ARTICLES

Lazaroa ISSN: 0210-9778

http://dx.doi.org/10.5209/LAZAROA.51054

ForesttypologyofbroadleafforestsfromSierraMaestra,EasternCuba

OrlandoJ.Reyes 1

Received: 28January2015/Accepted:23March2016

Abstract. AphytosociologicalstudyoftheforestsfromSierraMaestraisconducted,followingthemethodologyof theZurich-MontpelierSchool.TheyaretransformedintoaforesttypologyusingthestandardsoftheInstituteofAgro- ForestryResearch.Ingeneral,35typesand/orsubtypesarepresented.Fromthisgroup,themostabundantonesbelong tosemi-microphyllforest,followedbythosefrommangrovesandmountainrainforest,respectively. Silviculturaltreatmentsareneeded;amongthem,theprotectionforestsarethosefoundabove800maslandman - groves. Keywords: Forests;foresttypology;SierraMaestra;EasternCuba.

[es]TipologíaforestaldelosbosqueslatifoliosdelaSierraMaestra,CubaOriental

Resumen. UtilizandolametodologíadelaEscuelaZürich-Montpellierserealizaunestudiofitocenológicodelos bosquesdelaSierraMaestra,elcualestransformadoenunaTipologíaForestalusandolosnormasdelInstitutode InvestigacionesAgro-Forestales.Engeneral,sepresentan35tiposy/osubtipos,deloscualeslosmásabundantes pertenecenalosbosquessemideciduosmicrófilos,siguiéndolelosmanglaresylapluvisilvamontana.Seprecisanlos tratamientossilvícolas,entreloscualesconstituyenbosquesdeprotecciónlosqueseencuentranporencimadelos800 msnmylosmanglares. Palabras clave: Bosques;Tipologíaforestal;SierraMaestra;CubaOriental.

Introduction 2012).ThisshowsthesignificancetheCuban stategivestoforests. Forestsprovidemanygoodsandservicesto TheCubanforestareahasincreasedsignifi - humanity,mainlyrelatedtotheconservation cantly:froma13.4%ofdegradedforestsin ofbiodiversity,soilandwater.Inthecurrent 1959(RussóMilhet,2015),toa26.2%in2009 situationofclimatechangetheyalsoworkas (Alvarez& al .,2012)and28.6%in2014 carbonsinks,soitisessentialtheirconserva - (CITMA,2014;Parada&Torranzo,2014)in tionandsustainableuse.ThenumberofCuban relationtothesurfaceoftheCubanarchipelago. forestformationsdependsontheclassification Whilestudyingnature,itisimportantto recognizethesetofecologicalconditions used:Bisse(1988)identified16forestforma - (oecotope)thatareexpressedinthecharacte- tionslocatedthroughoutthecountry;while risticsofthevegetationanditsfloristiccom - Reyes(2011-2012)found23,forEasternCuba position.Foresttypologyisofgreatsignifi - (withsixsub types)and11,forSierrra cancewhentheresultsofsuchinterrelationship Maestra,withfoursubtypes(Reyes,2006). constituteinaparticulartypeofforest. TheforestrylawinCubacategorizedCuban Therefore,itisconsideredasforesttypology forestsasproducers,forconservationand thestudyofforestcommunitiesinitsrelation protective.The46.1%oftheCubanforest to environmentalconditions;sincethey heritageareprotectiv eareas,whilethe32.3% establish thequalityandproductivepotential arefortimberproduction(Alvarez& al ., oftheforeststations(sites).Alvarez&Varona

1 EasternCenterofEcosystemsandBiodiversity(BIOECO),MinistryofScience,TechnologyandEnvironment(CITMA),José A.SacoNr.601,esq.Barnada,SantiagodeCuba,Cuba.Email[email protected]

Lazaroa 37 2016: 43-103 43 44 Reyes O.J. Lazaroa 37 2016: 43-103

(1988)consideredasaforestsitetoalledaphic, ecosystemvalues,whichcontributetoforest clim aticandbioticfactorsthatdeterminethe typingandplanning. persistenceandintensityofbiomassproduc tion (grossprimaryproductivity)ofcertainfo rest community,beitnaturalorcreatedbyman. Materials and Methods Therearetwobasicmethodsforstudying forestsitesdependingontheenvironmental Natural conditions of the study area situation:withemphasisontheoecotope (Kopp,1965;Thomasius,1965;Schwanecke, SierraMaestra(Figure1)isaverycomplex 1970)orprioritizingvegetation(DelRisco, area,geologicallyspeaking.There,the 2000a,b;DelRisco&Samek,1984).The Palaeocene-Eocenevolcano-sedimentaryrocks methodthatprioritizestheoecotopeisrecom - oftheCobreGrouppredominate,mainly mendedinareaswherethevegetationhasbeen andesitesandtuffs(Mendez et al .,1994). destroyed,whiletheotherisappliedinsta - Relativelylargegranitesintrusivesoccurin tionswheretheexistingvegetationexpresses isolatedareasofthesouthernslope(around thepotentialofthatoecotope.Inthiswork,the Potrerilloriver,PinardelasCanas,LaFrancia, secondmethodisappliedbecauseitiscon - NimaNima,Juraguá-ElOlimpo,etc). ductedindevelopedvegetation,which, Likewise,theformationsMayaRiverand althoughitisoftensecondary,alsoexpresses Jaimanitasprevailinthecoastalareas(Com. theenvironmentalconditions,including CubanoHúngara,1976).

Figure1. MainforeststypesinSierraMaestrarange(Cuba)

From1500masl,theleachedyellowish acidstohighlyac idic(pHH 2O4.0-5.5)and ferralliticsoilpredominates.Accordingto theSvalueissmallerthan7cmol(+).kg -1 Renda(1989),thepHintheupperhorizonsis (Renda,1989;Renda& al .,1981b). fromacidtoveryacidic;itreachesvaluesof Sialliticbrownandfersialliticreddish- 4.45to5.3inwaterand3.5to4.15inClK. brownsoilsdominatethelowandpre-mountain Thesumofbasic cations(ChangeBases areas.Thesialliticbrownsoils(withoutcarbo- Capacity,CCB,Svalue)isverylow,generally nate)haveavariabledepth:theyusuallyhave rangingbetween1. 44and 5.02cmol(+).kg -1 . anSvaluerangingfrom13to18cmol(+).kg -1 Thecationexchangecapacity(CIC,Tvalue) onthesouthside,andfrom24to50cmol variesfrom2.5to18.75cmol(+).kg -1 .There (+).kg -1 (rarely)inthenorth,wheregen erally areverylowcontentofcalcium,magnesium havehighersaturation.Thesialliticbrown andpotassium.Beneaththissoil,toabout600 soils(withcarbonate)areabundantinthe ma.s.l,thesoilsaremainlyleachedredferra- northside,andtheyarericherandmoresatu - llitic,fromdeeptoverydeep,andoften ratedthantheabove.CCBisusuallybetween derivedfrom ferralliticweatheringcrustin 47and78cmol(+).kg -1 andtheCICranges moreorlessstableplaces.Theyarefrom from49to80cmol(+).kg -1 (Renda& al ., Reyes O.J. Lazaroa 37 2016: 43-103 45

1980,1981a,1981b,1982).Thefersiallitic Thecoastalandpre-coastalstrip,inthe reddish-brownsoilsoftenerodedandusually southofSierraMaestra,isoneofthemost withrockyoutcropsandstonesintheprofile, climaticallyextremeareasofCuba.Inthe haveanapproximatedSvalueof13cmol eastofSantiagodeCuba’sBay,theweather (+).kg -1 onthesouthernslope(Renda& al ., isbixeric.Theaveragerainfallhardlygoes 1980-81).Inthenorth,the yaresomewhatri- beyondthe700mmayear.Exceedingthe cher,whereintheCCBrangesfrom21to43 100mm,therainiestmonthsareMay, cmol(+).kg -1 .Theyaremainlyobservedin SeptemberandOctober;whilethedriest rocksfromtheCobreGroup. monthsareDecember,MarchandJuly,in Overthegranitoids,thereareusually whichrarelyexceed30mm.Therainydays gr ayishbrownsoils,sometimessandyand areusuallylessthan40peryear.Tothewest poor.TheirpHgoesfromslightlyacidicto ofthebay,rainincreasesgraduallytoabout acidic,withanSvalueof4to5cmol(+).kg -1 1200mminfrontofPicoTurquino,where inthefirsthorizon,andevenlowerbelowit. mountainsplungestraightintothesea.From Incoastallimestoneterraces,thevery thispoint,rainthengraduallydecreasesup shallowredsoils(rendzine)occupythefrac - to800mmnearCaboCruz. turesandcrevicesofthedogtoothrock.Ared TheairmeantemperatureinGranPiedra ferrallitics oilisobservedincoastalterraces, (1100masl)is18.4ºC;theaverageminimum limestonehillsandcumulativeplainsfrom is15.7ºCandtheaveragemaximum,22.4ºC. SierraMaestra’swesternmost.It’sCCBinthe OntopofPicoRealdelTurquino,theairmean firsthorizonis19cmol(+).kg -1 andtheT temperaturereachesaboutde13ºC valueis24.3cmol(+).kg -1 .Bothvalues (Montenegro,1991a).Onthenorthside, decreaseinthelowerhorizons(Renda& al ., aroundthe500and700masl,themeanannual 1981c). temperatureis20-22ºCintheupperparts InthehigherareasofSierraMaestra,it and22-24ºCinthelowest(Lapinel,1989). rainsbetween1600and2000mm,withaless Bycontrast,themeanannualtemperaturein rainyseasonfromNovembertoApril(Trusov thecoastalzonerangesfrom24-26ºC,with & al .,1983).Itisnoteworthythatcloudcon - anaveragemaximumof30-32ºC,andan densationisproducedbetween800and1200 averageminimumof22-24ºC(Montenegro, masl(horizontalprecipitation,moistshade, 1991b).Thedecreaseintemperatureper100 lowclouds).Hence,whenitoccurs,a100%of mofelevationinwesternSierraMaestrais relativehumidi tyisreached;sometimesit 0.62ºC,and0.66ºCinthenorthernand runsbythestem( BoytelYambú,1972).On southernsidesrespectively(Montenegro, average,inGranPiedra(weatherstationat 1991b). 1100masl)thereare239daysayearaffected AtthetopofSierradelaGranPiedra, byhorizontalprecipitation,ofwhich163days relativehumidityreacheshighlevels havedensefog,fromoctobertomayoccur20 throughouttheyear,fromjustover87%to daysormorepermonth(Montenegro,1990). about92%amongthedifferentmonths.At InthehigherpartsofwesternSierra 13:00h,ithasaminimumbetween80and Maestra,above1500masl,theinfluenceof 86%,andamaximum,at07:00h,from88to suchhorizontalprecipitationsisalmostdaily. 92%(Montenegro,1990). Themeanannual Airsaturation,however,isalmostlittle,even relativehumidityinCarsodeBaireisfrom withdensefog,whichisformedlowerdown 80to85%(Montenegro,1991c)andin andgoesupwiththewind.Theseconditions Contramaestre(bottom,northside),isof52 produceahightensiononthefoliage,which to60%.Inthecoastalarea,themeanannual mayturnitintomicrophyllandthedecompo - relativehumidityvariesfrom75and80%, sitionfromwitheredisveryslow themeanannualat13.00hisfrom70to (Reyes&Fornaris,2011). 75%,andat07:00h,itisfrom80to85% InCarsodeBaire(northside),around500to (Lapinel,1989;Montenegro,1991c).In 700masl,themeanannualrainfallvariesfrom SantiagodeCuba(lowerpart,southern around1200mminthelesselevatedareasupto slope)itvariesbetween47and60%.At morethan1600mminthehighest(Gagua& 13:00h,suchhumidityincreaseswithalti - al .,1989).TheaveragerainfallinLaTabla(600 tudefromthebottomupto1400masl, masl)is1634mm,varyingfrom1216to2320 remainingconstanttoabout1600masl. mmthroughtheyears:onaverage,thereare94 ThenitdecreasestothetopofPicoRealdel rainydaysayear.ArainyperiodfromMarchto Turquino(Montenegro,1991c). November,with85%ofrainfallisclearly EvaporationinGranPiedraislowandfor defined,aswellasotherlessrainyfromdecem - eightmonthsintheyear,itdoesnotexceed bertofebruarywith15%.Theheaviestrainfall 100mmpermonth(Montenegro,1990). rangesbetween5.1and30mm. Themonthswiththehighestevaporation 46 Reyes O.J. Lazaroa 37 2016: 43-103

(Piche)areapril,june,julyandaugust(from Herbaceous:under1m.Fernswere 75to82mm),andwiththesmallerone,from consideredseparately;therearedifferenttypes octobertojanuary(lessthan60mm) ofterrestrialferns.Insomeforest,theyare (Montenegro(1991d).Inturn,inthecoastal foundinthethreelayersdescribedbefore, zone,evapora tionfluctuatesbetween1900 mainlyintheherbaceouslayer. and2000mm(Crespo,1989).FromMarch Lianas:Itinvolvesthedifferenttypesof toSeptember,itisover110mm(Piche), vinesorlianas. March,April,JulyandAugustarethe Epiphytes:Plantslivingoverthephoro - monthswiththegreaterevaporation phyteswithoutbeingparasites.Sometimes (130-1400mm),andNovemberand theyplayanimportantroleasindicatorsin Decembershowthesmallestevaporation differentforesttypes,especiallyofenviron - records,with85mm(Montenegro,1991d). mentalhumidity. InCarsodeBaire,themeanannualevapora - Vertical structure of canopy layer (C/V). tionisof1300to1500mm(Montenegro, Componentsofthislayerwereindividually 1991d) . calculated,directlyontheplots.Kraftclassifi - cation(inSamek,1974)wasused: Predominant(P):Thebeyondthepri - Methodology marylevel. Dominant(D):Thosemakingupthemain Thepresentpaperusedsomedataobtained level. fromphytosociologicalstudies(205phytoso - Codominant(C):Thoseinthelowerpartof ciologicalplotsfrom625m 2)conductedinthe theprimarylevel,butwhicharenotwell forestsofSierraMaestra;someofthem developed. alreadypublished(Reyes&Acosta,2003, Intermediate(I):Thosebelowthemain 2004;Reyes& al .,2004a,b;Reyes& level,whichoftenformasub-layer. Martínez,2005)andsomeothersstillunpub - Oppressed(O):Theyarepartofthebottom lished.Thesesyntaxaaretransformedintothis sub-layer.Theyhavelittlevitality. foresttypologyusingthestandardsofthe Abundance-Coverage (A-C). Theprojec - InstituteofAgro-ForestryResearch(Del tionofthecrownorthenumberofindividuals Risco,2000a,b).Thescrubsyntaxa,grass - (onlyinthefirsttwoindices)wascalculated, lands,etc.werenottakenintoaccountinthis accordingtothemodifiedscaleofBraun study. Blanquet(1979).Inthisstudy,thescalefromone Twomaintypesofindicatorswereconsi- tofivewasused.Eachnumberrepresentsarange dered: fromtheA-Casdiscussedbelow.Iftherearetwo numbers,itmeansthattherangevariesbetween • Forestscharacteristics. thesetwovalues.Thescaleisasfollows: • Oecotopecharacteristics. 1.Shortlyrichorpoorcoverage.Itcovers upto5%ofthesampleplot. 2.Littleortoonumerous,orcoverage Forest Characteristics between6and25%. 3.Coveragebetween26and50%. DelRisco(2000a,b)methodologyfor 4.Coveragebetween51and75%. strata,verticalstructureofcanopylayer, 5.Seventysixpercent(76%)orgreater abundance-coverage,presencedegreeand coverage. eachspecies’sparticipationwasappliedfor Presence Degree (P-D). Theexistenceofa studyingtheforestcharacteristics.Outofit, speciesinthedifferentsampledplotsiscon - thecharacteristiccombinationofeachforest ceivedaspresence.Valuesmeanaccordingto typewascreated.Incaseofanalogous thenumberoftimessuchspeciesarefoundin forests,havingasimilarfeaturecombination, thesampledplotsandtheyareexpressedinthe thedifferentialcombinationwasdetermined. followingscale: Inaddition,thecanopylayerorsub-layer V.Constantlypresent:whenspeciesare coveragewasconsidered,aswellasthe observedinover80%oftheplots. andherbaceouslayers. IV.Presentmostofthetimes:whenspecies Strata (layers): Samek(1973)criteriafor arefoundfrom61to80%. separatingthethreemainstrata(canopy, III.Oftenpresent:whenspeciesarefound shrub,herbaceousandferns)andtheother between41and60%. synusiaeswereusedaccordingto: II.Rarelypresent:whenspeciesarepresent Canopy:Treesgreaterthan5minheight. from21to40%. Shrub:andlittletreesbetween1 I.Rare:whenspeciesareobservedinless and5m. than20%oftheplots. Reyes O.J. Lazaroa 37 2016: 43-103 47

Type of participation. Itisthecombination relief,exposure,slopeindegrees,position oftheindicesofabundance-coverageandthe ontheslope(relativeelevation)and presencedegree.Itisconsideredasfollows: microrelief. Mainspecies:Theyarespecieshavinga Climatope characteristics: SierraMaestra presencedegreeofVandanabundance-co- climatopewasdeterminedusingtheBorhidi verageoftwoorgreater. (1996)bioclimate,withsomemodifications Subdominantspecies:Specieswithapresenc e fromSamek&Travieso(1968),Montenegro degreeofVandanabundance-coveragelower (1990,1991a,b,c,d,e)andTrusov& al . thantwoorspecieswithapresencedegreeof (1983)intheparameters:temperature,mean IVandanyabundance-coverage. multi-annualrainfallanddrymonths. AggregatedSpecies:Specieswithapresence Edatope characteristics: Intheplotand degreeofIII,andanyabundance-coverage. itssurroundings,thereareobserved:rockand ScatteredSpecies:Specieswithapresence soiltypes,depthofsolum,texture,structure, degreeofIIorIandanyabundance-coverage. stoniness,gravelnessanderosion.Besides,the Canopy layer coverage :Itisthefloorsur - organiclayers(L,FandH)andsoilprofileare faceoccupiedbythehorizontalprojectionofa described. foliage.Itiscalculatedaccordingtothe Thetrofotope(trophism)isstudied:A= classificationofSamek(1974). littlefertile,B=moderatelyfertile,C=fertile Superdense:Whencoverageisgreaterthan andD=veryfertile.Thehigrotope(soilmois - 100%.Thebranchesofsomeindividualsenter ture)isalsostudied,whichaccordingtothe inthespaceofthecrownsoftheneighboring Pogrebnyakmethod(1955,inSamek,1973)is individuals. classifiedas:0=verydry,1=dry,2=fresh,3 Dense:Coveragebetween91and100%; =moderatelywetand4=wet. branchestoucheachother. Thepercentageofstoneonthesurfaceand Slightlysparse:Coveragebetween81and thepresenceofdogtooth(lapiez)areclassified 90%. asfollows:lessthan40%( x),40to59%( xx ), Sparse:Coveragebetween51and80%. 60%ormore( xxx ).Therootmathasavitalrole Verysparse:Coveragebetween21and50%. innutrientcycleinsomeforeststypes,thereby Acoveragelowerthan/belowthe20% itisimportanttotakeintoaccountwriting correspondstoa“clearing”withscatteredtrees. (Rm*)whentheyhaveupto10cmand Coverage of the shrub and herbaceous (Rm**)formorethan10cm. layers. Thepercentageofeachoftheselayers Name of the forest type :Tonamethe iscalculatedasfollows: foresttype,itisusedthecommonname Veryhigh:Coveragebetween91and (Appendix1)oftwotypicalspeciesfromthe 100%. characteristiccombination(constant)ofthat High:Coverageof51to90%. type.Thefirstonemustbelongtothecanopy Media:Coveragebetween21and50%. layer(constantanddominant)followedby Low:Coveragebelowthe20%. anothertypicalandconstantspeciesfromany Characteristic combination: Itistheset stratum.Todifferentiatesomeothertypesof ofspeciesthatcharacterizeandtypifythe forest forestsmoreorlesssimilar,thenameofa type.Itisformedbymain,subdominantand typicalandconstantspeciesfromthedifferen - aggregatedspecies.Whenthereisagroupof tialcombinationisused,andthenitisaddedto speciesthatdifferentiatessimilarforesttypes thepreviousname. inoecotopeswithsomedissimilarity,itforms Throughthecharacteristicsoftheoeco - adifferentialco mbination.Inthesecombina - topesoccupyingthedifferentforesttypes,itis tions,constantsp ecies(thosehavingaV possibletodeterminetherelativequalitiesof presencedegree),andaccompanyingspecies theforeststationorsites,whichdependsonthe (thosehavingIVandIIIpresencedegree) potentialproductivecapacity (PPC),remaining weredifferentiated. definedfiveclasses:excellent(I);good(II); Oecotopes characteristics: Theoecotope regular(III);bad(IV)andverybad(V). istheclimatope(climaticconditions)and Forforestmanagementproposals,itis edatope(soilconditions:higrotopeandtrofo - takenintoaccount,inadditiontotheabove, tope)assembly.Sitesthatoccupytiltedareas thefloristic,syngeneticandphysiognomic areinfluencedbyphysiographiccharacte- characteristicsofforests(microphylly,meso - ristics.Theywillbethereforetakeninto phylly,size,degreeofcoverage,etc.);aswell account. asthedevelopmentstrategies(nutrientscircu - Physiographic characteristics: These lationtype),stagesofdegradationandstrate - observationsaremadeineachplotalongsideits giesforrehabilitationand/orrestoration. surroundingarea.Thefollowingparametersare Scientificnames(,speciesand determined:elevationabovesealevel,typeof author)arefullywritteninAppendix1. 48 Reyes O.J. Lazaroa 37 2016: 43-103

