwhelmtheenvironmentalsignalobtainedfromphenologicalobservaonsoflo-

calpopulaons,scienstspropagatedclonesof14speciesoftreesandshrubsandsentthemtogardenslocated

fromItalytotheNetherlands.Moregardenswereestablishedinsubsequentyears,andover50ofthemaresll

beingmonitoredtoday.Withnogenecvariaonamonggardens,eachspecies,throughitsphenologicaldevel

opment,hasrevealedsubstanalgeographicvariaoninseasonality.Collecvely,thephenologicalinformaon

providedbythesegardenshasledtothecreaonofclimatemapsaswellastopredicvecomputermodelsfo

theimprovementofagriculturalpracces.Even more,theInternaonalPhenologicalGardenshaveplayedacricalroleindetecngtheeectsofclimatechangeonthephenologyofplants.Infact,thestudymenoned

previouslyrevealinga10.8dayexpansionoftheEuropeangrowingseasonwaspublishedbyIPGscienstsde-

scribingthephenologyoftheirclonedtreesandshrubs.

Recognizingthecontemporaryvalueofthesedataandoftheremaininginfrastructureofindependentlyfunc-

oningphenology networks throughoutEurope,the European PhenologyNetwork (EPN) was establishedin

How does a clonal phenological garden work?

Earlierinthischapteryoulearnedhowscienstsquanfyphenologicalvariaonatavarietyofbiological

andgeographicalscales.Youalsolearnedthattheobservedvariaoninphenologyamongindividuals,populaon,andcommuniescanbeduetothreesourcesofvariaon:environmentalfactors,genec

factors,oracombinaonofenvironmentalandgenecfactors.Wecanexpressthisconceptinthefol-

lowingmathemacalequaon:

V=G+E+GxE

whereVistheobservedvariaonforatraitwithinapopulaon(suchaswithdateofrstowerinour

earlier examples),Gistheamountofvariaoninthetraitduetogenecdierencesamongindividuals

(e.g.,siblingstendtooweratasimilarme,therebyreducingtheamountofgenecvariaonwithin

thepopulaon),Eistheamountofvariaoninthetraitduetotheinuenceoftheenvironment(e.g.,

warmercondionstypicallyinduceearlierowering),andGxEistheinteraconbetweengenecand

environmentalfactors(e.g.,somegenotypes[individuals]inthepopulaonaremoresensivetowarmtemperaturesthanothergenotypes).

The sciensts whoestablishedtheInternaonal Phenological Gardens(IPGs) usedthis knowledge to

theiradvantage.ByplanngclonedtreesandshrubsindierentregionsofEurope,theycausedallge-

necvariaonamonggardenstobezero.Basedontheaboveequaon,then,allobservedvariaonbe-

tweengardensintheirphenologicalpaernsmustbeduetoenvironmentalfactors(E).Thisexperimental

approachisusedoentodaybyevoluonaryecologistsandecologicalgenecistswhoareinterestedin

understandingtheinuencesofgenecdiversityandenvironmentalchangeonecologicalinteracons

(suchasplant-pollinatorandpredator-preyinteracons).


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2000bythePhenologyStudyGroupoftheInternaonalSocietyofBiometeorology.TheEPNsgoalsweretocre-

ateaplaormfromwhichtocoordinatephenologicalmonitoringeortsandtostandardizeobservaonsacross

Europetomonitormoreecientlythechangingclimate.Othernetworksaroundtheworldhavealsodeveloped

orresurfacedoverthepastdecadeunderthesamecontext.Today,phenologynetworksexistinNorthandSouth

America,Europe,EastAsiaandAustralia.

PHENOLOGY NETWORKS AND THE CITIZEN SCIENTIST MOVEMENT

While thedatabases createdby contemporary phenology networks arelled by professional scienstswho

observeandreportthemingofseasonalevents,somenetworkshaveinvitedcizen scienststoparcipate

Theseincludenatureenthusiasts,homegardeners,parkdocents,botanicalgardensta,farmers,NaonalPark

rangers,families,teachers,andstudents.Whenpeopleacrossawidegeographicareaaretrainedtoobserve

andrecordphenophasesintheirbackyards,localparks,eldstaons,andnearbywildareas,thenumberof

observaonssubmiedtoadatabasecanincreasedramacally.Thisprocessalsoengagesthe publicso tha

theybecomemoreawareofnatureandofthemanifestaonsofclimatechange.TheEuropeanPhenologyNet-

work,NaturesCalendarinBritain,NatuurkalenderinHolland,andNatureWatchinCanadahaveeachhadgreat

successinrecruingthepublictoreportthemingofphenophases.Thedataaresynthesizedbyscienstsand

resultsarereportedinavarietyofwaysonthenetworkswebsites.RecenteortsintheUnitedStateshaveledtothedevelopmentofanaonwidephenologynetworkthatinvitesparcipaonfromprofessionalandcizen

sciensts.