Results and Discussion side,meanannualtemperatures ran ge between21.3-23.1ºC(topandbottom respec - Climatic type tively),and21.0-24.0ºConthesouthern slope(Montenegro,1991b).The rainfall Tropicalclimatewithtwodryperiods:Type rangesfrom1200to1800mm(medium rainy ofbioclimatewithlittleextensionin torainy),accordingtothelocality.Relative SierraMaestra,duetotherainshadowofthe humidityfluctuatesduringtheyearbetween mountains.Thefollowingsubtypeis 77and85%at500masland84to92%atthe observed: top(Montenegro,1991c).InCarsodeBaire, 5aTh.Extremelydrytropicalbixericcli - meanannualevaporationis1300to1500mm mate:Itisobservedinthecoastalstrip(zone (Montenegro,1991d).Inmountainousareas ofwetlandsandsemi-deciduousmicrophyll below800ma bovesealevel,intheevenings forest)eastofthebayofSantiagodeCuba.It andinmid-apriltolateOctober,relative isoneofthemostclimaticallyextremeareas sun shine(percentageofrealsunshineregard - inCuba;itrainsalittlelessthan800mma ingtheastronomicalone)variesbetween20 yearonaverage(aridclimate);therainy and30%,therestofthemonthsisfrom30to monthsareMay,SeptemberandOctober 40%.Duringthemornings,itrangesfrom40 exceeding100mm;onthecontrary,thedriest to60%(Montenegro,1991e). onesarefromdecembertoMarchandJuly,in Wetmountainoustropicalclimate:This whichhardlyevertherainfallgoesover30 kindofclimateoccursinhigherpartsofSierra mm(extremelydry).Therainydaysareusu - Maestra,from800maslandthetopofPico allylessthan40peryear.Evaporation Turquino(maximumelevationoftheCuban rangesfrom1900to2000mm(Crespo, archipelago,1972masl).Twosubtypesare 1989).Theincreasedevaporationmonthsare observed: March,April,JulyandAugustwith130to 7a-Wetmountainoustropicalclimate:It 140mm(Piche)andthosewithminorevap - occupiesthe“floorofthemountain rainfor - orationareNovemberandDecemberwith85 est”,i.e.between800and1500masl.Annual mm(Montenegro,1991d).Meanannualtem - rainfallis1600-2000mm,withalowerrainy peraturerangesfrom24-26ºC,withamean periodfromNovembertoApril(Regal,1988) annualmaximumof30-32ºCandamean andwithagreatinfluenceofthehorizontal annualminimumof22-24ºC(Montenegro, precipitation:239daysinGranPiedra(1100 1991b). masl;Montenegro,1990).Itisnoteworthy Tropicalclimatewithdrywinter:Thistype thatinthisareatheorographiccloudconden - ofbioclimateisthemostwidespreadinSierra sationoccurs,whichtakesplacebetween800 Maestra,withtheexceptionoftheupperparts and1200masl;therefore,a100%relative andthecoastalareainthesoutheastopposite humidityisobservedwhenitoccurs,some - side.Thetwofollowingsubtypesarefound: timesgettingtoflowthroughthestems 4bTh-Tropicalweatherwith5to6scarce - (BoytelYambú,1972).Duetothesehorizontal lyrainymonths(medium-dry):Itisobserved rainfallinabove800maslareas(evenings), onthe“floorofsemideciduousforestsand for9months,itdoesnotexceedthe20%of scrubs”,longerspreadtothewestofthebayof relativesunshine,includingfourmonthsin SantiagodeCuba.Thealtituderangesfrom whichitislessthan10%.Theothermonthsit sealevelto400-500maslinthenorthernand rangesbetween20and30%.Duringthemorn - southernsidesrespectively;also,themean ings,itvariesfrom30to50%.Throughoutthe annualtemperaturefluctuatesbetween24.5 year,suchrelativehumidityreacheshighval - and26.0ºCandrainfallvariesfrom800to ues,fromover87%tonearby92%amongdif - 1200mm(fromlowtomediumrainy). ferentmonths.Themeanairtemperaturein Meanannualrelativehumidityis75to80% GranPiedrais18.4ºC.Therefore,evaporation andevaporationis1700to1900mm islowandforeightmonthsintheyearitdoes (Crespo,1989;Montenegro,1991d).Tothe notexceed100mm/month(Montenegro, westofthebay,rainincreasesgraduallyup 1991d).Thezonalvegetationismountain toabout1200mm(mediumrainy)before rainforest. PicoTurquino(wherethemountainsplunge 7b-Tropicalclimateofhighmountain: straightintothesea),fromthispoint,it Thisclimatictypetakesplaceonthe“floor graduallydecreasesto800mmnearCabo ofcloudforests”from1500mabovesea Cruz(littlerainy). leveltothetopofPicoRealdelTurquino 4cTh-Tropicalclimatewith3to4little (1972masl).Themeantemperaturefluctuates rainymonths(moderatelydry):Itisobserved at1500 masl,between14.1ºCand19.3ºC in onthe“floorofevergreenforests”,i.e., januaryandjulyrespectively;inturn,atthe between500and800masl.Onthenorthern topofTurquino,itreachesaboutde13ºC Reyes O.J. Lazaroa 37 2016: 43-103 49

(Montenegro,1991b).Rainfallsreachabout conditions(higheraltitudesoftheCuban 2000mm(rainyweather).Theinfluenceof archipelago,sublatitudinalextension),Sierra horizontalprecipitations(orographicclouds Maestrahasagreatdiversityofforesttypes. andmists)isalmostdaily;however,onits Themainonesare: outside,theforestexperiencessignificant waterstressduetothescarceairsaturation (Reyes&Fornaris,2011).Comparedtothe “floorofthemountainrainforest”,relative Uva caleta with yana ( Coccoloba uvifera humiditydecreasesinthesealtitudes,avera- with Conocarpus erectus ; Table 1) gingbetween81and85%at1500masl,and 82-83%ontopofTurquinoindryandrainy Physiognomyandfloristiccomposition: periodsrespectively(Montenegro,1991c). Thistypeofforestisknownassea-grape Sometimes,valuesof36and40%are woodlandandconsistsonalongandnarrow reachedduringthehottesthours(Reyes& stripdominatedby Coccoloba uvifera al. ,2011). (Table1).Generally,itsheightdoesnot exceed8m,exceptinginbeaches,whereit Forest Typology canreachabout15m.Duetotheeffectofthe seawinds;itoftenformsaninclinedplane, Duetothewidevarietyofoecotopes(typesof fromnearthegroundtotheappropriateheight weather,rocksandsoils)andphysiographic level.

Table1. Characteristiccombinationofuvacaletawithyana (Coccoloba uvifera with Conocarpus erectus ,6plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Coccolobauvifera D4 V Conocarpuserectus D2 V

Scatteredtreespecies:majaguadelaflorida( Thespesia populnea ),ipilipil( Leucaena leucocephala ),vomitel( Cordia sebestena )andguacaloteamarillo( Caesalpinia bonduc ).

Oecotopecharacteristics.Duetoitslocation inthefarthestareasfromtheseaandthe nearthesea,itisremarkablyinfluencedbysalt estuaries. ladenseabreezes(saltspray)andbythe spl ashesduringheavyswell,whichchangesthe localclimate.Itisgenerallyobservedoverthe Red mangrove community ( Rhizophora CobreGroupandsandybeaches;soilsaresandy mangle ; Table 2) andgravelly,usuallyveryshallow. Termsforforestmanagement.Sincethey Physiognomyandfloristiccomposition. protectsurroundingvegetationfrommarine Thiscommunityispresentedasabeltbetween saltsprayandtheyareanarrowstrip,they 2and10mwidewherethereisaverystraight - mustbeprotectedagainstanyhumaninterven - forwardcontactwiththesea.Itmayreach tion,sincetheycanbeeasilydestroyed. highervaluesinflatareas.Itsheightisusual - lybetween5and10manditsstructureis rathercontinuous,withoutdefinedlayer. Mangrove stands Regenerationisvariable,sometimesscarce. Sinceitisexclusivelycomposedbyredman - Mangrovesareaffectedbymarineand grove( Rhizophora mangle ),thisforestforms freshwaterinfluence,whichdeterminesthe adistinctecosystem.Itsanchoringaerialroots correspondingtype.Therefore,thelocalcli - createspecialconditionsforestablishingand maticconditionshavetheirgreatestimpact developingvarioustypesoffauna(Table2). 50 Reyes O.J. Lazaroa 37 2016: 43-103

Oecotopecharacteristics.Itoccursinlow- Termsforforestmanagement.Itmustbe lyingplaceswithsedimentdeposition,usually protectedfromanyhumanintervention,since inareasprotectedfromoceanstreams.Itis itformsanarrowstripandprotectsthecoasts directlyincontactwiththeoceanwaves,and andthecommunitiesbehindthem.Thereare theplace swheretidevariationgetsitshighest importantareasthathavebeendestroyedand values.Thesubstrateconsistsofquaternary shouldbetherefore,restored.Recoverycanbe sedimentswhosecompositionvariesdepending madebymeansofthepropagules,whichare onthe surroundinggeology. borninthetree(viviparouscapacity). Distribution.IttakesplaceinalltheCuban archipelagocoasts.

Table2. Characteristiccombinationoftheredmangrovecommunity (Rhizophora mangle ,10plots) Species Vertical Abundance- PresenceDegree structure Coverage Alllayers Rhizophoramangle D5 V

Community of red mangrove with black whileintheshrublayeritisalmostalways mangrove ( Rhizophora mangle with low (5and10%);seedlings,ontheotherhand, Avicennia germinans ; Table 3) showanaveragecoveragefrom20to50%. Oecotopecharacteristics.Theyaresimilar Physiognomyandfloristiccomposition.In tothoseoftheredmangrovecommunity, SanMigueldeParadas,bayofSantiagode althoughwithlessinfluenceofwaves. Cuba,thismixedmangrovestandformsanar - Distribution.Itcomesaftertheredman - rowbeltofupto70m;beinglowerinsome grovecommunity,orindirectcontactwiththe otherplaces.Suchstripisonlycomposedby sea;therefore,itreceivesalongsidetheformer redandblackmangroves( Avicennia germinans ) community,thestrongesttidalinfluenceand altogether(Table3).Usually,thefirstoneis thegreatestdepthofwater.Atcertaintimes, moreabundantbeforethesea.Thisiswhere suchawaterheightisofatleast20to30cmat itformsthebulkofthecanopylayerinthe lowtide. first 7to15m;itappearslaterasseedlingsin Termsforforestmanagement.Thiskindof aforestdominatedbythesecondtypeofman - mangrovestandshouldbeprotectedfrom grove.Thecanopylayerisbetween10and 12 humanactivitiesduetoitscoastalprotection mtall,withadensecoverage(around90%), functionsandnarrowstripcondition.

Table3. Communityofredmangrovewithblackmangrove (Rhizophora mangle with Avicennia germinans ,10plots) Species Vertical Abundance- PresenceDegree structure Coverage Alllayers Rhizophoramangle D3 V Avicenniagerminans D2 V

Scatteredtreespecies:whitemangrove( Laguncularia racemosa )

Community of black mangrove ( Avicennia islocatedbehindtheredmangrove,and/or germinans ; Table 4) theredwithblackmangrovecommunities.It consistsofpurepopulationsof Avicennia Physiognomyandflor isticcomposition. germinans ,whichgivesitadistinctivelook.It Thistypeofmangrovestand isfoundthroughout showsitsgreatestheightinthepartincontact the studyareaandisgenerallypredominant.It withthecommunityofredmangrove,whereit Reyes O.J. Lazaroa 37 2016: 43-103 51 rea chesbetween10and12m.Behindthis ments.Itislocatedintheareaoftidalfluctuation point,itonlyreachesfrom3to6mandcan anditiscompletelyaffectedduringhightide. beconsideredasstuntedmangroves.Itis Distribution.Theycanbefoundinall considerablethehighdensityofpneu - coastsoftheCubanarchipelago. matophores,whicharesometimesusedas Termsforforestmanagement.Itmustbe attachmentpointsofseaweedandanimals, protectedlikeallmangrovecommunities. andoverwhichitispossibletowalkeasil y Therearesignificantareasaffectedbyhuman (Table4). impact.Suchareasshouldberestoredinorder Oecotopecharacteristics.Thesoilsare tore-establishtheirroleasprotectorofthe boggy(halomorphic)withquaternarysedi - coastsandthebiodiversity.

Table4. Characteristiccombinationoftheblackmangrovecommunity(10plots)

Species Vertical Abundance- PresenceDegree structure Coverage Alllayers Avicenniagerminans D5 V

Black mangrove with acrostic ( Avicennia sitydeterminestheabundanceofherbs. germinans with Acrostichum danaefolium ; Distribution.Itisdispersed.Itwasstudied Table 5) atthenorthwesternedgeofthemangrove standsofSanMigueldeParadas,whereinter - Physiognomyandfloristiccomposition. mittentstreamsflowwhichcausesignificant Thistypeofmangrovestand(Table5)hasa freshwaterinput.Furthermore,itislittle blackmangrovecanopylayerof12to15m influen cedbythetides,duetoitsgeographical high,withacoverfrom verysparsetosparce positioninthemangrovestand. (40to70%).Theshrublayerisofacrostic Termsforforestmanagement.Likeother (Acrostichum danaefolium ),whichreaches mangrovecommunities,itshouldbeprotected about5to6mandhasacoverbetweenhigh fromallhumanactivity. andveryhigh(51to100%). Oecotopecharacteristics.Itusuallyde- velopswithfreshwaterinfluence,whoseinten -

Table5. Blackmangrovewithacrostic (Avicennia germinans with Acrostichum danaefolium ,5plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Avicenniagerminans D3 V Shrublayer Acrostichumdanaefolium -4 V

Black mangrove community with white and10minheightanditoftenhasunderneath, mangrove ( Avicennia germinans with seedlingsofbothspecies(Table6). Laguncularia racemosa ; Table 6) Oecotopecharacteristics.Thesoilsare wea klyflooded.Insomeplaces,theyarejust Physiognomyandfloristiccomposition. directlyaffectedbysaltwaterduringhightides. Thistypeofforest,composedbywhiteman - Theyarealsoinfluencedbytherainwater. grove( Laguncularia racemosa )andblack Distribution.Itiscommonlyseenthroughout mangroveisverypoorlyrepresentedanditis thestudyarea. knownasmixedmangrove.Sometimesitis Termsforforestmanagement.Likeother putapartfromtheredmangrovebysalt mangrovecommunities,itshouldbeprotected marshesofvariablesize.Itreachesbetween4 fromallhumanactivity. 52 Reyes O.J. Lazaroa 37 2016: 43-103

Table6. Blackmangrovecommunitywithwhitemangrove (Avicennia germinans with Laguncularia racemosa ,7plots)

Species Vertical Abundance- PresenceDegree structure Coverage Alllayers Avicenniagerminans D3 V Lagunculariaracemosa C1 V Scatteredtreespecies:yana,cambrón( Prosopis juliflora )

Black mangrove with white mangrove and has avari ablecover,media(21-50%);while junco ( Avicennia germinans with intheherbaceouslayerishigh(51-80%).This Laguncularia racemosa and Eleocharis typeofmangrovestandistherichestinherba - mutata ; Table 7) ceousspecies(Table7). Oecotopecharacteristics.Waterusually Physiognomyandfloristiccomposition.In variesbetween2and10cmabovethesub - SanMigueldeParadas,thisforestisamong strate,stronglyinfluencedbyfreshwater. themangrovestandsofblackmangroveand blackmangrovewithacrostic,i.e.,itcomes Distribution.Itisinthenorthwesternpart afterthelatterintothemangrovestands .In ofthemangrovestandsofSanMiguelde additiontotheblackmangrove,thereis ParadasinthebayofSantiagodeCuba. oftenwhitemangroveinthecanopylayer.It Termsforforestmanagement.Likeother is8to14mhighanditscoversfrom very mangrovecommunities,itshouldbeprotected sparsetosparse(21-70%).Theshrubla yer fromallhumanactivity.

Table7. Blackmangrovecommunitywithwhitemangroveandjunco (Avicennia germinans with Laguncularia racemosa and Eleocharis mutata ,7plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Avicenniagerminans D3 V Lagunculariaracemosa C1 III Herbaceouslayer Eleocharismutata -2 V Typhadomingensis -2 V Eleocharis.elegans -1 V Cyperusgiganteus -1 V Scirpusolneyi -1 V Cyperusalternifolius -1 III Ludwigiaerecta -1 III Bacopamonnieri var. monnieri -1 III Alternantheramaritima -1 III Lianas Mikaniamicrantha -1 III

Scatteredtreespecies:yana. Reyes O.J. Lazaroa 37 2016: 43-103 53

Yana community ( Conocarpus erectus ; Oecotopecharacteristics.Usuallythe Table 8) groundismuddyandbrown.Itoftenoccupies lagoon-likechannelsandisusuallyoutoftidal Physiognomyandfloristiccomposition. influence. Thistypeofmangrovestand(Table8)is Distribution.Itisahighlydisperseman - exclusivelycomposedbyyana( Conocarpus grovestand. erectus ).Stratificationiscontinuousanddense Termsforforestmanagement.Likeother (100%)andcanreach8to12mhigh. mangrovecommunities,itshouldbeprotected fromallhumanactivity.

Table8. Yanacommunity( Conocarpus erectus ),6plots

Species Vertical Abundance- PresenceDegree structure Coverage Alllayers Conocarpuserectus D5 V

Jatia with no me toques ( Phyllostylon species.Amonglianas,bejucodepurgación brasiliensis with Oplonia tetrasticha ; Table 9) (Commicarpus scandens ) issubdominant; and asaggregatedspecies , therearebejucoan- Physiognomyandfloristiccomposition. garilla (Serjania diversifolia ), bejucodeberra - Thisforesttypeispartofthesemi-deciduous co( Chiococca alba ), amansaguapo microphyllforest.Thecanopylayeris (Triopteris rigida ), Cynanchum sp.andbejuco between8and10metershigh,unusualif uví( Cissus trifoliata ).Curujey( Tillandsia more.Itscoverageisbetween60and80%, fasciculata )andbarbadeindio( T. usneoides ) andconsideredasscarce.Withinthegroupof aresubdominantintheepiphytesynusiae. constantspecies,onlyjatia[ Phyllostylon Oecotopecharacteristics.Thealtitude brasiliensis (= Ph. rhamnoides )]isinthe variesbetween10and80maslwithanincli - categoryofmainspecies;baría( Cordia nationbetween25and45degreesandan gerascanthus ),carbonero( Colubrina elliptica ) exposuremainlysouthward.Itislocatedon andchicharrón( Pseudocarpidium avicennioides ) thebottomofthemountains,nearthesea,with aresubdominants.Intheshrublayer, with avariedmicroreliefandstronglyaffectedby acoverconsideredmediumtohigh(40-80%), thesalt-ladenseawinds(saltspray).This nometoques( Oplonia tetrasticha )and south-coastalareaofSierraMaestra,where Varronia globosa aremainspecies;while thisforesttype(“floorofsemideciduous pichadeperro( Capparis flexuosa )andpalo forestsandscrubs”)isobserved,isunderthe bronco( Malpighia suberosa )aresubdominant effectoftherainshadow(windscreen,Föhn species.Intheherbaceouslayer,cuabade effect)provokedbythesemountains;which ingenio( Croton lucidus )isthemainspecies, meanslowrainfall,verywarmtemperature, withamediaaveragecover(20-50%).As lowrelativehumidityandhighevaporations lianasandepiphytes,bejucosanpedro rate. (Stigmaphyllon sagreanum )andcurujey TotheeastofthebayofSantiagodeCuba, (Tillandsia recurvata )aresubdominant thereisanextremelydrytropicalbixericcli - species , respectively(Table9). mate(5aTh)andrainfallislessthan700mma Inthegroupofaccompanyingspeciesfrom yearonaverage.TherainiestmonthsareMay, thecanopylayer,agalladecosta( Randia SeptemberandOctober,withmorethan100 aculeata )isobservedassubdominant . mm.ThedriestmonthsarefromDecemberto Negracuba (Thouinia patentinervis ), majaguilla March,andJuly,whenbarelyexceedsthe30 (Carpodiptera cubensis ), aceituna( Picrodendron mm.Therainydaysareusuallylessthan40 baccatum )andyuraguana( Coccothrinax peryear,beingoneofthemostclimatically gundlachii )areaggregatedspecies.Intheshrub extremeareasCuba’s,hence,itisconsidereda layer, Pictetia mucronata , Tabebuia myrtifo lia hotclimatewithtwodryperiodsandmore andfrijolillo( Senna atomaria )areaggregated . thansixmonthsofdrought.Tothewestofthe Intheherbaceouslayer, Malpighia apiculata bay,rainincreasesgradually,changingto tropi - isconsideredassubdominant,whilejijira calclimatewithdrywinter,4bThsubtype, (Harrisia eriophora ), marilope (Turnera semi-dry,with5to6monthswithlittlerain. ulm ifolia ), jibá (Erythroxylum havanense ) and Precipitationfallsfrom900toabout1200mm anamú (Petiveria alliacea )areaggregated infrontofthePicoTurquino(wherethemoun - 54 Reyes O.J. Lazaroa 37 2016: 43-103 tainstumbledirectlyintothesea),fromwhere (Lapinel,1989;Montenegro,1991b,c). itgraduallydecreasesto800mmnearCabo Evaporationrangesbetween1900and2000 Cruz.Themeanannualtempera tureranges mm(Crespo,1989);fromMarchto from24to26ºC,withameanmaximumof Septemberitisover110mm(Piche),March, 30to32ºCandameanminimumof22to24 April,JulyandAugustarethemonthswith ºC(Montenegro,1991b).Themeanannual higherevaporation(130to140mm)and relativehumidityvariesbetween75and NovemberandDecembershowsthesmallest 80%,themeanannualat13.00his70to evaporationrates,85mm(Montenegro, 75%,andfrom80to85%at07:00h 1991d).

Table9. Characteristiccombinationofjatíawithnometoques (Phyllostylon brasiliensis with Oplonia tetrasticha ,7plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Phyllostylonbrasiliensis D4 V Collubrinaelliptica D1 V Pseudocarpidiumavicennioides D1 V Cordiagerascanthus C1 V Randiaaculeata I1 IV Picrodendronbaccatum C1 III Carpodipteracubensis C1 III Thouiniapatentinervis C1 III Coccothrinaxgundlachii I2 III Shrublayer Oploniatetrasticha -2 V Varroniaglobosasubsp. humilis - 2 V Capparisflexuosa - 1 to2 V Malpighiasuberosa -1 V Tabebuiamyrtifolia -1 III Pictetiamucronata -1 III Sennaatomaria -1 III Herbaceouslayer Crotonlucidus -2 V Malpighiaapiculata -1 IV Harrisiaeriophora -1 III Turneraulmifolia -1 III Erythroxylumhavanense -1 III Petiveriaalliacea -1 III Lianas Stigmaphyllonsagreanum -1 V Commicarpusscandens -1 IV Cynanchum sp. - 1 III Cissustrifoliata -1 III Serjaniadiversifolia -1 III Chiococcaalba -1 III Triopterisrigida -1 III Epiphytes Tillandsiarecurvata -1 V Tillandsiafasciculata -1 IV Tillandsiausneoides -1 IV

Scatteredspeciesinthecanopylayer: almácigo (Bursera simaruba ),guásima (Guazuma ulmifolia ),cafecillo (Bourreria virgata ),yamaquey( Belairia spinosa )andaguedita (Picramnia pentandra ). Localities:SouthernpartofSierradelTurquino. Reyes O.J. Lazaroa 37 2016: 43-103 55

Itisgeologicallylocatedonrocksfromthe darycommunityofthesemi-deciduous CobreGroup(Méndez& al .,1994)andasso - microphyllforestandisthestagefieraII ciatedtolimestoneofJaimanitasformation. undertheconditionsinwhichittakesplace. Therearemainlyrocksandstonesfromthe Thecanopylayerisverysparse(20%),occa - firstgroup,mixedwiththesecond´sones, sionallymore,andtheheightis7to8m,in althoughthereversesituationisalsoobserved, moredevelopedareasitcanreachupto10m. i.e.limestonemixedwithrocksandstonesof Asdominantspeciesthereiscarbonero,and theCobreGroup.Thesoilisreddishbrown asco-dominant(subdominantspecies)there fersiallitic(Renda,1989;Hernández& al ., aremajaguilla,almácigoandcafecillo.Inthe 1994),veryshallow,oftengravelly,containing shrublayer,coverageishigh(70to90%);fri - stonesandrocksrangingfrom60to90%. jolilloand Varronia globosa subsp . humilis Therefore,itcanbeconsideredasverydry(0) arethemainspecies,whilebayúa andlittlefertile(A). (Zanthoxylum elephantiasis )andpichade Becauseitgrowsinsuchdifficultenvi - perro,aresubdominantones.Theherbaceous ronmentalconditions(shallowsoil,high layercoverageishigh(60to70%),with rockiness,stoniness,gravelness,hightem - cuabadeingenio (Croton lucidus )asthe peratures,highevaporationandmarinesalt mainspecieswithhighabundance–cover - sprayinfluence),thisforestismuch age,andtapaculo( Helicteres semitriloba ) as stressed,whichisexpressedinitsmicro - subdominant.Amonglianas,bejucosan phyll,sclerophyllousandsemi-deciduous pedro( Stigmaphyllon sagreanum )andbejuco character(Reyes&Acosta,2004). depurgaciónaresubdominants. Therefore,thepotentialproductivecapacity Thegroupofaccompanyingspecies is shouldbeconsideredasverybad(V).The numerous.Inthecanopylayer,negracuba, typologyclassificationisasfollowsA xxx 0, chicharrón( Pseudocarpidium avicennioides ), 5aTh a -4bTh b-c ,V,Ph.b.,O.t. Pictetia mucronata andyuraguana Distribution.Thisforesttypehadamuch (Coccothrinax gundlachii )theyaresubdomi - widerdistributioninthiscoastalarea,yet nants,withhighabundance-coverage.Inthe wasstudiedinareasclosetothecoastof shrublayerjibáissubdominantandinthe SierradelTurquinoandwasobservedin herbaceouslayer,thefollowingspeciesare relativelylargeareasonbothsidesofthebay alsosubdominant:nometoques,palobronco ofGuantanamoandinscatteredpatches (Malphigia suberosa ),abrojodecosta elsewhere. (Castela spinosa ), Zapoteca gracilis , Lantana Termsforforestmanagement.Duetothe montevidensis ,marilope( Turnera ulmifolia ), ecologicalconditionsinwhichitdevelops, Crossopetalum sp.,yerbadeguinea( Urochloa thiscoastalforesthashardwoodtreespecies humidicola )andcamagueyana( Bothriochloa andveryslowgrowing,sohandlingshouldbe pertusa ).Barbadeindio( Tillandsia especiallycareful.Itmustbeprotectedinge- usneoides )andcurujey( Tillandsia recurvata ) neral,with30-year-spacedselectiveinterven - aresubdominantspecies amongtheepiphytes tions.Specialcaremustbetakenwithrege- (Table10). neration,whichisgenerallypoorands cattered. Oecotopecharacteristics.Itoccupiesthe Thereareplentyofcoastaloecotopesadequate upperareasofthemicro-basins,wherepartof torestorethistypeofforest,whichmustbe therainwaterdoesnotinfiltratebutdrainsinto plantedwithjatía,asamainspecies,mixed thelowerareas,hence,thehydraulicstressis withbaría,almácigo,carbonero,majaguilla verypronounced.Ittakesplacebetween70 andnegracuba.Climatechangemodelspredict and120maslTheslopeisverysteep,40to45 increaseddroughtineasternCuba(Álvarez& degrees;exposurevariesbetweenthesouth Mercadet& al .,2012).Thatiswhy,restoring andtheeast.Atropicalclimatewithdrywin - thistypeofforest(duetoitsgeographical ter,subtype4bTh,isobservedwith5to6 positioninthelandscape)constantlyraisesits monthswithlittlerain(semi-dry).Themean strategicvalueforprotectingthecoastalzone annualtemperaturerangesfrom24-26ºC , andtheecosystemsbehindfromtheeffectsof withaveragepicksof30-32ºCand22-24ºC suchclimatechanges. res pectively.Themeanannualrelative hum idityis75to80%,thisbeing70to75%at 13:00handfrom80to85%at07:00h. Carbonero with majagüilla de costa Evaporationsrateis1700-1900mm (Colubrina elliptica with Helicteres (Montenegro,1991b,c,d).Itrainsabout1000 semitriloba ; Table 10 ) mm,withalittlerainyseasonfromNovember toAprilandarainyseasonfromMayto Physiognomyandfloristiccomposition. October.Itisthereforeconsideredahot Thisforesttypecorrespondstoalatesecon- trop ica lclimatewithdryandsemi-drywinter, 56 Reyes O.J. Lazaroa 37 2016: 43-103 with5-6drymonths.Commonly,itisstrongly Forallthedifficultconditionsoftheoecotope: affectedbysaltsprayseabreezes. topographicposition,soil,hydraulicstress, Geologically,itisobservedonrocksfrom etc,thistypeofforestgrowsunderextreme theCobreGroup(Méndez& al .,1994).The conditions,whichisexpressedinitsmicro - soilisfersialliticreddishbrown(Renda, phyllandsclerophyllousnature.Therefore,it 1989),yellowish,veryshallow,verygravelly, isconsideredasverydry(0)andlittlefertile highlyerodedandhastruncatedprofiles;the (A).ThetypologyclassificationisA xxx 0, amountofstonesonthesurfaceis60to90%. 4bTh b,V,C.e.,H.s.

Table10. Characteristiccombinationofcarbonerowithmajagüilladecosta (Colubrina elliptica with Helicteres semitriloba ,3plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Colubrinaelliptica D2-5V Carpodipteracubensis C1 V Bursera simaruba C1 V Bourreriavirgata C1 V Thouiniapatentinervis C4 IV Pseudocarpidiumavicennioides C4 IV Pictetiamucronata I4 IV Coccothrinaxgundlachii I4 IV Shrublayer Sennaatomaria -2 V Varroniaglobosa subsp. humilis -2 V Zanthoxylumelephantiasis -1 V Capparisflexuosa -1 V Erythroxylumhavanense -1 IV Herbaceouslayer Crotonlucidus -3 V Helicteressemitriloba -1 V Oploniatetrasticha -1 IV Malphigiasuberosa -1 IV Castelaspinosa -1 IV Zapotecagracilis -1 IV Lantanamontevidensis -1 IV Turneraulmifolia -1 IV Crossopetalum sp. - 1 IV Urochloahumidicola -1 IV Bothriochloapertusa -2 IV Lianas Stigmaphyllonsagreanum -1 V Commicarpusscandens -1 V Epiphytes Tillandsiausneoides -1 IV Tillandsiarecurvata -1 IV

Scatteredspeciesinthecanopylayer:yarúa( Caesalpinia violacea ),liriosantana( Exostema caribaeum )andjatía. Localities:inthepremountainareaofthesouthernslopeofSierradelTurquino. Reyes O.J. Lazaroa 37 2016: 43-103 57

Distribution:Itcoversthebottomofthe NW),soitisprotectedfromthemarinesalt southernslopeofSierradelTurquino. spray. Termsforforestmanagement.Asitisalate Inthisarea,rainfallisabout800mm, secondarycommunitywithsuchdifficult mainlyconcentratedfrommaytooctober;in topographicandecologicalconditionsitmust turn,thehumidityislowandtemperaturesand beprotected,withtheprimaryobjectiveof sunshineareveryhigh,causingalargeevapo - preservationofthesoil,thewaterandthe ration(Montenegro,1991a,b,e;Crespo, micro-basins. 1989).Itisconsideredasthebeginningofthe tropicalclimatezonewithdrywinter,semidry 4bThsubtype,with5to6monthswithlittle Cuyá with guairaje ( Sideroxylon rain. salicifolium with Eugenia maleolens ; ItisdevelopedonmarlsfromLaCruz Table 11 ) formation.Soilsaresialliticbrown(sensu Hernández& al .,1994),loamy,fromvery Physiognomyandfloristiccomposition. shallowtoshallow,sometimeswithgravel Evenwhenithasbeenrelativelyanthr opized, fromthemarl.Thereforethisedatopemust thecanopylayerofthissemideciduous beconsidereddry(1)andlittlefertile(A), microphyllforesthasbetween8and12m andthepotentialproductioncapacityiscon - tallinitshighestextension;exceptionally sideredbad(IV).Thetypologyclassification from7to8mwithemergingfromabout10 isasfollowsA1,4bTh b,IV,S.s.,E.m. m.Coverageissparsetodense(70-100%). Duetothesteepslope,thewithered Inthegroupofconstantspecies,themain leavescoverabout60%ofthesurface.It speciesare:cuyá( Sideroxylon salicifolium ), oftenaccumulatesintheanteriorpartofthe almácigo,yaití andguairaje (Eugenia stem,whereitisthicker.TheLlayerranges maleolens ).Assubdominantspecies,there between1and1.5cm.TheFlayerisabout are:ciguilla( Guettarda elliptica ),liriosan - 0.5cm,butbeforethestems,itsometimes tanaandcarbonero . Intheshrublayer, maybe1.0cm.TheHlayerismissing whosecoverishigh(50-80%),thesubdomi- (Reyes&Martinez,2005). nantspeciesareyuraguana( Coccothrina Distribution.Itislocatedinthecoastalhills fragans ),cuabilla, carnedevaca (Maytenus aroundthebayofSantiagodeCuba,inthe buxifolia ) andyarúadecosta (Erythroxylum rotundifolium );andintheherbaceouslayer southcoastalterracesofSierraMaestra (mediumcover),thesubdominantisthejibá. (Núñez&Viña,1989). Therearedifferentspeciesamonglianas Termsforforestmanagement.According (Table11).Inthegroupofaccompanying totheecologicalconditionsinwhichit species,inthecanopylayer:yaya( Oxandra develops,thiscoastalforesthassemi-hard lanceolata ),ipilipil,guatapaná( Acacia andhardwoodtreespecies,aswellasslow macracantha )andmacaguey(toad)( Guapira andveryslowgrowing.Therefore,itshould obtusata )aresubdominantspecies.The becarefullymanaged.Itshouldbepreserved shrublayerispoorandtheherbaceousrela - forbeinganecosystemofgreatimportance tivelyrich(Table11).Aggregatedspecies forcoastalprotection.Interventionsshould areconcentratedintheshrubandherbaceous beselectiveandspacedatleast,30years. layersandinthelianassynusiae. Regenerationshouldbeprotected,sinceitis generallydispersed.Thereareoecotopes Oecotopecharacteristics.Theelevationis showingadequateconditionstorestorethis 40to80ma.s.l,butnotdirectlyfacingthesea. kindofforest.Theyshouldbeplantedcuyá, Itisusuallyfoundonslopesof28from40 jatía,almácigo,baría,carbonero,liriosan - degreesandexposureistothenorth(NEto tanaandyarúa( Caesalpinia violacea ). 58 Reyes O.J. Lazaroa 37 2016: 43-103

Table11. Characteristiccombinationofcuyáwithguairaje (Sideroxylon salicifolium with Eugenia maleolens ,5plots ) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Sideroxylonsalicifolium (Dipholissalicifolia)D2V Burserasimaruba D2 V Gymnantheslucida I2 V Eugeniamaleolens I3 V Guettardaelliptica I1 V Exostemacaribaeum I1 V Colubrinaelliptica I1 V Oxandralanceolata I1 IV Leucaenaleucocephala I2 IV Guapiraobtusata I1 IV Acaciamacracantha O1 IV Shrublayer Randiaaculeata -1 V Coccothrinaxfragans -1 V Amyriselemifera -1 V Maytenusbuxifolia -1 V Erythroxylumrotundifolium -1 V Herbaceouslayer Erythroxylumhavanense -1 V Randiaspinifex -1 IV Melicoccusbijugatus -1 IV Crotonlucidus -1 IV Drypetes sp. - 1 IV Sclerialithosperma -1 IV Panicum sp. -1 IV Zanthoxylumfagara -1 III Celtistrinervia -1 III Thouiniatrifoliata -1 III Canellawinterana -1 III Ravenialeonis -2 III Zapotecagracilis -1 III Lianas Chiococcaalba -1 V Smilaxhavanensis -1 V Stigmaphyllonsagreanum -1 V Passiflorasuberosa -1 V Capparisflexuosa -1 IV C.cynophallophora -1 IV Lasiacisdivaricata -1 IV Gouanialupuloides -1 III Capparisgrisebachii -1 III Dalechampiascandens -1 III