A BRIEF HISTORY OF PHENOLOGY IN THE UNITED STATES

Afewaemptswere made inthe early20thcenturyto iniate

coordinated phenological measurements in parts of the United

States,butitwasntunlthemid-20thcenturythattheconcept

tookroot.Inthe1950s,movatedbythenoonthatthesimul-

taneousmonitoringofawidespreadspeciescouldbeusedasan

indicatorfortheprogressionoftheagriculturalgrowingseason,

Montana State University agricultural climatologist Dr. Joe M.

Capriostartedwhatwouldeventuallybecomethenaonsphe-

nologicallegacy.CapriosoughtobserversacrosstheWesternUS

whowouldreporttohimthelocalmingofleangandowering

oftheCommonLilac,Syringa vulgaris.By1957hehadrecruited

over2500volunteersin12westernstates,andwithinadecade

headdedtwohoneysuckleculvars(Lonicera tataricaArnoldRed

and L. korolkowii Zabeli)totheobservaonprogram.Observa-

Figure 6. Locaons in North America

withveyearsormoreoflilacphenology

data,1956-2001(Qubecstaonsnotin-

cluded).FromSchwartz2003.

Manyculturesthroughouthistoryhaveusedoweringmesascuesforhunngandculturalacvies.In

thePacicNorthwest,themanynaonscomprisingtheNuu-Chah-NulthofwesternVancouverIslandhave

lookedtotheripeningofsalmonberries(Rubus spectabilis)toalertthemthatanadromoussockeyesalmon

(thosethatmigratefromsaltwatertofreshwater)areabundantinnearbystreams.TheBlackfootofAlberta

wouldmetheirannualmid-summertribalceremonytotheripeningofSaskatoonberries(Amelanchier

alnifolia),andreportedlywouldmetheirspringhuntforbisonbullstotheoweringmeoftheBualo

Beanplant(Thermopsis rhombifolia).

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onsinthiswesternUSnetworkconnuedthrough1994atwhichmeCaprioreredandmanyoftheobserv-

ersdroppedoutastheyagedanddied,underscoringthefactthataphenologicalnetworkisalivingsystemthat

mustberepopulatedeachgeneraontoensureitslongevity.

CapriosearlysuccessmovatedtwootherresearcherstoiniateregionalphenologynetworksinNebraska(W.

L.Colville,UniversityofNebraska)andintheNortheast(R.Hopp,UniversityofVermont)inthe1960s.Usingthe

sameapproachastheInternaonalPhenologicalGardensinEurope,thesetwonetworksrecordedobservaons

foraclonedculvaroflilac, Syringa chinensis(RedRothomagensis)atabout300sites.Theyalsocollecteddata

ontheaforemenonedhoneysuckleculvars.Administraonofthesetwonetworkswasjuggledforacoupleofdecadesunlfundingwasulmatelylostin1986,aerwhichUniversityofWisconsin-Milwaukeeclimatologist

andgeographyprofessorDr.MarkD.Schwartzcontactedtheirvolunteernetworkstomanagetheremaining

monitoringacvies.

Thedatafromthelilacandhoneysucklephenologynetworkshaveprovenusefulonmanycounts.Schwartzhas

developedsophiscatedcomputermodelscapableofpredicngtheonsetofspringacrosstheUnitedStates

fromacombinaonofphenologicalandclimatologicaldata.Thephenologicaldatahavealsohelpedtodem-

onstratethatspringinNorthAmericaisarrivingearlierthaninthepast,andSchwartzhasesmatedthatthe

onsetofspringisadvancingby1.2daysperdecadeacrosstheNorthernHemisphere.Thereisevenpreliminary

evidencethatphenologicalschedulesare,insomeregionsinthewesternU.S.,stronglycorrelatedwiththesize

andfrequencyofwildresduringthesubsequentreseason.