Scatteredspeciesinthecanopylayer: aguedita (Picramnia pentandra ),peonía( Citharexylum sp.),yarúa,robleprieto (Ehretia tinifolia ), Licaria sp.,abeymacho( Jacaranda coerulea )andespueladecaballero( Bucida spinosa ). Reyes O.J. Lazaroa 37 2016: 43-103 59

Macaguey (toad) with curujey and Oecotopecharacteristics.Thiskindoffor - chicharrón ( Guapira obtusata with estispresentatthetopofthemogotes(karstic Hohenbergia penduliflora and Drypetes domehill)witha600to670maslaltitude. mucronata ; Table 12) Themicroreliefisdiverse,composedbybig, sharp,withlotsofholesandrupturesdogtooth Physiognomyandfloristiccomposition. rocks.Theaverageexposuretosunlightin Thec anopylayerisirregularinheight,usually thiszoneisof3to5hoursperday,withlower bet ween7and12m,itscoverissparse(50- andhigherpeaksduringtherainyanddrysea - 60%)andisconsideredaclearmicrophyll sonsrespectively.Themeanannualtempera - evergreenforest;theleavesaremainlyxero - tureis20-22ºC,withameanminimumof 16- morphicandmostlymicrophyllous.Inthe 18ºCandameanmaximumof26-28ºC groupofconstantspeciesthemacaguey(toad)is (Montenegro,1991b,e).Theclimateistropical themainspecie , andassubdominantthereare with4cThsubtypedrywinterand3to4 lirioamarilloorsuchel( Plumeria emargi nata ), monthswithlittlerain(moderatelydry),the ayúadesierra (Zanthoxylum cubense ),chichar - averagerainfallis1634mminLaTabla(1216 rón( Drypetes mucronata ),bone (Drypetes to2320mm),soitcanberegardedasrainy alba ),yuraguana( Coccothrinax elegans )and (sensuBorhidi,1991).Onaveragethereare94 Tabebuia bibracteolata .Theshrublayerhasa rainydaysintheyear.Arainyseasonisclearly mediumtohighcovers(30-60%),occasionally definedfrommarchtooctoberwith85%ofthe less.Thesubdominantspeciesaresigua( Ocotea rainfallwhilethedryseasontakesonlythe coriacea ),hicaquillo( Savia bahamensis )and 15%ofit.Themeanannualrelativehumidity yarúadecosta( Erythroxy lum rotundifolium ). is80-85%,rangingbetweena60-70%at13.00 Theherbaceouslayer,inwhichtherearemany h,and85-90%at07.00h(Montenegro,1991c, speciesconsideredasepiphytesinother d).Themeanannualevaporationis1300to ecosystems,hasahighcover(80-90%).Themain 1500mm,withadailyaverageof2.5-3mm speciesaremaguey( Agave underwoodii ),nome (Piche)andupto3.5-4mminsunnydays toques, curujey( Hohenbergia penduliflora ), (Montenegro,1991d). curujey( Tillandsia fasciculata ),andassub - Thesemogotesarestronglycarsifiedand dominant,flordelcáliz( Selenicereus grandi - dissecteddenudeplateaus(ViñaBayes,1991), florus ),jijira( Pilosocereus brooksianus ), withverticalwalls,composedbylimestones bejucolombricero (Philodendron lacerum ), Polypodium polypodioides , Epidendrum andmarblesfromCharcoRedondoformation. pygmaeum andflordesanpedro (Epidendrum Becauseofitstopographicposition,thisforest nocturnum ).Inthelianas,bejucodeberraco, isinaveryirregularmicrorelief,withbig, alambrillo( Smilax havanensis ),bejucosan sharp,withlotsofholesandrupturesdogtooth pedro, Platygyne dentata andbejucoman- rocks,wheresometimesthereisared,very teca (Passiflora penduliflora )aresubdominants . shallowsoil.Thatiswhyitisconsidereddry Amongtheepiphytesthemainspeciesisbarba (1),withalittlefertile(A)trophism. deindioandthecurujeyes( Tillandsia recurvata However,witheredleavesaccumulatesin and T. pruinosa )aresubdominants. theholes,sometimesreachingupto20cm, Inthegroupofaccompanyingspecies,the andformarootmat,wherenutrientrecycling subdominantsincanopylayerare:carbonero, takesplace.Theecosystemdependsonthis carmín( Alvaradoa arborescens )andcuabilla recycling.Duetotheparticularconditionsof (Amyris elemifera ).Inturn,intheshrublayer thisforest,thepotentialproductivecapacityis aguedita( Celtis trinervia ),cuabaprieta( Erithalis consideredverybad(V).Thetypologyclassi - fruticosa ),negracuba,yaití( Gymnanthes lucida ), ficationisasfollowsA xxx 1,4cTH d,V,G.o., brasil( Caesalpinia vesicaria )andpichajutía H.p.,D.m. (Hyperbaena paucinervis )havethesamecate - Distribution.Thisareaofmogotes,where gory;aswellasaguedita( Picramnia pentan - thisforesttypedevelops,occursinthenorth - dra ),cañuela( Cyrtopodium punctatum ),disci - ernslopeofSierraMaestra,formingthecen - plinilla( Rhipsalis cassutha ),amongothers tralpartofatypicalterritoryknownasCarso withintheherbaceouslayer.Aggregated deBaire(LaTabla,PalmadelPerro). speciesarealsodiverse(Table12). Termsforforestmanagement.Duetothe Thiskindofforesthavearichdifferential difficultenvironmentalconditionswherethese combinationthatdistinguishitfrommacaguey forestsevolvedandtheirtopographicalposi - (toad)withcurujeyandliriosantana( Guapira tion,th ehardwoodspecieswithaveryslow obtusata with Hohenbergia penduliflora and growthpredominate.Therefore,itisconsidered Exostema caribaeum )(Table11),duetosome a typicalobjectofconservationandshouldnot rainandaltitudinaldifferences. beexploitedatanycircumstances. 60 Reyes O.J. Lazaroa 37 2016: 43-103

Table12. Characteristiccombinationofmacaguey(toad)withcurujeyandchicharrón (Guapira obtusata with Hohenbergia penduliflora and Drypetes mucronata ,4plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Guapiraobtusata D2-3V Zanthoxylumcubense C1 V Drypetesalba C1 V Coccothrinaxelegans I1 V Tabebuiabibracteolata I1 V Plumeriaemarginata I1 V Colubrinaelliptica I1 IV Alvaradoaarborescens I1 IV Amyriselemifera I2 IV Spirotecoma sp. O 1 III Shrublayer Ocoteacoriacea -1 V Saviabahamensis -1 V Celtistrinervia -1 IV Hyperbaenapaucinervis -1 IV Herbaceouslayer Agaveunderwoodii -3 V Hohenbergiapenduliflora -3 V Tillandsiafasciculata -3 V Oploniatetrasticha -2 V Pilosocereusbrooksianus - 1V Philodendronlacerum - 1V Selenicerusgrandiflorus -1 V Epidendrumnocturnum -1 V Grisebachianthusplucheoides -1 IV Zamiaangustifolia -1 IV Vanilla sp. - 1 IV Arthrostylidiumangustifolium -2 IV Crotonlucidus -1 III Vernonia sp. - 1 III Ilex sp. - 1 III Pharusglaber -1 III Lianas Smilaxhavanensis -1 V Chioccocaalba -1 V Stigmaphyllonsagreanum -1 V Platygynadentata -1 V Capparisflexuosa -1 III Cynanchum sp. - 1 III Epiphytes Tillandsiausneoides -2 V Tillandsiarecurvata -1 V Differentialcombinationfrommacaguey(toad)withcurujeyandchicharrón Drypetesmucronata C1 V Erythroxylumrotundifolium -1 V Polypodiumpolypodioides -1 V Epidendrumpygmaeum -1 V Passiflorapenduliflora -1 V Tillandsiapruinosa -1 V Erithalisfruticosa -1 IV Thouiniatrifoliata -1 IV Reyes O.J. Lazaroa 37 2016: 43-103 61

Gymnantheslucida -1 IV Caesalpiniavesicaria -1 IV Picramniapentandra -1 IV Trichocentrumundulatum -1 IV Cyrtopodiumpunctatum -1 IV Rhipsaliscassutha -1 IV Polypodiumaureum -1 IV Encyclia sp. - 1 IV Octomeriatridentata -1 IV Pleurothallisgelida -1 IV Campiloneurumphyllitidis -1 IV Gesneriaheterochroa -1 III Auerodendroncubensis -1 III Solandralongiflora -1 III Isidoreapolyneura -1 III Peperomiarotundifolia -1 III Guzmaniamonostachya -1 III Tillandsiaschiediana -1 III T.balbisiana -1 III Scatteredspeciesinthecanopylayer:jaguey( Ficus laevigata ),ramoncow( Dendropanax arboreus ),liriosantanaand jaguey( Ficus perforata ). Localities:CarsodeBaire.

Macaguey (toad) with curujey and lirio Oecotopecharacteristics.Becauseitisalso santana ( Guapira obtusata with developedinthetopofthemogotesofCarso Hohenbergia penduliflora and Exostema deBaire,theecologicalconditionsandforest caribaeum ; Table 13) managementarethesameoftheforestpre- viouslydescribed[macaguey(toad)withcuru - Physiognomyandfloristiccomposition. jeyandchicharron].However,therearesome Thecanopylayerofthisclear-microphyll differencesinitsdistribution,sincethisforest evergreenforestvariesbetween7and12min islocatedinLasManuelasandinLaPimienta. heightanditscoverissparse(50-60%).In Thus,itissomewhatlowerandhigherthanthe thegroupofconstantspecies,themainspecies otherdescribedforest,respectively,which ismacaguey(toad)andthesubdominantsare mayresultinsomerainfalldifferences.The yuraguana( Coccothrinax elegans )and typologyclassificationisasfollowsA xxx 1, Tabebuia bibracteolata .Intheshrublayer, 4cTh d,V,G.o.,H.p.,E.c. withvariablecoveragebetweenlowandhigh (15and80%),isonlysubdominantsigua (Ocotea coriacea ).Intheherbaceouslayer, Sigua with carmín ( Ocotea coriacea with withahighcover(50and80%),nometoques Alvaradoa arborescens ) andcurujeyes( Hohenbergia penduliflora and Tillandsia fasciculata )arethemainspecies . Physiognomyandfloristiccomposition. Concerninglianas,alambrillo,bejucodever - Thistypeofforest(2plots)isanearlycom - raco,bejucolombriceroandflordelcáliz are munity;itcomesfromexcessivedisturbance subdominants.Inthegroupaccompanying andfire,soitiscurrentlyahomeostasisI. species,inthecanopylayer,cuabilla,boneand Carmín( Alvaradoa arborescens )isthemain suchelaresubdominants.Otherspeciesfrom speciesthatmakethecurrentstability.The theshrubandherbaceouslayers,andlianasare canopylayerisquiteuneven,withaheight foundinTable13.Inthecanopylayer,among between6and11m;thecoverageissparse, theaggregatespeciesgroup,thereareayúade rangingbetween60and70%.Themostabun - sierra( Zanthoxylum cubense ),carmínandfew dantspeciesarecarmín(D, A-C3-4),sigua speciesinotherstrataandsynusiaes.Also, (D,A-C1-2)andsometimeslirioamarillo(C, liriosantana( Exostema caribaeum ),panetela A-C1),macaguey(toad)(I,A-C1),ayúade (Phyllanthus epiphyllanthus )andpasionaria sierra(I,A-C1),pichajutía( Hyperbaena decerca( Passiflora sexflora )arepresentin paucinervis ,0,A-C1),penda(I,A-C1), lirio thedifferentialcombination. santana(I,A-C1),carbonero(I,A-C1) and 62 Reyes O.J. Lazaroa 37 2016: 43-103 guamá (Lonchocarpus longipes ,0,A-C1)are Intheshrublayer,coverageishigh alsoseeninthislayer.Otherscatteredtree (50-60% ),anditismediaattheherbaceous speciesare:raspalengua( Casearia hirsuta ), (30-40%),whichisalsotherichestin ag ueditasweet,aguedita( Picramnia penta ndra ), species. yaití,cuabillaandpolebox( Allophyllus cominia ).

Table13. Characteristiccombinationoftheforestmacaguey(toad)withcurujeyandliriosantana (Guapira obtusata with Hohenbergia penduliflora and Exostema caribaeum ,5plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Guapiraobtusata D3 V Coccothrinaxelegans C1 V Tabebuiabibracteolata I1 V Amyriselemifera I1 IV Hyperbaenapaucinervis O1 IV Drypetesalba O1 IV Plumeriaemarginata O1 IV Zanthoxylumcubense C1 III Alvaradoaarborescens.I1III Shrublayer Ocoteacoriacea -1 V Saviabahamensis -1 III Herbaceouslayer Oploniatetrasticha -2 V Hohenbergiapenduliflora -2 V Tillandsia fasciculata -2 V Agaveunderwoodii -2 IV Vernonia sp. - 1 IV Grisebachianthusplucheoides -1 IV Pilosocereusbrooksianus -1 IV Epidendrumnocturnum -1 III Lianas Philodendronlacerum -1 V Selenicerusgrandiflorus -1 V Smilaxhavanensis -1 V Chioccocaalba -1 V Vanilla sp. - 1 III Stigmaphyllonsagreanum -1 III Epiphytes Tillandsiausneoides L. - 1 IV Differentialcombinationfrommacaguey(toad)withcurujeyandliriosantana Exostemacaribaeum -1 III Phyllanthus epiphyllanthus -1 III Passiflorasexflora -1 III Scatteredspeciesinthecanopylayer: baría,mabóadesabana( Cameraria retusa ),penda( Citharexylum caudatum ), carboneroandguairaje( Eugenia asperifolia ). Reyes O.J. Lazaroa 37 2016: 43-103 63

Ecologyanddistribution.Itcomesontop Oecotopecharacteristics.Thisforestis ofthestronglydissectedcarsifiedplateaus, presentinsmallaltitudes,between230and mogotes,whichoccupiesthegeographical 240maslTheslopeissmall,about5degrees, areaknownasCarsodeBaire.Thealtitude witharelativelyuniformmesoandmicrore - variesbetween600and650maslWeather lief.Theexposureisvaried.Climatically,itis conditionsaredescribedinmacaguey(toad) intheareawheretheinfluenceofSierra withcurujeyandchicharrón,sincetheyare Maestraends.Ithasatropicalclimatewithdry locatedinthesameterritory.Althoughitisat winter,4bThsubtype,with5to6monthswith thetopofthemogotes,itoccupiesthe littlerain(semidry).Inthearea,itrains edatopeswithshallowtoveryshallowsoils, between1000and1200mm(mediumrainy), withabout40%ofrocksonthesurface. distributedinarainyseasonfromMaytoO cto- Becauseofitstopographicposition,itis ber,andalessrainyfromNovembertoApril. exposedtotheaverageclimaticconditionsof Ithasfrom40to60dayswithprecipitation theregion.TheLlayerisvariableinsite,it annually.Themeanrelativehumidityis75to rangesbetween0.5to2cm,theFlayerfluctu - 80%(Montenegro,1991b,c)andevaporation atefrom0.2to1cmandtheHlayeroccasion - variesbetween2000and2200mm(Crespo, allyitreaches1.5cm,withrootlets. 1989).Themeanannualtemperatureranges Asthegeneralconditionsaresimilarto from24-26ºC,withanaveragehighof32-34ºC macaguey(toad)withcurujeyandchicharrón, andanaveragelowof18-20ºC. thereforeitisalsoconsideredatypicalobject ItoccursinlimestonefromCharco ofconservationandshouldnotbeexploitedat Redondogeologicalformation.Thesoilis anycircumstances.Thetypologyclassifica - brown,veryshallow,sometimesrockscover tionisconsideredB1,4cTh d,IV,O.c.,A.b. upto80%ofthesurface.Witheredleaveshave anLlayerbetween2and2.5cm,andanF layerof0.5to1cm.TheHlayeristhinand Jocuma with guao ( Sideroxylum formsarootmatofaround1cminsome foetidissimum subsp. foetidissimum with places.Itishenceconsideredasadryand Comocladia dentata ; Table 14) moderatelyfertileedatope.Theref ore,its potentialproductivecapacityisconsideredbad Physiognomyandfloristiccomposition.This (IV).Thetypologyclassificationisa sfollows semideciduousmicrophyllforestwasaltered B1,4bTH c,IV,Rm*,S.f.f.,C.d. byintenseselectivelogging,whichhas Distribution.Thisforesthasamorewide - changeditsstructure.However,itmaintains spreaddistributioninthehillsofthenorthern itsfloristiccomposition;therefore,nowadays, slopeofSierraMaestra.Itwasstudiedatthe itcanberehabilitated. placeknownasElGranizo,betweenJiguaní Thecanopylayerisbetween7and10m andBaire,inthehillsofnorthernedgeof inheight,soitisconsideredalowforest.It SierraMaestra. hasalsoemergingabout15m.Coverisvery Termsforforestmanagement.Asinterven - sparse(30-50%).Themainspeciesare tioncontinues,thisforesthaslostpartofits jocuma( Sideroxylon foetidissimum subsp. structure.However,therearemanyvaluable foetidissimum )andguao( Comocladia dentata ). speciesfromtheforestrypointofview,which Thesubdominantspeciesarealmácigo,sigua haveanacceptableparticipationinthecanopy (sometimesabundant),cuyá,liriosantana layerandaverygoodregeneration(jocuma, (locallyabundant),guara( Cupania glabra ), cuyá,sigua,guara,yaicuaje).Therefore,inter - jíablanca( Adelia ricinella )andaromablanca ventionsshouldbelimitedtorestrictguaoand (Alvaradoa amorphoides ).Inthecanopy frijolillointhecanopylayerandreleasethe layerofthegroupofaccompanyingspecies, seedlingsofthemostvaluablespeciesinthe yaicuaje( Exothea paniculata )issubdomi - lowerones.Itcanbealsoenrichedwith nant. mahogany( Swietenia mahagoni ),cedar Theshrublayerhasahighcover(80to (Cedrela odorata ),yarúa,etc. 90%).Subdominantandabundantareguairaje (Eugenia sp.)andguao,therearetooraspalen - guaandthejibá,amongothersspecies.Inthe Group of cuyá with cúrbana ( Sideroxylon herbaceouslayerwithanaveragefrom20to salicifolium with Canella winterana ) 30%,arepresent Zapoteca gracilis ,amorseco (Desmodium incanum ),cortadera( Scleria Physiognomyandfloristiccomposition.This lithosperma )andcanutillo( Commelina foresttypeconsistsofthreestagesofsucce- elegans ).Lianasarediverse.Thecomposition ssionaldevelopmentcorrespondingtoalate oftheotherstrataandsynusiaes(whichare community(sensuBudowski,1985;Capote& many)isfoundinTable14. al .,1988;Reyes&Acosta,2005).Differences 64 Reyes O.J. Lazaroa 37 2016: 43-103

Table14. Characteristiccombinationofjocumawithguao (Sideroxylon foetidissimum subsp. foetidissimum with Comocladia dentata ,3plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Sideroxylonfoetidissimum subsp. foetidissimum P3 V Comocladiadentata D2 V Burserasimaruba P1 V Sideroxylonsalicifolium C1 V Ocoteacoriacea C1 V Exostemacaribaeum C1 V Cupaniaglabra C1 V Adeliaricinella O1 V Alvaradoaamorphoides O1 V Exotheapaniculata C1 IV Shrublayer Eugenia sp. - 3 V Picramniapentandra -1 V Caseariahirsuta -1 V Erythtroxylumhavanense -1 V E.rotundifolium -1 V Varroniaglobosa -1 V Capparisflexuosa -1 IV Coccolobadiversifolia -1 IV Erithalisfruticosa -1 IV Chrysophylumoliviforme -1 IV Guettarda sp. - 1 IV Sennaatomaria -1 IV Zanthoxylumfagara -1 IV Herbaceouslayer Desmodiumincanum -1 V Zapotecagracilis -1 V Sclerialithosperma -1 V Commelinaelegans -1 V Lasiacisdivaricata -1 IV Selenicereus grandiflorus -1 IV Rivinahumilis -1 IV Oeceoclades maculata -1 IV Agave sp. - 1 IV Lantanacamara -1 IV Urochloahumidicola -1 IV Lianas Pisoniaaculeata -1 V Smilaxhavanensis -1 V Stigmaphyllonsagreanum -1 V Gouanialupuloides -1 V Serjaniadiversifolia -1 V Chiococcaalba -1 V Turbinacorymbosa -1 IV Tragiahexandra -1 IV Epiphytes Tillandsiafasciculata -1 IV

Scatteredspeciesinthecanopylayer:cedar( Cedrela odorata ),carbonero,bijáguara( Colubrina ferruginosa ),guamá (Lonchocarpus domingensis ),penda,roble( Tabebuia sp.),mamoncillo( Melicoccus bijugatus ),mahogany( Swietenia mahagoni ),yaití,boneandcúrbana( Canella alba ). Reyes O.J. Lazaroa 37 2016: 43-103 65 aregivenbythedegreeofsuccessionaldeve- Cuyá with cúrbana and guatapaná lopmentandtheecologicalconditions,mainly (Sideroxylon salicifolium with Canella exposureandpositionontheslope.Therefore, winterana and Acacia macracantha ; Table 15 ) astheyarelocatedinthesamearea,theywill bestudiedtogether,specifyingonlythe Thistypeofforestistheleastadvancedtransi - diffe rences.Thecanopylayerisfromsparseto tionalsuccessionalstage.Here,frijolilloand slightlysparse(70to90%)andreaches guatapanástillplayanimportantroleinthe between9and12mtall.Floristically,itis canopylayer,whichisslightlysparse.The quiteheterogeneous;atotalof41specieswas coverofguairaje andcuyáisstillverylow.In foundreachingthislayer. thegroupofconstantspecies , onlyguatapaná Oecotopecharacteristics.Thestudiedsiteis iswithinthemainspeciesgroup.Frijolillo, hillsbetween20and100maslMostofitis cuyá,cúrbanaandalmácigoaresubdominants. locatedonthenorthsideoftheCoastalTerraces Intheshrublayer,whosecoverageisfrom southofSierraMaestra,sotheirsecondary mediumtohigh(30-70%),onlyguairajeis watershedanddells(thatonlyrunwithcon - consideredasmainspecies,theothersaresub - siderablerainfall)arepreferablynorth-northwest dominants.Thesamehappensintheherba - directedandtoalesserdegree,northwest. ceouslayer(itisfromlowtohigh,10-90%). Themostfrequentinclinationisbetween Usually,inthistypeofforest,soilshavea 20-40º .Themicroreliefonthesideisgenerally higherdegreeofdevelopmentandmaybe flat.Themeanannualtemperatureisaround consideredasshallow.Theypreferentially 26°Candtheaveragemaximumisabout32°C. occ upythebottomoftheslope(lowerbasin) Annualprecipitationisnear800mm(little andalthoughtheyareusuallysteep(30-45º) rainy)andthemeanannualevaporationis are directedtowardsthenorth(NE-NW). between1700and1900mm(Montenegro, Therefore,thehumidityhereismorefavorable 1991a,b,d).Therefore,itisconsideredatropi - thaninothertypesofforestinthisseries(dry calclimatewith4bThsubtypedrywinter,with withatendencytofresh). 5to6monthswithlittlerain(semidry). Thedifferentialcombinationhascabode Fromthegeologicalpointofview,thesehills hacha( Trichilia hirta )asamainspeciesand arecomposedbymarlsfromLaCruzformation jocumaanduvita( Cordia alba )assubdominants. TherestofthespeciesarefoundinTable15. (ComisiónCubano-Húngara,19 76).Soilsare brown(withcarbonate),andtheyarefrom sh allowtoveryshallow.Theyevenreactwith Cuyá with cúrbana and black ebony saliva.Itisusuallyclayloam,sandyclayor (Sideroxylon salicifolium with Canella sandy;stonyingreaterorlesserextent,with winterana and Diospyrus grisebachii ; Table 16 ) smallandmediumstones.Therefore,ingeneral, theedatopeisconsideredasdry(1)andlittlefer - Itisfoundinthelowerslopesoftheentirecom - tile(A),andthepotentialproductioncapacityis munity,asitsinclinationonlyoccasionally inferredasbad(IV).Thetypologyclassification exceeds20degrees.Althoughthepositionon isconsideredA1 x,4bTh b,IV,S.s.,C.w. theslopeisvariable,itismostcommonlyfound Regardingthewitheredleaves,theLlayer intheupperpartthereof;then,therelative reachesabout2cm.TheFlayerisgenerally heightisusuallygreaterthan20m.Theexpo - absent,exceptforareaswithhigherearth - sureistothenorth(NE-NW).Here,theearth - wormactivityandtheHismissing.Thisacti- wormactivitydescribedaboveiscommon.This vity(whichishigherinthemoreadvanced typeofforestisamoreadvancedsuccessional successionalstages)isveryimportanthere, stagethantheabove(latesecondarycommuni - becauseitincorporatesorganicmatterintothe ty).Inthecanopylayer,cuyáisthemainspecies soil.Theyformamixedlayerof1to1.5cmof andcúrbana,guatapaná,frijolillo,almácigoand soilandhalf-decomposedwitheredleaves, liriosantanaarethesubdominantsones.The whichinfluencestheecosystemnut rients mainspeciesfromtheshrublayerisguairaje cycle.Duetothisrapiddecompositionof fallen (Eugenia axillaris ),withahighcover(50-80%). leavesandthetotalabsenceofrootmat,the Fromtheherbaceouslayer,cuabadeingenio fermentationprocesscanbeconsideredas (mediaoccasionallyhighcovers,40-70%) quick(Herrera&Rodríguez,1988). wasthemainspecie.Here,thenumberof Thethreesubtypes(successionalstages)are: treeseedlingsishigherthanintheprevious typeof forest.Inthedifferentialcombination, Cuyáwithcúrbanaandguatapaná, thesubdominantsspeciesfromthecanopy Cuyáwithcúrbanaandblackebony, layer are:blackebonyandnegracuba.Other Cuyáwithcúrbanaandliriosantana. subdominantspeciesfromtheshrubandherbaceous Specialfeaturesofthesubtypes. layersandlianasynusiaeareshowninTable16. 66 Reyes O.J. Lazaroa 37 2016: 43-103

Table15. Cuyáwithcúrbanaandguatapaná (Sideroxylon salicifolium with Canella winterana and Acacia macracantha ,6plots) Species Vertical Abundance- Presence structure Coverage Degree Canopylayer Acaciamacracantha D2 V Sennaatomaria D1 V Sideroxylonsalicifolium C1 V Canellawinterana C1 V Burserasimaruba C1 V Lonchocarpusdomingensis I1 III Shrublayer Eugeniaaxillaris -2 V Coccothrinaxfragans -1 V Comocladiadentata -1 V Bourreriavirgata -1 V Randiaaculeata -1 IV Herbaceouslaye r Sclerialithosperma -1 V Lasiacisdivaricata -1 V Paspalumbreve -1 V Zapotecagracilis -1 IV Lantanacamara -1 III Tremamicrantha -1 III Lianas Stigmaphyllon sagreanum -1 V Chiococcaalba -1 V Smilaxhavanensis -1 IV Cissampelospareira -1 III Differentialcombinationfromcuyáwithcúrbanaandguatapaná Canopylayer Trichiliahirta D2 V Sideroxylonfoetidissimum subsp. foetidissimum C1 V Cordiaalba C1 V Shrublayer Ehretiatinifolia -1 V Turneraulmifolia -1 V Herbaceouslayer Erythroxylumhavanense -1 V Chrysophyllumoliviforme -1 V Caseariahirsuta -1 V Urochloahumidicola -1 V Heliotropiumindicum -1 IV Commelinaerecta -1 IV Desmanthusvirgatus -1 IV Lianas Gouanialupuloides -1 V Serjaniadiversifolia -1 V Salacianipensis -1 IV Epiphytes Tillandsiarecurvata -1 IV

Scatteredspeciesinthecanopylayer: carbonero,jiablanca( Adelia ricinella ),guara( Cupania glabra ), Citharexylum ternatum ,guaguasí( Zuelania guidonia ),sigua, blackebony (Diospyrus grisebachii ),guamá( Lonchocarpus longipes ) andbrasil( Caesalpinia vesicaria ). Reyes O.J. Lazaroa 37 2016: 43-103 67

Table16. Cuyáwithcúrbanaandblackebony (Sideroxylum salicifolium with Canella winterana and Diospyrus grisebachii ,8plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Sideroxylonsalicifolium D3 V Canellawinterana C1 V Acaciamacracantha C1 V Sennaatomaria C1 V Burserasimaruba C1 V Exostemacaribaeum I1 V Sideroxylonfoetidissimum subsp. foetidissimum C1 IV Colubrinaelliptica C1 IV Bourreriavirgata I1 IV Shrublayer Eugeniaaxillaris -3 V Comocladiadentata -1 V Randiaaculeata -1 V Lantanamontevidensis -1 IV Zanthoxylumfagara -1 IV Herbaceouslayer Crotonlucidus -2 V Coccothrinaxfragans -1 V Lasiacisdivaricata -1 IV Paspalumbreve -1 IV Sclerialithosperma -2 IV Zapoteca gracilis -1 III Varroniaglobosa subsp. humilis -1 III Morindaroyoc -1 III Erytrhroxylumrotundifolium -1 III Lianas Tragiahexandra -1 V Chiococcaalba -1 V Smilaxhavanensis -1 V Stigmaphyllonsagreanum -1 IV Salacianipensis -1 IV Passiflorasuberosa -1 IV Gouanialupuloides -1 III Distictisrhynchocarpa -1 III Triopterisrigida -1 III Differentialcombinationfromcuyáwithcúrbanaandblackebony Canopylayer Diospyrusgrisebachii I1 V Thouiniapatentinervis I1 IV Shrublayer Tremamicrantha -1 IV Erithalisfruticosa -1 IV Lantanacamara -1 IV Herbaceouslayer Spermacocelaevis -1 IV Bourreriasetoso-hispida -1 IV Varronialineata -1 IV Euphorbiaheterophylla -1 IV Lianas Cissusverticillata -1 IV Scatteredspeciesinthecanopylayer: cuabilla( Amyris elemifera ),guaguasí,robleprieto,guamádecosta (Lonchocarpus longipes ),guamá( Lonchocarpus domingensis ),guara,sigua, Citharexylum ternatum ,brasil,jiablan - caandraspalengua( Casearia hirsuta ). 68 Reyes O.J. Lazaroa 37 2016: 43-103

Cuyá with cúrbana and lirio santana Intheshrublayer(highcover,50-80%) (Sideroxylon salicifolium with Canella guairaje( Eugenia axillaris )isthemainspecie; winterana and Exostema caribaeum ; Table 17 ) whileintheherbaceous(mediatoheigcovers, 40-70%)itiscortadera( Scleria lithosperma ). Itisgenerallyobservedinexposurestothe Theothersubdominantspeciesinbothlayers, west(NtoSSW),onslopesof20-40º and aswellaslianasareshowninTable17.The generally,inthemidtoupperhillside,sothe proportionoftreeseedlingsishighinthese relativeheightisgreaterthan20m.The strata,whichalsoconfirmedhowadvanced microreliefisoftenslightlywavy. thesuccessionalstageis. Thisforestisalsoalatesecondarycommu - Thesoilisveryshallow,sometimesgra- nity;themaintreepioneers(guatapanáandfri - vellyandwithsomestonesonthesurface. jolillo)generallyhavediminishedvitalityand Duetoitsinclination,positionontheslope poororverypoorregeneration.Thedominant andexposure,itcanbeconsideredasadry(1) speciesinthecanopylayerarecuyáandcúr - andlittlefertile(A)oecotope. banaandtheabovementionedspeciesandthe almácigoarealsosubdominant.

Table17. Cuyáwithcúrbanaandliriosantana (Sideroxylon salicifolium with Canella winterana and Exostema caribaeum ,6plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Sideroxylonsalicifolium D2 V Canellawinterana D1 V Burserasimaruba C1 V Acaciamacracantha I1 V Sennaatomaria I1 V Shrublayer Eugeniaaxillaris -2 V Randiaaculeata -1 V Lantanamontevidensis -1 V Amyriselemifera -1 V Caesalpiniavesicaria -2 III Comocladiadentata -1 III Herbaceouslayer Sclerialithosperma -2 V Coccothrinaxfragans -1 V Crotonlucidus -1 V Zapotecagracilis -1 V Zanthoxylumfagara -1 V Paspalumbreve -1 V Simaroubalaevis -1 V Lasiacisdivaricata -1 IV Oploniatetrasticha -1 III Lianas Tragiahexandra -1 V Stigmaphyllonsagreanum -1 V Chiococcaalba -1 V Smilaxhavanensis -1 V Tragiavolubilis -1 IV Salacianipensis -1 III Distictisrhynchocarpa -1 III Differentialcombinationfromcuyáwithcúrbanaandliriosantana Exostemacaribaeum I1 V Colubrinaelliptica C1 IV Passiflorasuberosa -1 IV

Scatteredspeciesinthecanopylayer:negracuba,raspalengua,blackebony,jocuma,robleprieto,sigua,guara, Citharexylum ternatum andcuabaprieta. Reyes O.J. Lazaroa 37 2016: 43-103 69

Termsforforestmanagement.Asthese Guatapaná with frijolillo and raspalengua subtypesaresuccessionalstages,enrichment (Acacia macracantha with Senna atomaria withvaluablespeciesfromtheforestrypoint and Casearia hirsuta ; Table 19) ofviewshouldbeachievedinsilvicultural treatment,mainlyinthelessadvancedstages. Physiognomyandfloristiccomposition.This Someofthesespeciesare:cedar,bijáguara, foresttypeisanearlycommunity(ho- roble( Tabebuia sp.),mahogany,bone,cuyá, meostasisI),i.e.therelativeequilibriumstage jocuma,brasil,sabicú,dagameandbaría. takingplaceafterthefirstphasefiera(Reyes & al .,2004b).Thecanopylayerreaches between6and9mandisslightlysparse Guatapaná with frijolillo and cafecillo (80-90%).Themainspeciesareguatapanaand (Acacia macracantha with Senna atomaria frijolillo,especiallythefirstwhichdetermine and Bourreria virgata ; Table 18) thephysiognomyoftheecosystem.Theyform acontinuousandclearcanopythatallowsthe Physiognomyandfloristiccomposition.Low passageoflight.Uvitaisthesubdominant secondaryforestshowingdominanceofthorny species.Theshrub(20-60%)andherbaceous legumes.Itisanearlycommunity(homeostasisI) (30-70%)layershaveacoversfrommedium aftertotalorneartotaldestructionoftheprevious tohigh,exceptionallyless.Theyaremainly vegetation.Thecanopylayerislowandfairly composedbywoodymicroandnanophanero - opensparse(50to60%);itsheightisabout7to8 phytes,someherbaceousandlianas,which m.Themainspeciesatthegroupofconstant canbefoundinTable19. speciesareguatapanáandfrijolillo;whilethe Sinceguatapanaisthemaincontributorto subdominantareipilipilanduvita.Thespecies thewitheredleaves,andithasleptophyll fromtheotherstrataandsynusiaes,includingthe leaflets,theLlayerisgenerallylessthanacen - groupofaccompanyingspeciesandofdifferen - timeterandoftenscattered.TheFlayerisbare - tialcombinationareshowninTable18.The lynoticeableandtheHonlycontains(some - differencewiththeshrublayerisoftenunclear. times)remainsontheground.Forthisandthe Oecotopecharacteristics.Itcanbeseen absenceofrootmat,fermentationcanbecon - below150ma.s.l,showingaflatmicrorelief; sideredquick(Herrera&Rodriguez,1988). exposuresarevaried,mainlytothesouth. Oecotopecharacteristics.Thestudiedareas Theslopevariesbetween15-25º.Themean arehillsbetween20and100maslTheirse- annualtemperaturerangesfrom24-26ºC.The condarywatershedanddells(whichonlyrun meanannualrelativehumidityis75to80%and withconsiderablerainfall)arepreferably evaporationis1700to1900mm(Montenegro, north-westdirectedandtoalesserdegree, 1991a,b,c).Itrainsabout1000mm,withlittle northwest.Themostfrequentinclinationis rainyseasonfromNovembertoAprilanda between20and40degrees,themicroreliefon rainyfromMaytoOctober.Itisthereforecon - thesideisgenerallyflat.Itpreferssunny sideredatropicalclimatewithdrywinter, exposures,especiallytothesouthwest.The 4bThsubtype,with5-6drymonths(semi-dry). relativeheightislessthan20mingeneral. Itis alsoheavilyinfluencedbythesaltspray Themeanannualtemperatureisaround26°C seabreezes.Itoccursonsoilsderivedfrom andthemeanmaximumisabout32°C.The volcanogenicsedimentaryrocksandlimestones. annualprecipitationisnear800mm(little Thesoilisfersialliticreddishbrown(Renda, rain)andthemeanannualevaporationis 1989),veryshallowandhighlyeroded. between1700and1900mm(Montenegro, Therefore,itisconsideredasverydry (0)and 1991a,b,c);soitcanbeconsideredatropical littlefertile(A).Thepotentialproductioncapa- climatewithtwodryperiods,5aThsubtype, cityisinferredasverybad(V)andthetypology extremelydrytropicalbixeric. classificationisA0,4bTh b-c ,V,A.m.,S.a.,Bv. Geologically,thesehillsarecomposedby Distribution.Itwasstudiedinthesouthern PliocenemarlfromLaCruzformation.Soils partofSierradelTurquino,nearthecommunity arebrown(carbonate),fromshallowtovery ofLasBrujas(SantiagodeCubaProvince). shallowandevenreactwithsaliva.Theyare Termsforforestmanagement.Astheyare commonlysandyloamandsometimeslocally anearlycommunity(homeostasisI)afterthe gravelly.Itisoftenrockywithsmallandmid- destructionoftheoriginalforest,ithasvery stonesonthetopoftheprofile.Thelaminar littlediversityoftreespeciesintheshruband erosionisvariable;insomeplacesitisweak, herbaceouslayers.Thatiswhy,anenrichment inotherisstrongthough.Regardingthe withspeciesadaptedtotheoecotopeis edatopemoisture,itcanbeconsideredasdry advised;suchas:jatía,baría,carbonero, (1)andlittlefertile(A).Thetypologyclassifi - majaguilla,almácigo,sigua,cuyá,yarúaand cationisasfollowsA x1,5aTh b,IV,A.m.,S.a., macaguey(toad). C.h. 70 Reyes O.J. Lazaroa 37 2016: 43-103

Table18. Characteristiccombinationofguatapanáwithfrijolilloandcafecillo (Acacia macracantha with Senna atomaria and Bourreria virgata ,4plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Acaciamacracantha D3 V Sennaatomaria D2 V Cordiaalba C1 V Herbaceouslayer Bothrichloapertusa -2 V Zapotecagracilis -1 V Commelinadiffusa -1 V Spermacocelaevis -1 V Malpighiacnide -1 IV Sidarhombifolia -1 IV Malvastrumcorchorifolium -1 III Urochloahumidicola -1 III Helicteres sp. - 1 III Coccothrinaxfragans -1 III Desmanthusvirgatus -1 III Zanthoxylumfagara -1 III Corchorus sp. - 1 III Bourreria sp. - 1 III Wissadulaperiplocifolia -1 III Sidaacuta -1 III Desmodium sp. - 1 III Lianas Stigmaphyllonsagreanum -1 V Dalechampiascandens -1 V Cissustrifoliata -1 V Chiococcaalba -1 IV Tournefortiavolubilis -1 IV Triopterisrigida -1 IV Centrosemavirginianum -1 III Commicarpusscandens -1 III Galactia sp. - 1 III Ipomoea sp. - 1 III Epiphytes Tillandsiarecurvata -1 IV Differentialcombinationfromguatapanáwithfrijolilloandcafecillo Bourreriavirgata I2 V Leucaenaleucocephala C1 V Varroniaglobosa subsp. humilis -3 V Wissadulaamplissima -1 V Lantanareticulata -1 V Phyllostyllonbrasiliensis -1 IV Adeliaricinella -1 IV Randiaaculeata -1 IV Scatteredspeciesinthecanopylayer: Casearia sp.,cuabaprieta. Reyes O.J. Lazaroa 37 2016: 43-103 71

Table19. Guatapanáwithfrijolilloandraspalengua( Acacia macracantha with Senna atomaria and Casearia hirsuta )inthemarlsfromtheSouthCoastalTerracesofSierraMaestra(6plots).