SchwartzandothershaverecentlyleveragedthisphenologicallegacyintheU.S.to rallysciencandpublic

interestaroundanaonal-levelphenologynetwork.The USA Naonal Phenology Network (USA-NPN) is cur-

rentlybeing designedand organizedtoengagefederal agencies, environmental networks andeld staons,

educaonalinstuons,andmassparcipaonbycizensciensts.Itisanaonwidecollaboraonamongpro-

fessionalandcizenscienstsdesignedtotracktheprogressionoftheseasons.

THE USA NATIONAL PHENOLOGY NETWORK (NPN)

ThevisionoftheUSANPNistointegrateavailabledatafromexisnginfrastructures(e.g.,NASAssatelliteim-

agery, NaonalWeather Service Co-Op staons, Naonal Park Services annual inventories, andAmeriFluxs

biogeochemicalmonitoringstaons),whileengagingcizenscienstsacrossthecountrytomonitorlocalorgan-

isms.ThecomponentoftheNPNthatisacvelyworkingtoinformtheAmericanpublicandsolicittheirhelpis

theEducaon,CizenScience,&Outreachworkinggroup(ECSO).

The Educaon, Cizen Science & Outreach (ECSO) working group of the NPN

Intheglobalcampaigntoreducethefactorsthatpromoteglobalwarming,therearemanyorganizaonsthat

providecizenswithopportuniestodonatemoneyormewiththegoalofreducingenergyuseandgreen-

Figure 7.PhenologicalprogressionforthecommonLilac, Syringa vulgaris.Leafbudburst(le)oc-

cursearlyinthespring,therstowersopenaboutonemonthlater(middle),andfullowering

occursafewweekslater(right).PhotoscourtesyMarkSchwartzandtheUSANPN.


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housegasemissionsthrougheducaonandoutreach.Thereareonlyafeworganizaons,however,thatinvite

cizenstocontributetheirmeandenergytowardsciencmonitoringandthedeteconoflocalclimate-

relatedpaerns.TheECSOworkinggroupoftheNPNoperateswiththeunderstandingthatphenology,andthe

abilitytomeasureandtorecorditsprogressovertheseasons,isaccessibletoeveryoneinthecountry.Thepri-

marygoalofECSOprogramsistobringtogetherprofessionalandcizensciensts,students,stewardsofpublic

andprivatelands,andoutdoorenthusiaststocompilebasicinformaononphenologicalresponsestoclimate

change;tostudyitsnature,paceanditseectsonecosystemfuncon;andtounderstandsynchronyandeco

logicalinteraconsamongspecies.

TheNPNsECSOworkinggrouphasdesignedarangeofprogramsandproductsthroughwhichthepublicand

sciencprofessionalscanobserveandinterpretthephenologicalworldonanunprecedentedscale.Eachpro

gramisdesignedtoachieveoneormoreofthefollowingobjecves:

To engage the publicinlong-termphenologicaldatacolleconandanalysisthroughbothformal

andinformalscienceeducaonprograms;

To enge nderself-directed,voluntarylearningusinginquiry-basedapproaches;

To engage currentlyacvelandstewards,networks,andmasternaturalistprogramsinthecollec-

onandcontribuonofphenologicaldatatotheNPN;

Toprovide traininginthetoolsandapplicaonsofphenologicalstudiestocizensandtoscien-stsatallstagesintheircareers;

To enha nceopportuniesforthepublictointeractwithprofessionalsciensts;

Topromot enoveltooldevelopmentandcollaboraonamongscienstsrepresennghistorically

disnctdisciplines(e.g.,climatology,hydrology,remote-sensing,engineering,ecology,popula-

onbiology,andevoluonaryecology).

PROJECT BUDBURST THE NPNS FIELD CAMPAIGN FOR CITIZEN SCIENTISTS

Thison-line educaonal anddata-entry program is ECSOs agship naonwide program forthe public

(hp://www.budburst.org).ThesitewentliveonApril1,2007,targengstudents,natureenthusiasts,and

theirfamilies.Thesiteprovides:informaononthehistoryandecologicalsignicanceofphenology;its

linktoclimatechange;materialsforteachersandstudents;andaneasy-to-useonlineulityforuploading

botanicalphenologicaldataobtainedinbackyards,botanicgardens,naturepreserves,andthewild.For

the2007implementaonofProjectBudburst,60widespreadandeasilyidenablewildandculvated

speciesweretargetedforobservaonsthroughouttheU.S.Conveniently,thedatamanagementsystem

allowsuserstoenterphenologicalobservaons(includingthedatesofbudburst,fullleaf-out,rstower,

peakowering,andcompleonofowering)foranyspecies.BytheendofthersteldseasonforProject

Budburst(June2007)almost1000phenologicalobservaonsweresubmiedfromalloverthecountry.The2008seasonofProjectBudburstkickedoinJanuaryandisexpectedtoaractawideraudienceof

cizenscienst parcipants.ProjectBudburstparcipantsarenow ableto establish a useraccount at

www.budburst.org,wheretheycanuploadaswellasmanagetheirdatathroughouttheseason.