Species Vertical Abundance- Presence structure Coverage Degree Canopylayer Acaciamacracantha D4 V Sennaatomaria D2 V Cordiaalba C1 V Ehretiatinifolia C1 IV Shrublayer Coccothrinaxfragans -1 IV Lantanacamara -1 IV Herbaceouslayer Waltheriaindica -1 V Zapotecagracilis -1 V Lantanamontevidensis -1 V Wissadulaperiplocifolia -1 V Melochianodiflora -1 V Euphorbiaheterophylla -1 V Spermacocelaevis -1 V Bothriochloapertusa -1 V Urochloahumidicola -1 V Sclerialithosperma -1 V Lasiacisdivaricata -1 V Diclipteravahliana -1 IV Jatrophagossipifolia -1 IV Cenchrusciliaris -2 IV Commelinaerecta -1 IV Varroniaglobosa subsp. humilis -1 III Erythroxylumhavanense -1 III Hibiscusbrasiliensis -1 III Comocladiadentata -1 III Chromolaenaodorata -1 III Pavoniaspinifex -1 III Lianas Passiflorasuberosa -1 V Stigmaphyllonsagreanum -1 V Tournefortiavolubilis -1 V Centrosemavirginianum -1 V Desmanthusvirgatus -1 V Smilaxhavanensis -1 V Triopterisrigida -1 V Passiflorafoetida -1 IV Salacianipensis -1 IV Merreniabisecta -1 III Chiococcaalba -1 III Serjaniadiversifolia -1 III Epiphytes Tillandsiarecurvata -1 IV Tillandsiafasciculata -1 III Differentialcombinationfromguatapanáwithfrijolilloandraspalengua Caseariahirsuta -1 V Turneraulmifolia -1 V Paspalumbreve -1 V Echitesumbellata -1 V Plumbagoscandens -1 V

Scatteredspeciesinthecanopylayer: cuyá,jubaban( Trichilia hirta ),brasil,cúrbana,jíablanca,jocuma,penda,ague - ditadulce,carbonero,agalladecostaandcafecillo. 72 Reyes O.J. Lazaroa 37 2016: 43-103

Distribution.Itpresentsitsgreatestextent deredatropicalclimatewithdrywinter,sub - inthenortheasternpartofthemarlyhillsfrom type4bTh,with5-6drymonths(semi-dry). theSouthCoastalTerracesofSierraMaestra. Themostimportantsoilisthereddish Termsforforestmanagement.Asitisa brownfersiallitic;itsdepthvariesfromvery communitysimilartothepreviousforest shallowtomoderatelydeep.Itdrainswell, (homeostasisI),ithasverylittlediversityof bothinternalandexternal.Therefore,the treespeciesintheshrubandherbaceouslay - edatopeisconsideredasdry(1)andfromlittle ers.Thatiswhy,enrichmentwithspecies tomoderatelyfertile(B).Witheredleavesare adaptedtotheoecotopeisadvised;suchas: unevenlydistributed,theLlayerranges jatía,baría,carbonero,majaguilla,almácigo, between1.5and3cm,occasionallymore;in sigua,cuyá,yarúaandmacaguey(toad). isolatednanodepressionscanreachupto15 cm.FandHlayersarealmostimperceptible, andonlythelastisobserved,mixedwiththe Yaya with almácigo ( Oxandra lanceolada horizonA.Inturn,thepotentialproductive with Bursera simaruba ; Table 20) capacityisconsideredfrombad(IV)toregu - lar(III).Thetypologyclassificationisasfol - Physiognomyandfloristiccomposition. lowsB1,4bTh c,III-IV,O.l.,B.s. Althoughthisvegetationtype(semideciduous Distribution.Thisforestisatypicalsemi - mesophyllforest)hasbeenexploitedforlive - deciduousmesophyllforest,whichhadalarge stockdevelopment,theanalyzedareahasa areaintheflatandpremountainareas.Itwas highdegreeofnaturalness.Thecanopylayer studiedonthesouthernslopeofthemountain hascoveragefromsparsetoslightlysparse rangeofSierraMaestra.Nowadays,themost (50-90%)andaheightbetween7and15m.In extensiverelictislocatednearthemouthof thegroupofconstantspecies,almácigoand Turquinoriver. yaya( Oxandra lanceolada )arethemain Termsforforestmanagement.Almostall species;whilebaríaandguamácandelón areasfromthemesophyllsemideciduousfor - (Piscidia piscipula )arethesubdominant. esthavebeendestroyedforagricultureand Thesespeciesfromtheshrubandherbaceous livestockpurposes,soitisnecessarytopre - layers,aswellasthesynusiaesoflianasand servetheserelics,whicharerelativelyrichin epiphytesarefoundinTable20. treespecies. Inthegroupofaccompanyingspecies, jocuma,guárano( Cupania americana ),guási - ma,jobo( Spondias mombin ),jatía andsabicú Jubaban with guásima ( Trichilia hirta with (Lysiloma sabicu )aresubdominantspecies. Guazuma ulmifolia ; Table 21) Bone,yaití,guara,guaguasí,tengue (Poeppigia procera ),royalpalm( Roystonea Physiognomyandfloristiccomposition. regia ),ramonhorse( Trophis racemosa ),cuyá Thissecondaryforestisalatesecondarycom - andmajaguillaareaggregatedspecies.Plants munity(fieraIIandhomeostasisII)fromthe fromotherlayerareshowninTable20. successionalseriesthattakesplaceinthe Thespeciesfromtheshrublayerwitha mesophyllsemideciduousforest;themost coverfrommediumtohigh(40-90%)andthe advancedonehasmorespeciesinallstrata. herbaceouslayerwithmedium(30-50%)are Thecanopylayerisirregularinheightand foundinTable20.Thecompositionofbiolog - variesfrom7to12m.Coverisfromsparseto icaltypesisasfollows:40.5%ofthespecies dense(60-100%).Amongthegroupofcon - aretrees,24.3%areshrubs,13.5%formthe stantspecies,guásima(themostabundant) herbaceouslayer,2.7%areferns,16.2%are andfrijolilloarethemainspecies.Thesub - lianasandthe4.1%,epiphytes. dominantsarejubaban,baría,jobo,majagui- Oecotopecharacteristics.Themicrorelief lla,negracubaandyuraguana( Coccothrinax isgenerallyflat.Itlocallyshowsrockyout - gundlachii ).Inthegroupofaccompanying cropsonthesurface.Theslopeisalsovery species,almácigoissubdominantandaguedi - variable,rangingbetween20and40degrees. ta( Celtis trinervia )aggregated.Thespeciesin Theexposurehastwomaintrends:onetothe theshrublayerwithacoversfrommediumto eastandtheothertosouth.Thealtitudevaries high(40-80%)areinTable21,aswellasthe between120and200maslThemeanannual specieswithmediumcoverage(30-50%)from rainfallinthisareais1000to1200mmper theherbaceouslayerandlianas. year,i.e.moderatelyrainy.Therearetwodis - Oecotopecharacteristics.Itoccurs tinctperiods:thedryseason,fromNovember between40and100masl,withaninclination toApril(between300and400mm)andthe of20-37ºandinvariousexposures.The rainyseason,fromMaytoOctober(800and microreliefisrelativelyflat.Themean 1000mm)(Montenegro,1991a).Itisconsi- annualtemperaturerangesfrom24to Reyes O.J. Lazaroa 37 2016: 43-103 73

Table20. Yayawithalmácigo (Oxandra lanceolada with Bursera simaruba ,9plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Burserasimaruba D2 V Oxandralanceolata D2 V Cordiagerascanthus C1 V Piscidiapiscipula C1 V Sideroxylonfoetidissimum subsp. foetidissimum C1 IV Cupaniaamericana C1 IV Guazumaulmifolia C1 IV Spondiasmombin C1 IV Phyllostylonbrasiliensis I1 IV Lysylomasabicu I1 IV Drypetesalba C1 III Gymnantheslucida I1 III Cupaniaglabra var. glabra C1 III Zuelaniaguidonia C1 III Roystonearegia C1 III Trophisracemosa C1 III Sideroxylonsalicifolium C1 III Poeppigiaprocera I1 III Carpodipteracubensis I1 III Shrublayer Erythroxylumhavanense -1 V Picramniapentandra -1 V Coccothrinaxgundlachii -1 IV Trichiliahirta -1 IV Capparisflexuosa -1 III Caseariasylvestris var.myricoides -1 III Herbaceouslayer Olyralatifolia -1 V Oeceocladesmaculata -1 IV Urochloahumidicola -1 III Zapotecagracilis -1 III Rivinahumilis -1 III Lianas Serjaniadiversifolia -1 V Gouanialupuloides var lupuloides -1 V Pisoniaaculeata -1 V Stigmaphyllonsagreanum -1 V Epiphytes Tillandsiarecurvata -1 V

Scatteredspeciesinthecanopylayer:yaicuaje( Exothea paniculata ),yaba( Geoffroea inermis ),dagame (Calycophyllum candidissimun ), carbonero( Collubrina reclinata ),almendrillo( Prunus myrtifolia ),jagueymacho (Ficus membranacea ),ayua( Zanthoxylum martinicensis ),robleblanco( Tabebuia angustata )andceiba( Ceiba pen - tandra ). Localities:SouthernslopeofSierraMaestra.

26ºC,withameanannualhighof30-32ºC (slightlyrainylimit),withalittlerainyperi - andameanannuallowof22-24ºC.Themean odfromnovembertoaprilandarainy,from annualrelativehumidityis75-80%. maytooctober.Itisconsideredatropicalcli - Evaporationis1700-1900mm(Montenegro, matewithdrywinter,4bThsubtype,with5-6 1991a,b,c,d).Itrainsabout1000mm drymonths(semi-dry). 74 Reyes O.J. Lazaroa 37 2016: 43-103

Thesoilisreddishbrownfersiallitic,very Therefore,itisconsideredasadry(1)andli- shallow,gravelly,withapercentageofstones ttlefertile(A)edatope,soitspotentialproduc - from50-80%,exceptionallylower.This tivecapacityisconsideredbad(IV).The foresttypeoccupiesthemiddleandlower typologyclassificationisasfollowsA xx-xxx 1, zonesofthemicrobasin,sotheirwatersitua - 4bTh ,IV,T.h.,G.u. tionisrelativelymorefavorablethaninthe b-c upperbasins.Duetothelargeslope,withered Distribution.Itwasstudiedatthebottomof leavesareverysparsewhereitaccumulates. thesouthernslopeofSierradelTurquino. Sometimesamongthestones,anLlayer Termsforforestmanagement.Thisforest (1-2cm)andagenerallyimperceptibleFare canbeenrichedwithcedar,mahogany,baría, observed.Sometimessuchlayersaremixed. yarúaandayúa.

Table21. Characteristiccombinationofjubabánwithguásima (Trichilia hirta with Guazuma ulmifolia ,6plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Guazumaulmifolia D4 V Sennaatomaria I2 V Trichiliahirta D1 V Spondiasmombin C1 V Cordiagerascanthus C1 V Carpodipteracubensis I1 V Thouiniapatentinervis I1 V Coccothrinaxgundlachii I1 V Burserasimaruba C1 IV Celtistrinervia I1 III Shrublayer Varroniaglobosa subsp.humilis -2 V Phyllostylonbrasiliensis -1 V Erythroxylumhavanense -1 V Herbaceouslayer Melochianodiflora -1 V Lantanamontevidensis -1 V Desmodiumincanum 1V Commelinaelegans -1 V Wissadulaamplissima -1 V Bothriochloapertusa -1 V Urochloahumidicola -1 IV Pavoniafructicosa -1 IV Abutilon sp. - 1 IV Eupatorium sp. - 1 III Harrisiaeriophora -1 III Malpighiasuberosa -1 III Picramniapentandra -1 III Bidenspilosa -1I II Lianas Centrosemavirginianum -1 V Gouanialupuloides var. lupuloides -1 V Stigmaphyllonsagreanum -1 V Serjaniadiversifolia -1 IV Plumbagoscandens -1 IV Passiflorasuberosa -1 III Epiphytes Tillandsiarecurvata -1 V Scatteredspeciesinthecanopylayer:cuabilla,yarúa,bayúa (Zanthoxylum elephantiasis ),yabaandliriosantana. Localities:SouthernslopeofSierradelTurquino. Reyes O.J. Lazaroa 37 2016: 43-103 75

Table22. Characteristiccombinationofguáranowithguásima (Cupania americana with Guazuma ulmifolia ,4plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Guazumaulmifolia D2 V Cupaniaamericana D2 V Cordiagerascanthus C1 V Poeppigiaprocera C1 V Trichiliahirta C1 V Zanthoxylummartinicense C1 V Roystonearegia C1 V Sennaatomaria I2 IV Cupaniaglabra C1 IV Samaneasaman C1 IV Cecropia peltata C1 IV Cedrelaodorata C1 IV Guareaguidonia C1 IV Piscidiapiscipula I1 III Shrublayer Caseariasylvestris var.sylvestris -1 V Picramniapentandra -1 V Eugeniafloribunda -1 IV Erythroxylumhavanense -1 IV Caseariahirsuta -1 IV Sideroxylonfoetidissimum subsp. foetidissimum -1 IV Chrysophylumoliviforme -1 IV Cordiacollococca -1 III Psidiumguajava -1 III Oxandralanceolata.-1III Herbaceouslayer Desmodiumincanum -1 V Urochloahumidicola -1 V Lasiacisdivaricata -1 V Adiantumtrapeziforme -1 V Pavoniafruticosa -1 IV Spermacoceassurgens -1 IV Pharusglaber -1 IV Randiaaculeata -1 III Chromolaenaodorata 1III Oplismenussetarius -1 III Sidarhombifolia -1 III Capsicumfrutescens -1 III Petiveriaalliacea -1 III Campiloneurumphyllitidis -1 III Lianas Gouanialupuloides -1 V Ipomoeaalba -1 V Pithecocteniumechinatum -1 V Passiflorasuberosa -1 IV Serjaniadiversifolia -1 IV Tournefortiahirsutissima -1 IV Pisoniaaculeata -1 IV Salacianipensis -1 III Trichostigmaoctandrum -1 III Cissampelospareira -1 III Cissusverticillata -1 III Aristolochialindeniana -1 III Plumbagoscandens -1 III Hippocrateavolubilis -1 III Scatteredspeciesinthecanopylayer: bijáguara,penda,jagua( Genipa americana ),jatía,mije( Eugenia asperifolia ), majaguilla,yaití,jobo, fustete( Clorophora tinctoria ),yaba,anacahuita( Sterculia apetala ),dagame,bienvestido (Gliricidia sepium ),robleprieto,guatapaná,almácigo,guasimilladelpinar( Casearia arborea ),yagrumahembra (Schefflera morotottoni ),jagueymacho( Ficus membranacea ),dairy( Sapium jamaicense )andguaguasí. Localities: SierraMaestra. 76 Reyes O.J. Lazaroa 37 2016: 43-103

Guárano with guásima ( Cupania americana Amoroso with guásima ( Zanthoxylum with Guazuma ulmifolia ; Table 22) fagara with Guazuma ulmifolia ; Table 23)