Visit www.budburst.org today!

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THE UCSB PHENOLOGY STEWARDSHIP PROGRAM

AttheUniversityofCalifornia,SantaBarbara(UCSB),wehavedesignedaprogramtorecruitandtoengagelocal

families,teachers,natureenthusiasts,andstudentsatalllevelsinphenologicalmonitoring.The UCSB Phenol-

ogy Stewardship Programisagrowingnetworkofobserversdedicatedtotrackingandrecordingseasonalbio

logicaleventsintheSantaBarbararegion.WithfundingfromUCSBsCoastalFund,thePhenologyStewardship

ProgramwasiniatedintheSpringof2007byPhDstudentBrianHaggertyandProfessorofEcology&Evoluon

Dr.SusanMazer.Throughacombinaonofsciencresearch,environmentaleducaon,anddynamicoutreachpresentaons,theirmissionistoengageCaliforniaresidentsinthecolleconofphenologicaldataandthere-

porngofthoseobservaonstotheProjectBudburstonlinedatabase.

TheUCSBPhenologyStewardshipProgramprovidesinternshipsforundergraduatesinsciencresearch,envi

ronmentaleducaon,graphicdesign,andecologicalmapping.TheProgramalsoprovidestrainingandteaching

materialsforK-12teacherstobringphenologicalobservaonanditsinterpretaonintotheclassroomandout-

doorschoolyardorphenologygarden.AcviesalsoincludetheuseoftheherbariumatUCSBsCheadleCenter

forBiodiversityandEcologicalRestoraontodemonstratetheusefulnessofherbariumrecordsforreconstruct

inghistoricaloweringmesanditsrelaonshiptoclimate,elevaon,latude,andhabitat.

UCSBs Coal Oil Point Natural Reserve (COPNR) isthelocalsiteforphenologicalresearch,training,andoutreach.Here,aseriesofPhenologyTrailshavebeenestablishedthataretrackedbyundergraduatePhenology

Stewards(researchinterns)whocollectoweringdatafrom21plantspeciesalongwiththemigraonandresi-

dencemesofshorebirdsandpasserines.

LookfortheUCSBPhenologyStewardshipProgramonlineatwww.Facebook.comandvisitourpreliminaryweb-

siteat:hp://www.uweb.ucsb.edu/~cmcosner/phenology/

Figure 8.ImagesofUCSantaBarbarasCoalOilPointNaturalReserve.Ontheleisasatelliteimage(fromGoogleEarth)

ofa poronof thereserve;redlines markthe Phenology Trailsthathavebeenestablishedinthreehabitatsincluding

foredune,backdune,andcoastalblu/scrub.Thebodyofwaterintheupper-lecornerisaporonofDevereauxLagoon,

whereavianphenologicalsurveysareconducted.Ontherightisaphotograph(May2007;courtesyMaWilson)ofthe

backduneatCoalOilPoint.WiththePacicOceanasabackdrop,oweringphenologyof21plantspeciesismonitored

byPhenologyStewards(includingthepurple-oweredBeachSandVerbena,Abroniaumbellata,andtheyellow-owered

CaliforniaPoppy,Eschscholziacalifornica.NotethelocaonofSantaBarbara(reddot)ontheinsetmapofCalifornia.


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TheUSA-NPNPlantPhenologyPrograms:

TheUSA-NPNNaonalCoordinangOce:

USA-NPNNaveSpeciesObservaonProgram:

ECSOContacts:

ProjectBudburst:

www.uwm.edu/Dept/Geography/npn/

www.usanpn.org/

www.uwm.edu/Dept/Geography/npn/naveintro.html

www.usanpn.org/workinggroups/outreach.html

www.usanpn.org/about/contact.html

www.budburst.org

Where to obtain more informaon about the USA Naonal Phenology Network and ECSO:


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28

PROTOCOLS FOR PHENOLOGICAL MONITORING

INTRODUCTION

Allphenologicalstudiesshareonesimplegoal:toobtainprecisedatesofkeylifecycleevents.Thereis,however,

substanalvariaonamongspeciesinhowtheycompleteagivenphenophaseandthiscancomplicatethepro

cessofidenfyingandassigningadateforagivenphenophase.Considerthedateofrstowerdoesaplantreachthephaseofrstowerwhenthepetalsofitsrstowerbegintobecomevisibleasthebudopens,oris

itwhentheowerisopenenoughtoseetheanthersorsgmainside(thecorrectansweristhelaer)?Howdo

wedeterminethisphenophasefordierentspecieswhoseowersopenindierentwaysoratdierentrates?

Furthermore,notallowershavethesamereproducvestructuresinside,soourdenionofrstowermus

alsoaccommodatestructuraldierencesamongspecies.

Asanotherexample,considerthespaalarrangementorarchitectureoftheowersproducedbyaplant

Someplantsproduceasingleoweroneachoweringstemwhereasothersproducelooseordenseclusters

(calledinorescences)ofafew,ofdozens,orofhundredsofowersperstem.Withininorescencesofsome

species,theindividualowersmaybesosmallthatevenamagnifyingglasscannotmakethemclearlyvisible.

How,then,shouldwequanfythedateofrstowerforaplantwhoseindividualowerswecannotevensee!?

Theanswerstothesequesons,andmanymore,areprovidedinthischapter.

Thegoalofthischapteristoprovidedenionsofphenophasesforoweringplantssothatobservaonscanbe

standardizedamongobservers.Giventhetremendousvariaoninplantarchitectureamongspecies,wehave

madeaneorttofacilitatetheidencaonandrecordingofphenologicalinformaon.First,wehaveaimedto

minimizethenumberofdenionsandtopresenttheminamannerthatisintuivetoallreaders,regardlessof

theirbotanicalexperience.Second,wehaveidenedseveralclassesofplantarchitecture,eachofwhichhasits

ownsetofprotocolsfortherapididencaonandmeasurementofitsphenophases.Mostspeciesofplants

caneasilybeassignedtooneofthesearchitecturalclasses,soitwillbeasimplemaerforreaderstondthe

protocolthatismostappropriatefortheindividualplantorspeciesthattheyareobserving.We have condensedall phenophase denions into a 3-page set of reference tables that we call the Phenophase Quick-Guide

which is located at the end of this chapter.

ThemethodsanddenionspresentedherehavebeentestedbyStewardsintheUCSBPhenologyStewardship

Programandwerecondentthat,withalilepracce,evennoviceswillsoonbeabletoidenfythepheno-

phasesofmostspeciesthattheyencounter.Inthischapterwewillconsidergeneralpoints,suchaswhereand

whentoobserveplantphenology,aswellasspecicpointssuchasphenophasedenionsbywhichphenolo

gistscanstandardizetheirobservaonswithotherphenologistsintheSantaBarbararegionandacrossthecoun

try.

PLANT PHENOPHASES

Beforeweaddressgeneralpointsaboutphenologicalmonitoringsuchassiteandplantselecon,wewillrstre-

mindthereaderaboutplantphenophases.Althoughthereisconsiderablevariaonamongoweringplantspe-

cieswithregardtowhenandhowrapidlyseasonaleventsoccur,thegeneralizedplantphenologicalsequence

canbesummarizedasfollows:

leafbudburst

rstfullleaf(rstleafisfullyexpanded)


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enreplantleaf-out(allleavesontheplantareexpanded)

rstower

peakowering(largestoraldisplay)

lastower

fruitmaturityandseeddispersal

leafsenescence(colorchangeandabcissiontheprocessoflosingleavesattheendofthe

growingseason)Keepinmindthatnotallplantsexhibiteachofthesephases,andnotallplantscompletephasesinthisorder.

Moreover,thesephasescanoverlapsubstanallywithinagrowingseasonbothwithinandamongindividu

als.Forexample,incontrasttodeciduoustreesandshrubs,evergreenspeciesdonotdropalloftheirleavesin

theautumnnordotheyexhibitthesimultaneousproduconofnewleavesinthespring(leafbudburst).Also,

manywindpollinatedplantssuchasbirchandaldertreesproduceowersbeforeleafbudburstoccurswithout

leavestobreakupthewindcurrents,pollentransportismuchmoreecient.Addionally,fruitsoenbecome

fullyripe(andtheseedsreadyfordispersal)midwaythroughthegrowingseasonbeforeleaveshavesenesced

andoenbeforeoweringhasstopped.InSantaBarbara,California,wherethereislessseasonalchangein

climaterelavetomanyotherregionsoftheU.S.,manyspeciesofplantsowerformonthsatameleading

tooverlapbetweenleang,owering,andfruingphenology.Keepthisinmindasyouconnuereadingand

thinkingaboutstarngyourownmonitoringeorts.