Physiognomyandfloristiccomposition.This Physiognomyandfloristiccomposition. secondaryforestgrowsintheoecotopesfromthe Thissecondaryforestsuccessivestageknown semideciduousmesophyllforestandformsthe asfieraII.Thecanopylayerincreasesinsize latecommunities(fieraIIandhomeostasis withthedevelopmentoftheforest,from6-8m II)undertheseconditions.Thecanop y to10-12mandisconsidereddense(100%);in layervariesaccordingtothestageofdevelop - thelowerparts,norealdifferenceisobserved ment.Inplaceswhereitislower,itisbetween betweenthecanopyandtheshrublayers.Inthe 7and8m,whileinthemoresyngeneticaly groupofconstantsspeciesguásimaandfrijolil - evolvedplacesitreachesfrom12to15m. loaredominantwhileguarano,guara,amoroso Coverisslightlysparse(usually90%),occa - (Zanthoxylum fagara ),guatapaná,algarrobo, sionallyless.Inthegroupofconstantspecies, raspalenguaandcaimitillo( Chrysophyllum themainspeciesareguásimaandguárano oliviforme )aresubdominants.Inturn,inthe (Cupania americana ),whilebaría, tengue, groupofaccompanyingspeciesaresubdomi - jubabán, ayúaandroyalpalmaresubdomi - nants:jatia,ipilipil,mije( Eugenia floribunda ), nants.Inthegroupofaccompanyingspecies, aguedita( Picramnia pentandra ),guamá frijolillo, algarrobo( Samanea saman ),guára - hediondo( Lonchocarpus blainii ),aguedita nohembra, yagruma( Cecropia peltata ), cedar sweet,guayaba( Psidium guajava )and andyamagua( Guarea guidonia )aresubdomi - jubabán.Thecompositionoftheotherlayers nants,whileguamácandelon( Piscidia piscipula ) andsinusiaesarefoundinTable23. isaggregated.Thespeciesfromtheshrub Oecotopecharacteristics.Themicrorelief (mediatohighcovers,20-80%)andherba - isirregularandoccupiespreferablythemiddle ceous(media,20-50%)layers,aswellas andlowerbasins.Thestudywasrunatanalti - lianasandepiphytesarefoundinTable22. tudeof220to240masl,witha15to39 Oecotopecharacteristics.Ittakesplaceat degreeslopeandchangingexposure.Theclimat e analtitudebetween200and625masl,on istropicalwithdrywinter,4bThsubtype, slopesof20to40degreesandinvariousexpo - with5to6monthsoflittlerain(medium-dry). sures.Itlocatesinlowandpremountainareas The averagerainfallrangesfrom1000to fromSierraMaestra.Rainfallisseasonal, 1200mmandtheaveragetemperatureisaround between900and1200mm,i.e.,moderately 25ºC. rainy.Theclimateistropicalwithdrywinter, Itliesonrocksoftheundifferentiatedthe 4cThsubtype,3to4monthswithlittlerain CobreGroup.Soilsareyellowishbrown,shal - (moderatelydry).Soilsvaryfromfersiallitic low,withnoticeablesheeterosionandabout andbrowntoferrallitic,andfromshallowto 5%ofrockatthesurface.Thewitheredleaves moderatelydeep,sometimeswithstoneson areoftenscattered,inthelessstructurally thesurface.Humusisunderdeveloped.TheL evolvedpartsLandFlayersaremixedandare layerisbetween1and3cm.TheFlayerusu - nothickerthanacentimetre;onthecontrary, allypresentsfromvestigesto0.3cm,onlyin atthemoredevelopedlayerstheLisabout1.5 structurallycomplexareascanreach2cm cmandF0.5cm,Hismissing.Theedatopeis (withrootlets).Hlayerisnotcommon;excep - considereddry(1)andmoderatelyfertile(B); tionally,intheabovedescribedsitescanbe,in thereforethepotentialproductioncapacityis dispersedform,upto0.3cm.Therefore,the consideredbad(IV).Thetypologyclassifica - edatopeismoderatelyfertile(B),fromdryto tionisasfollowsB1,4bTh c,IV,Z.f.,G.u. fresh(1-2),andthepotentialproductive Distribution.Thestudywasrunatthesur - capacityisconsideredasregular(III).The roundingsofthetownElCobre(Santiagode typologyclassificationisB x1-2,4cTh b-c ,III, Cubaprovince). C.a.,G.u. Conditionsofforestmanagement.As Distribution.Thistypeofforestistheave- advancedsuccessivestage,itcanbeenriched rageconditionsofsemideciduousmesophyll withspeciesofforestvaluesuchascedar, forestintheSierraMaestra. mahogany,yarua,baria,jocuma,bijáguaraand Termsforforestmanagement.Itisrichin dagame,amongothers. species,44treespeciesareobserved.Then,its treatmentistokeepitassuch,sothatitcan developitsfullpotential.Correspondingoeco - Sigua with guara ( Ocotea coriacea with topestosuchtypeofforestcanberestored Cupania glabra ; Table 24) withspeciessuchascedar,mahogany,yarúa, baría,ayúa,jocuma,bijáguaraanddagame, Physiognomyandfloristiccomposition. formingamixedforest. Thecanopylayervariesbetween7and15m Reyes O.J. Lazaroa 37 2016: 43-103 77 heightanditscoverissparse,occasionally grows,managingonlyallowslong-termselec - dense(50-100%).Inthegroupofconstant tiveextractions(30orolder)toavoiddecrea- speciesitisonlysothesigua,whiletheguara, singratesofvaluablespeciesfromtheforestry jaguey( Ficus laevigata ),macaguey(toad), pointofview. yellowlilyandaguedita( Picramnia pentan - dra )aresubdominants;asforthegroupof accompaningspecies,thiscategoryispresent Yamagua with ramon cow ( Guarea guidonia withramoncow( Dendropanax arboreus ), with Dendropanax arboreus ; Table 25) bone,aguedita( Celtis trinervia ),cuabilla, morurorojo,raspalenguaandguairaje Physiognomyandfloristiccomposition. (Eugenia asperifolia ).Asaggregatedspecies, Thistypeofforest,relativelyrichintree thereare:abey( Abarema glauca ),ayuade specieswith58,isalatesecondarycommuni - sierra,baríaandpenda.Thespeciesinthe ty(fieraII),resultingfromtheneglectofco- shrublayer,lowsometimeshigh(10-90%), ffeeplantationsand/oralterationoftheforest. herbaceous(mediumtohigh,30-95%)andthe Itoccupiesscatteredareas,becausetheco- sinuciasoflianasandepiphytesarefoundin rrespondingsitesarecurrentlyinusebyagri - Table24. culturalecosystems.However,itisconsidered Oecotopecharacteristics.Thealtitude thatthefull-growncommunitywillhavea variesfrom480to645masl.Theslopeis floristiccompositionprettysimilartothecur - irregular,asitistheexposure.Themeanannu - rentone. altemperatureis20to22ºC(Montenegro, Thecanopylayerisdense(90-100%)and 1991a).Inthecentralareaofthestudiedterri - irregularinheight,between8and20m.Inthe torymeanannualrainfallis1634mm,varying groupofconstantspeciestherearemain intheanalyzedyearsbetween1216and2320 ramoncowandyamaguawhereas,thesub - mm.Thereare94rainydays(average), dominantare:ayúa,baría,sigua,yagrumaand rangingbetween76and111.Evaporation royalpalm.Inthegroupofaccompanying rangesfrom1300-1500mmandrelative speciesguásimaissubdominantandaggregate humiditybetween80and85%(Montenegro, are:redcedar( Cedrela cubensis ),jobo, 1991c,d).Accordinglythereisatropical jocuma,jaguey( Ficus membranacea ), weatherwithdrywinter,4cThsubtype,with macagua,guaguasi,jaguey( Ficus suffucans ), 3-4drymonths(moderatelydry). yaya,yaba,jubabán,agueditasweetandbone. Thisforestgrowsonlimestoneandmarble. Intheshrublayerwithafundamentally Itispresentinboththetopandbottomofthe mediacovers(20-30%,exceptionally60%) mogotesinareasofreddishbrownsoil,orin andherbaceouslowtohigh(10-90%)are dogtoothwithcavitiesanddiaclaseswithred observedasseedlingsandsmalltrees:aguedita soils,bothveryshallow.Generally,inthis (Picramnia pentandra ),raspalengua,ramon ecosystem,soilishighlyinvolvedintherecy - horse,guara,guarano,leviza,almendrillo( Prunus clingofnutrients.However,itoccursinless myrtifolia ),ocuje( Calophyllum antillanum ), extremeconditionsthatmacaguey(toad)with guayabillo( Guettarda calyptrata ),sweetpotato curujeyandchicharrón,occupyingthesame laurel( Ocotea f loribunda ),caimitillo,pole territory.Soitisconsideredalittlefertile boxandaguedita( Celtis trinervia ).Other (A xxx )anddrytrophism(1).Thewithered speciesoftheselayersandsinusiaesarefound leaveswelldeveloped,withanLlayer inTable25. between1.5and5cm;Fis0.2to3cm,andan Oecotopecharacteristics.Thisforestis Hsometimesconvertedintorootmat0.5to developedintheupperandlowermogotes 2.5cm,ofrootsandrootletsembeddedina whichareformedbydenudationplateaus,and matrixofhumus.Inplaceswithmoresoil,this dissectedsharply(ViñaBayés,1991),composed rootmatissmaller,withlittlehumus;inthis forlimestonesandmarbles.Thealtitudevaries casetherearemorerootletsinthefirstsoil inthetwostudyareas(LasManuelas,190-200 horizon.Consistentwiththedifficultcondi - masl)and(LaPimienta,565-650masl),the tionsinwhichthisforestdevelopsthepoten - slopeisbetween8and35degreesandexposure tialproductivecapacityisconsideredbad isnorthtosouth-southeast. (IV).Thetypologyclassificationisasfollows Theannualrainfallmeaninthisareais1 Axxx 1,4cTh d,IV,Rm*,O.c.,C.g. 634mm;frommarchtonovembertherainfall Distribution.Thistypeofforestislocatedin rateisover100mmwhilethedryseasontakes thegeographicalareaknownasCarsodeBaire placefromdecembertofebruary,withonly (Núñez&Viña,1989;ViñaBayés,1991).Itis 15%ofrainfall,sothisareacanbeconsidered presentinLasManuelasandLaPimienta. atropicalclimatewithdrywinter,subtype Conditionsofforestmanagement.Dueto 4cTh,with3to4drymonths(moderately theconditionsunderwhichthistypeofforest dry). 78 Reyes O.J. Lazaroa 37 2016: 43-103

Table23. Characteristiccombinationofamorosowithguásima (Zanthoxylum fagara with Guazuma ulmifolia ,3plots). Species Vertica Abundance- PresenceDegree structure Coverage Canopy+shrublayers Guazumaulmifolia D3 V Sennaatomaria C3 V Cupaniaamericana C1 V Cupaniaglabra C1 V Acaciamacracantha C1 V Samaneasaman C1 V Zanthoxylumfagara I1 V Caseariahirsuta I1 V Chrysophyllumoliviforme I1 V Phyllostylonbrasiliensis D1 IV Leucaenaleucocephala C1 IV Shrublayer Eugeniafloribunda -1 IV Picramniapentandra -1 IV Lonchocarpusblainii -1 IV Caseariasylvestris var. sylvestris -1 IV Psidiumguajava -1 IV Herbaceouslayer Chromolaenaodorata -1 V Erythroxylumhavanense -1 V Rivinahumilis -1 V Commelinaelegans -1 V Petiveriaalliacea -1 V Abutilon sp. - 1 V Pavoniatyphalea -1 V Trichiliahirta -1 IV Spermacocelaevis -1 IV Desmanthusvirgatus -1 IV Phylanodiflora -1 IV Turneraulmifolia -1 IV Rauvolfiatetraphylla -1 IV Desmodiumincanum -1 IV Adiantumtrapeciforme -1 IV Varroniaglobosa ssp. humilis -1 IV Oeceocladesmaculata -1 IV Lianas Gouanialupuloides -1 V Pisoniaaculeata -1 V Serjaniadiversifolia -1 V Turbinacorymbosa -1 V Trichostigmaoctandrum -1 IV Smilaxhavanense -1 IV Lasiasisdivaricata -1 IV Cissampelospareira -1 IV Chiococcaalba -1 IV Tournefortiahyrsutissima -1 IV Centrosema sp. - 1 IV Stigmaphyllumlineare -1 IV Stigmaphyllumsagreanum -1 IV

Scatteredspeciesinthecanopylayer:anacahuita,ayúa,yamagua,yagruma, Citharexylum sp.,jobo,sigua,royalpalm, pomarrosa( Zyzygium jambos ),jagua,cedar,tengue,corojo( Gastrococos crispa )andjocuma. Reyes O.J. Lazaroa 37 2016: 43-103 79

Table24. Characteristiccombinationofsiguawithguara (Ocotea coriacea with Cupania glabra ,6plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Ocoteacoriacea D2 V Cupaniaglabra var.glabra C1 V Ficuslaevigata C1 V Guapiraobtusata C1 V Plumeriaemarginata I1 V Amyriselemifera C1 IV Drypetesalba C1 IV Celtistrinervia I1 IV Dendropanaxarboreus C1 IV Abaremaglauca C1 III Zanthoxylumcubense C1 III Cordiagerascanthus C1 III Citharexylumcaudatum I1 III Shrublayer Hyperbaenapaucinervis -1 V Picramniapentandra -1 V Cojobaarborea -1 IV Caseariahirsuta -1 IV Eugeniaasperifolia -1 IV Herbaceouslayer Farameaoccidentalis -1 V Pharusglaber -1 IV Zamiaangustifolia -1 IV Lianas Chiococcaalba -1 V Philodendronlacerum -1 V Selenicereusgrandiflorus -1 V Smilaxhavanensis -1 V Lasiacisdivaricata -1 IV Pisoniaaculeata -1 III Smilaxlanceolata -1 III Vanilla sp. - 1 III Epiphytes Hohenbergiapenduliflora -1 V Trichocentrumundulatum -1 V Tillandsiafasciculata -1 V Tillandsiausneoides -1 IV Peperomiamagnoliaefolia -1 III Rhipsaliscassutha -1 III Pleurothallissertularioides -1 III Tillandsiapruinosa .1 III

Scatteredspeciesinthecanopylayer: carmín,roblemacho( Tabebuia hypoleuca ),agueditadulce,palodecaja,yaya, yaicuaje, Ficus perforata , guayabillo( Guettarda calyptrata ),yaití,royalpalm, combretaceae ,leviza( Licaria jamai - censis ), Pithecellobium sp.,guaguasí,canilladevenado( Citharexylum fruticosum ),cupey( Clusia rosea ),macagua (Pseudolmedia spuria ),yellowjucaro( Buchenavia (tetraphylla ) capitata ),dominguito( Chionanthus domingensis ), cuyá, Tabebuia sp.,carbonero,maboa( Cameraria latifolia ),chicharrón,ayúa,yagruma,ceiba , guásimaandcedar. Localities:CarsodeBaire. 80 Reyes O.J. Lazaroa 37 2016: 43-103

Table25. Characteristiccombinationofyamaguawithramoncow (Guarea guidonia with Dendropanax arboreus ,3plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Dendropanaxarboreus D3 V Guareaguidonia D2 V Cordiagerascanthus D1 V Ocoteacoriacea D1 V Zanthoxylummartinicense D1 V Cecropia peltata C1 V Roystonearegia C1 V Guazumaulmifolia C1 IV Cedrelacubensis C1 III Spondiasmombin C2 III Sideroxylonfoetidissimum subsp. foetidissimum C1 III Ficusmembranacea C1 III Pseudolmediaspuria C1 III Zuelaniaguidonia C1 III Burserasimaruba C1 III Ficussuffucans C1 III Oxandralanceolata I2 III Geoffroeainermis I1 III Trichiliahirta I1 III Caseariasylvestris var.sylvestris I1 III Drypetesalba I1 III Shrublayer Picramniapentandra -1 V Caseariahirsuta -1 V Trophisracemosa -1 V Cupaniaglabra -1 V Licariajamaicensis -1 IV Prunusmyrtifolia -1 IV Calophyllumantillanum -1 III Guettardacalyptrata -1 III Coffeaarabica -1 III Ocoteafloribunda -1 III Erythroxylumareolatum -1 III Herbaceouslayer Cupaniaamericana -1 V Pharusglaber -1 V Adiantumtrapeziforme -1 V Erythroxylumhavanense -1 IV Pavoniaspinifex -1 IV Oeceocladesmaculata -1 IV Oplismenussetarius -1 IV Chrysophyllumoliviforme -1 III Psychotria sp. - 1 III Lithacnepauciflora -1 III Allophylluscominia -1 III Celtistrinervia -1 III Farameaoccidentalis -1 III Anthuriumcubense -1 III Urerabaccifera -1 III Lianas Cissusverticillata -1 V Pithecocteniumechinatum -1 IV Reyes O.J. Lazaroa 37 2016: 43-103 81

Pisoniaaculeata -1 IV Gouanialupuloides -1 IV Abrusprecatorius .1 IV Trichostigmaoctandrum -1 III Hyperbaena sp. - 1 III Passiflorasuberosa -1 III Tournefortiaglabra -1 III Chiococcaalba -1 III Serjaniasubdentata -1 III Philodendron lacerum -1 III Epiphytes Tillandsiafasciculata -1 IV Tillandsiausneoides -1 III Campiloneurumphyllitidis -1 III

Scatteredspeciesinthecanopylayer: bucare( Erythryna poeppigiana ), Ficus laevigata ,dominguito,yaicuaje,toad (Ocotea leucoxylon ),penda,tabaco( Cordia sulcata ),bienvestido( Gliricidia sepium ),lechero( Tabernaemontana amblyocarpa ),comecará( Eugenia aeruginea ),siguaralla( Trichilia havanensis ),aceitunillo( Beilschmiedia pendula ), ayúadesierra,morurorojo( Cojoba arborea ),dagame, Thouinia sp.,cupey,guamácandelón, Eugenia sp.,tengue, mije( Eugenia floribunda ),yellowjucaro,cuajaní( Prunus occidentalis ),dairyandguairaje( Eugenia axillaris ).

Itoccurswheretheecologicalconditions mumnumberofsamplesisnonexistent(2 aremostfavorable.Itstopographicposition, plots).Thecanopylayerhastwosublayers,the protectedbynarrowvalleysbetweenthe upperoneisabout20mhigh,withemergents mogotes,orrockmassesintheupperpart of25m,sometimesthislayerreachesthelat - thereof,makesforlimitedexposuretothesun - terheight;thelowerlayerontheotherhandis lightandnaturalwindshield;thisreducesthe from8to10mhigh,occasionallybetween12 aircirculationanditseffectonevaporation- to15m;withadensecovert(100%).Inge- transpiration.Becausethissetoffactors,the neralthereare41treespeciespresent,the waterregimeisrelativelyfriendly. mostabundantare:jobo(D,AC3),yaya(I, Thesoilisbrownishsialític(Hernández& AC2-3),yagruma(C,AC2),guásima(D,AC al .,1994),reddish,fromshallowtomoderately 2),yamagua(D,AC2),ramoncow(C,AC2) deep,withlimestonerocksonthesurface, andmacagua(D,AC2).Therearealsocon - whichcoverbetween5and80%.TheLlayer stant,althoughnotabundant:sigua(D,AC1), hasbetween1.5and3cm,theFlayervaries guarano(C,AC1),guara(I,AC1),royalpalm fromtracesto0.2cm(withrootlets)andHis (P,AC1),jocuma(C,AC1),jagueymale missing.Inthisforest,whosecycleofnutrients (Ficus membranacea ,I,AC1),jubabán(I,AC dependsonthesoil,therateofwitheredleaves 1),ramonhorse(C,AC1),ayua(C,AC1), decompositionseemstobeveryquick(Herrera guaguasi(I,AC1),almacigo(I,AC1),yaba &Rodriguez,1988).Therefore,theedatopeis (I,AC1),raspalengua(I,AC1),baria(I,AC consideredfresh(2)andmoderatelyfertile(B), 1),ocuje(C,AC1)andcaimitillo(0,AC1). sothatthepotentialproductioncapacityiscon - Asspreadtreespeciestherecanbeobserved: sideredasregular(III).Thetypologyclassifica - morurorojo(I,AC1),cupey( Clusia rosea ,I, tionisconsideredB x-xxx 2,4cTh d,III,G.g.,D.a. AC1),dagame(I,AC1), Thouinia sp.(0,AC Distribution.Itispresentintheareaknown 1),leviza(0,AC1),guamácandelon(I,AC1), asCarsodeBaire. washingjaguey( Ficus suffucans ,I,AC1), Termsforforestmanagement.Thistypeof guairaje( Eugenia sp.,0,AC1),aguedita forestisrichintreespecies,soitmustbehan - sweet(I,AC1),saithearabic( Erythroxylum dledinordertoobtainmaximumdevelop - areolatum ,I,AC1),tengue(0,AC1),mije ment,freeingupthetopsofthevaluable (Eugenia floribunda ,0,AC1),bone(0,AC1), species.Subsequently,extractionswillbeper - yellowjucaro(C,AC1),cuajaní(C,AC1), formedselectively. almendrillo(I,AC1),dairy( Sapium jamaicense ,I,AC1)andguairaje( Eugenia axillaris ,0,AC1). Yaya with jobo ( Spondias mombin with Theshrublayerhasamediacovert(20- Oxandra lanceolata ) 30%)anditiscomposedalmostentirelyof treespecies.Thiscovertislowintheherba - Itconsistsofagalleryforestwhosecorre - ceouslayer(approx.10%)andnearhalfofits spondingtableisnotpresentedsincethemi ni - compositionistreespecies. 82 Reyes O.J. Lazaroa 37 2016: 43-103