A FEW CONSIDERATIONS ABOUT PHENOLOGICAL MONITORING

Consistency and accuracy are crical

Thesuccessofallphenologicalstudiesdependsontwothings:consistentobservaons,andaccurateidenca

onsofphenophases.Especiallyinthecaseofaphenologynetworkwherehundredstothousandsofobservers

aresimultaneouslyrecordingthedatesofphenophasesforthesamesetofspecies,consistencyisthekey.Herewedescribehowtobestestablishandmaintainyourphenologicalstudies,andlaterweprovideyouwiththe

explicitinstruconstoaccuratelyascribedatestophenophases.

Where To Do Phenological Monitoring

Theshortansweris:anywhere!Natureisallaroundusalltheme,wejusthavetomakeaconsciouseortto

look.Therststepistobecomefamiliarwithyournaturalsurroundingsbyexploringtheoutdoorsregularlyand

withakeeneyetowardsweek-to-weekandday-to-daychangesintheabundancesanddiversityofplantsand

animals.Phenologicalobservaonscantakeplaceinurban,rural,andwildenvironments;parksandbackcountry

trails;schoolyards,backyards,andprisonyards;andbotanicgardens,zoos,anduniversies.Evenalongtheroadandbikepathswhilecommungtoandfromwork.Themoreyoustartlooking,themoreyouwillnocehowna-

tureisdynamicbutfarfromrandom.Eachyear,thesequencesinwhichplantspeciesower,mammalsemerge

fromhibernaon,insectsemergefromdormancy,andbirdsmigrateoenoccurinrepeatablepaerns.

Themoreappropriateanswertowheretodophenologicalmonitoringis:Itdepends.Ifyourgoalsaresimply

tobeeracquaintyourselfwiththeseasonalityof your region,then anywhere willdoyourneighborhood

outsideyourocebuilding,oralongtheroadwayorbikepathonwhichyoucommutetowork.If,however,your

goalsaretocollectaccuratephenologicaldatatocontributetotheProjectBudburstdatabase,thenyoullneed

tochooselocaonsbasedonafewfactors,including:


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theconvenienceof,andaccessto,thesitemakesurethatyouarenottrespassingandthatits

alocaonthatseasytovisitevenwhenyourweeksareheccoryourdisposionislazy;

thepermanenceofthehabitatifthatopenspaceisslatedfordevelopmentinacouplemonths,

itsbesttolooktowardprotectedareas,parks,botanicgardens,andbackandfrontyards;and

theinuenceofhumanacvitymakeeveryeorttominimizethis source of variaon.Ifacity

parkismanicuredtothepointthatyouneverseedriedstemsandfruits,orifshrubsaretrimmed

regularlysothatleangandoweringareconstantlydisrupted,gosomewhereelse.Youmay

wishtoavoidlocaonsthatyouknowareheavilywatered,ferlized,orsprayedwithchemicalsbecausetheseacviesimpactphenology(andpossiblyyourownhealth);attheveryleast,take

noteofthesehuman-directedmodicaonsofyourplantshabitatssothatyoucanincludethem

whenyoucontributeyourobservaons(youllhavetheopportunitytocommentonthesefactors

whenuploadingyourdatatotheProjectBudburstdatabase).

When To Do Phenological Monitoring

Theonlythingthatdoesntchangeaboutthenaturalworldisthatit isconstantlychanging!Thatmeansyou

canobservephenologyduringanymeoftheyear.Thespringandautumnareparcularlyrewardingperiods

becausethebiospherechangessoquickly,buteveninthemiddleofwinterorduringthelongdaysofsummethenedetailsofplantdevelopmentcanbesurprisingandfascinang.Currently,thenaonaldatabasehosted

attheProjectBudburstwebsite(www.budburst.org)isfocusedonspringgreen-upandowering,andinvites

anycarefulobserver

The Phenology Handbook-Haggerty Mazer 2008 v1

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Transcript of The Phenology Handbook-Haggerty Mazer 2008 v1