Oecotopecharacteristics.Locatednearthe speciesobservedaremorurorojo,aguacatillo riverinthebottomoftheslopeandconse - (Alchornea latifolia ),yaicuaje( Matayba quentlyreceivesthecontributionoftherunoff oppositifolia ),dairy, Ocotea globosa ,marañon thereof.Besides,duetoitstopographicposi - decosta( Talauma orbiculata ),jagueymale tionitisaffectedbythegravitationalnight (Ficus membranacea )andbone. winds.Therefore,thewaterbalanceisfavo rable. Intheshrublayer,withamediacovertand Theheightisbetween190and200ma.s.l,the 12.2treespecies(averagesamples)thereare inclinationfrom8to35degreesandexposure asaggregate:polebox,cupey,casmagua betweeneastandnorth.Themeanannual (Wallenia laurifolia )andmanaca temperatureisaround24ºC.Themeanannual (Calyptronoma plumeriana ).Theother rainfallisbetween1400and1600mm,soitis speciesoftheshrubandherbaceouslayers consideredmoderatelyrainy.Therelative (covertmediatohighand13.5averagetree humidityvariesfrom80to85%,whereasthe species),fernsandlianasarefoundinTable evaporationvariesfrom1500and1700mm. 26. Forthatreason,itisconsideredatropical Oecotopecharacteristics.Itusuallyde velops climatewithdrywinter,subtype4cTh,with between500and800masl (locallyupto900 3-4(moderatelydry)drymonths. mabovesealevelonthesouthside),the ItislocatedonrocksoftheCobreGroup, slopesvarybetween30and40degreesand thesoilissialíticbrown,deep,withgravel theexposurearevaried.Themicroreliefis (20%);inadditionithas40to100%ofrocksat generallyflat.Thisforestoccursinevergreen thesurface.Thereforetheedatopeis considered forestfloor;climateistropicalwithdrywinter, moderatelywet(3)andfertile(C)andthe 4cThsubtype,3to4monthswithlittlerain potentialproductivecapacityasgood(II).The (moderatelydry).Therainfallvariesbetween layersofhumusarenotwelldeveloped,theL 1200and1800mmofrain,amoderatelyrainy, isbetween3.5and5cmthick,coveringabout withlargedifferencesintheirlength;thedry 80%ofthesurface,Fisalmostimperceptible, seasonisfromNovembertoAprilandrainy scatteredamongtherocksandgravel,whileH fromMaytoOctober(Montenegro1991a,b). islacking.Thistypeofwitheredleaves Theaverageannualtemperatureisrelatively decompositionisknownasquick. cool,fluctuatesbetween22and23ºCatthe Thetypologyclassificationisasfollows bottomandaround20ºCatthetop.Atthese Cxx-xxx 3,4cTh c-d ,II,O.l.,S.m. Distribution.Thestudywasrunatthemiddle altitudes(below800masl),intheevenings oftheTurquinoriver. andinmid-apriltolateOctober,relativeinso - Termsforforestmanagement.Duetoits lationvariesbetween20and30%,therestof smallsizethisforestshouldbepreservedfrom thesemonthsis30to40%.Inthemorning anyhumanactivity.Inthezonethereareareas rangesfrom40to60%(Montenegro,1991e). whereitshouldberestoredtomixedforests Thislowerstheeffectofdrought. withthesamespeciesascurrentlypresented. Itoccursonsoilsderivedfromrockthe CobreGroup,mostlyandesites,andtheseare mainlyferraliticbrown,lowtoveryshallow, Almendrillo with yamagua ( Prunus sometimeswithstonesonthesurface.The myrtifolia with Guarea guidonia ; Table 26) witheredleavesarewelldeveloped,theL layerisbetween2and7cm thick;Fvariesfrom Physiognomyandfloristiccomposition. 0.5to3cm,whereasHismissing ;nanodepre - Thisevergreenforest(mesophytic)hasa tionssometimesinhumusaccumulationsare canopylayerwith24speciesoftrees(average observedwithrootsandrootlets.Therefore,it samples),itisfrom12to25mhighandits isconsideredasfreshedatopes(2)andmode- covertisdense(90-100%).Inthegroupof ratelyfertile(B),sothattheirpotentialpro - constantspeciesthereareasmainspecies ductioncapacityisratedasregular(III).The yamaguaandramoncow;assubdominants: typologyclassificationisasfollowsB2, boniatillo( Cinnamomum elongatum ),toad 4cTh c-d ,III,P.m.,G.g. (Ocotea leucoxylon ),ayua,guarano,caimitil - Distribution.Thistyperepresentsthesub - lo,ramonhorse,almendrillo,dominguitoand mountainforestsoftheSierraMaestra,i.e. aceitunillo( Beilschmiedia pendula ).Inthe evergreenmesophyllousforests(mesophytic), groupofaccompanyingspeciesthereareas whichweredesignatedasmanacalesbyLeón subdominantspecies:cuajaní,royalpalm, (1946)andascanelonesbySamek(1974).At sweetpotato,laurel,macagua,palmaboba thetoptransitionsofthemountainrainforests (Prestoea acuminata var. montana ),sweet andbottomwithmesophiticsemi-deciduous aguedita,mije( Eugenia floribunda )andcopal forests.TheywerestudiedintheSierraof (Protium cubense ).Inturn,theaggregate GranPiedraandTurquinoriverbasin. Reyes O.J. Lazaroa 37 2016: 43-103 83

Table26. Characteristiccombinationofalmendrillowithyamagua (Prunus myrtifolia with Guarea guidonia ,4plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Guareaguidonia D2 V Dendropanaxarboreus D2 V Prunusmyrtifolia C1 V Ocotealeucoxylon C1 V Cinnamomumelongatum C1 V Zanthoxylummartinicense C1 V Cupaniaamericana C1 V Trophisracemosa C1 V Chionanthusdomingensis C1 V Beilschmiediapendula C1 V Chrysophyllumoliviforme I1 V Prunusoccidentalis C1 IV Roystonearegia C1 IV Ocoteafloribunda C1 IV Pseudolmediaspuria C1 IV Prestoeaacuminata var. montana I1 IV Caseariasylvestris var. sylvestris I1 IV Eugeniafloribunda I1 IV Protiumcubense I1 IV Cojobaarborea C1 III Alchornealatifolia C1 III Mataybaoppositifolia C1 III Sapiumjamaicense C1 III Ocoteaglobosa C1 III Talaumaorbiculata C1 III Ficusmembranacea C1 III Drypetesalba I1 III Shrublayer Allophylluscominia -1 III Gesneria sp. - 1 III Palicoureaalpina -1 III Clusiarosea -1 III Wallenialaurifolia -1 III Anthirearadiata -1 III Piper sp. -1 III Calyptronomaplumeriana -1 III Herbaceouslayer Oplismenussetarius -1 IV Oeceocladesmaculata -1 IV Desmodium sp. - 1 III Farameaoccidentalis -1 III Psychotria sp. -1 III Psychotriauliginosa -1 III Pavoniaspinifex -1 III Lithachnepauciflora -1 III Peperomia sp. - 1 III Ferns Campyloneurumphyllitidis -2 V Blechnumoccidentale -1 V Cyatheaparvula -1 III Adiantumtrapeciforme -1 III 84 Reyes O.J. Lazaroa 37 2016: 43-103

Lianas Vitistiliaefolia -1 V Pisoniaaculeata -1 V Gouanialupuloides -1 V Lygodiumvolubile -1 V Hyperbaenapaucinervis -1 IV Trichostigmaoctandrum -1 IV Davilarugosa -1 IV Smilaxlanceolata -1 IV Hippocrateavolubilis -1 III Cissampelospareira -1 III Passiflorasexflora -1 III Smilaxhavanensis -1 III Serjaniadiversifolia -1 III

Scatteredspeciesinthecanopylayer: bucare,camaguilla (Myrsine coriacea ),cuero( Guettarda monocarpa ),penda, Clusia sp., Colubrina sp., Cassia sp.,aguedita( Picramnia pentandra ),yareicillo( Myrcia fenzliana = Gomidesia lin - deniana ),pomarrosa,yaya, Ocotea globosa , jubilla( Sideroxylon jubilla ),robleblanco( Tabebuia angustata ),gua - guasí,jocuma, Thouinia sp.,yaicuaje,sarnillacimarrona( Casearia sylvestris var. myricoides ),cedar, yagrumahem - bra( Schefflera morotottoni ),purio( Oxandra laurifolia )andateje( Cordia collococca ).

Forestmanagement.Theseforestsarerich Oecotopecharacteristics.Ittakesplace intreespeciesandhavebeenveryimpo- between800and1120masl,theinclinationis verished,especiallyforgrowingcoffee,meats among30and45degrees,andexposuresare andpastures,sothefewremnantsareseverely mainlytothenorth. threatenedandmustbeprotected.Manyfor TheweatherstationinGranPiedrareports thistypeofforestoecotopesshouldbe thefollowingdata(Montenegro,1990):29ºC restoredtomixedforests,mainly:cedar,jubi- asabsolutemaximumtemperature,6ºCas lla,jocuma,yamagua,boniatillo,almendrillo, absoluteminimumtemperature,18.4ºCas cuajaní,sweetpotatolaurel,coastmarañon, averagetemperature,relativehumidityat ateje,purio,guarano,ayua,etc. 89.6%,meanannualrainfallof1664mm (rainy),139dayswithrainonaverage,sunshine average5to7hours/day.Prevailingwindsare Dominguito with palma boba ( Chionanthus thetrade´sfromthenortheastandnorth- domingensis with Prestoea acuminata var. northeast,whichinturnarealsothestrongest. montana ; Table 27) Theaveragenumberofdayswithhorizontal precipitation(fogandlowcloud)is238;of Physiognomyandfloristiccomposition. them162daysarewithdensefog(Montenegro, Thistypeofforestisamountainrainforest. 1990).Thesehorizontalprecipitations,accordin g Theheightanddensityofthecanopylayer BoytelYambú(1972),cometomoistenthesoil (averaging15.5speciesamongsamples)are andvegetation;whileSamek&Travieso presentedfairlyuniform;heightfluctuates (1968)exposedthattheycanrepresentup to between12and15m,whilecoverageisdense 50%ofrainfall.Therefore,theirimportance (90-100%).Inthegroupofconstantspecies, asmoistureprovidersiscritical,aswellas palmaboba,ramoncowanddominguitoare forlimitingitslossintheecosystem. mainspecies;whiletoad, canelón( Ocotea Bioclimatically,theclimateismoistTropical cuneata ), guárano, yaicuaje, aceitunilloand Mountain(7a).Vilamajó& al. (1989) yareicillo( Myrcia fenzlian )aresubdominants. considereditasoftermoxerictype(wet Inthegroupofaccompanyingspecies,cupey equatorial). (Clusia tetrastigma ),barril,pomarrosaand Soilisleachateredferralitic(sensu boniatilloaresubdominants,whileinthe Hernández& al .,1994)oryellowishbrown.It groupofaggregatedspeciesare,cuajaníand derivedfromrocksoftheCobreGroup,main - puriofangar( Guatteria neglecta ). lyfromandesites.Itisverypoor(Renda, Theshrubandherbaceouslayerswitha 1989;Renda& al .,1980,1981),moderately coversfrommediatohigh(40-70%)andhigh fertile(B)andfresh(2),soitspotentialpro - (50-90%)respectively,ferns,lianassynusiaes ductivecapacityisconsideredregular(III). andepiphytesarefoundinTable27. Ithasgoodinternalandsurfacedrainage.The Reyes O.J. Lazaroa 37 2016: 43-103 85

witheredleavesarewelldistributedonthe typologyclassificationisasfollowsB2,7a e, sur face;theLlayerreachesfrom3to5cm; III,Rm*,Ch.d.,P.a. FandHlayersaremixed, sometimesforming Distribution.ItcomesinthetopofSierra aweftthatcanreachupto10cmthick, ofGranPiedra,from800maslinthenorth sometimesitgetstobearealrootmat.The sideandlocally900maslinthesouth.

Table27. Characteristiccombinationofdominguitowithpalmaboba (Chionanthus domingensis with Prestoea acuminate ,9plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Chionanthusdomingensis D2 V Prestoeaacuminata var. montana D2 V Dendropanaxarboreus D2 V Ocotealeucoxylon D1 V Cupaniaamericana C1 V Mataybaoppositifolia C1 V Ocoteacuneata C1 V Beilschmiediapendula C1 V Myrciafenzliana (Gomidesialindeniana)I1V Clusiatetrastigma I2 IV Cyrillasilvae C1 IV Syzygiumjambos C1 IV Cinnamomumelongatum C1 IV Prunusoccidentalis C1 III Guatterianeglecta C1 III Shrublayer Gesneria viridiflora -1 V Lyonialatifolia subsp. calycosa -1 III Ilex sp. - 1 III Herbaceouslayer Callicarpaoblanceolata -1 V Psychotriagrandis -1 V Phaiustankervilliae -1 IV Palicoureaalpina -1 IV Hedyosmungrisebachii -1 IV Meriania leucantha var. nana -1 IV Oplismenussetarius -1 IV Columneacubensis -1 IV Peperomia sp.1 - 1 III Coccoloba sp. -1 III Peperomia sp.2 - 1 III Ferns Cyatheaparvula -1 V Blechnumoccidentale -1 V Cyatheaaspera -2 IV Lianas Philodendrum lacerum -1 III Epiphytes Guzmaniamonostachya -1 V Campiloneurum phyllitidis -1 III Diplazium sp. - 1 III Scatteredspeciesinthecanopylayer: morurorojo,sarnillacimarrona, Ficus sp.,palodecaja,aguacatillo,cletra (Clethra cubensis ),guayacanejo( Coccoloba diversifolia ),camaguilla( Myrsine coriacea ),brunelia( Brunellia como - cladifolia ),roblemacho( Tabebuia hypoleuca ),almendrillo,robleblanco( Tabebuia angustata ),marañon( Talauma sp.),penda, Ocotea globosa ,yagruma,mije( Eugenia floribunda ),cuero( Guettarda monocarpa ),macagua,aguedita dulce, Hieronyma pallida ,cupey( Clusia rosea ),achotillo( Sloanea curatellifolia ). 86 Reyes O.J. Lazaroa 37 2016: 43-103

Table28. Characteristiccombinationofyareicillowithcopey (Myrcia fenzliana (Gomidesia lindeniana )with Clusia tetrastigma ,8plots

Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Clusiatetrastigma D3 V Myrziafenzliana (Gomidesialindeniana)I2V Eugeniascaphophylla C1 V Mataybaoppositifolia C1 V Ocotealeucoxylon C1 V Coccolobadiversifolia C1 V Alchornealatifolia C1 V Clethracubensis I1 V Cyrillasilvae C1 IV Lyonialatifolia subsp. calycosa I1 IV Myrica cerifera I1 III Shrublayer Ilexmacfadyenii -1 V Ossaeagranulata -1 V Palicoureaalpina -1 V Gesneriaviridiflora -1 V Myrsinecoriacea -1 V Ilexrepanda -1 IV Citharexylum sp. - 1 IV Guettardavalenzuelana -1 III Coccolobawrightii -1 III Herbaceouslayer Sclerialithosperma -1 V Panicumglutinosum -1 V Psychotria sp. - 1 IV Chionanthusdomingensis -1 IV Prestoeaacuminata var. montana -1 IV Eupatorium sp. - 1 IV Phaiustankervilliae -1 IV Olyralatifolia -1 IV Ichnanthusmayarensis -1 IV Vernoniahieracioides -1 III Oplismenussetarius -1 III Cinnamomumelongatum -1 III Piper sp. - 1 III Chaptalia sp. - 1 III Ferns Cyatheaparvula -3 V Sticherusbifidus -1 IV Blechnum occidentale -1 IV Cyatheaaspera -- III Odontosoriascandens -1 III Lianas Smilaxhavanensis -1 V Smilaxlanceolata -1 IV Arthrostylidiummultispicatum -1 IV Epiphytes Isochiluslinearis -1 IV Tillandsiafasciculata -1 IV Guzmaniamonostachya -1 IV Dichaeahystricina -1 III Scatteredspeciesinthecanopylayer: agueditadulce, ramoncow, cuero, guárano,cedar,cupey,roblemacho,poma - rrosa,purioprieto( Guatteria blainii ),palodecaja,aceitunilloandcanelón. Reyes O.J. Lazaroa 37 2016: 43-103 87

Forestmanagement.Thisrainforestisof thissuccessionalstage,suchrootmatisthe inestimablehydrologicvalue,andalsoaspro - mainelementintherecirculationofnutrients tectorofaveryerodiblesoil.Ithasbeenhigh - inthisecosystem.Decompositionisveryslow lyaltered;onlyrelativelypreservedrelictsare sinceplantsthatprovide75%ormoreofthe left.Itmustbeprotectedfromanyhuman observedwitheredleaves[copey,yareicillo, interventionsinceitispartofaBiosphere yaicuje,clavellina,fern( Cyathea parvula )and Reserve(alsoProtectedAreaofManaged sometimes,tapacamino( Palicourea alpina ) Resources)anditisitsmain objectofconser - andaguacatillo]mainlyprovideasclerophyl - vation.Onlycontrolledobservationofitsvalues lousmaterial.Thetypologyclassificationisas andscientificresearchsh ouldbeallowed. followsB2,7a e,III,Rm**,M.f.,C.t. Distribution.ItlocatesinthetopofSierra delaGranPiedra,up1000masl. Yareicillo with copey [ Myrcia fenzliana Forestmanagement.Itmustbeprotected (Gomidesia lindeniana ) with Clusia fromanyhumaninterventionasitisanearly tetrastigma ; Table 28] community(homeostasisIandearlyfieraII) fromthemountainrainforest,typicalofthe Physiognomyandfloristiccomposition. place,andthereareonlyisolatedremnants. Thisforestisasuccessionalstage(homeosta - sisIandearlyfieraII)(sensuCapote& al ., 1988;Reyes&Acosta,2005)ofthemountain Pomarrosa with palma boba ( Syzygium rainforestintheGranPiedra.Structurally,it jambos with Prestoea acuminata var. canbeconsideredasashrubbyforestand/or montana ) atreescrub.Alowcanopylayerispresentin themostevolvedpart(fieraII),between7and Physiognomyandfloristiccomposition. 10mheightandwithaslightlysparsetodense Thissecondaryforest,consistingofagallery (80-100%)cover.Inthegroupofaccompa- forestorisolatedforestfragments(2plots),is nyingspecies,barrilandclavellina( Lyonia observedinSierradelaGranPiedra,about latifolia subsp. calycosa )aresubdominants, 1000maslinoecotopesofmountainrainfor - whileinthegroupofaggregatedspecies, est.Thecanopylayercovers100%ofthearea arraigán( Myrica cerifera )is.Intheshrub (densecoverage)andisfrom12to15mhigh. layer,withacoverfrommediatoveryhigh Pomarrosa(D,A-C5)isobservedasmain (40-100%)camaguillaissubdominantand speciesandcopey(I,A-C1)assubdominant. Citharexylum sp.,yaicuaje , yareicillo , vigueta Theshrublayerisvariable,between20and (Guettarda valenzuelana ) and uverillo 80%(mediumcoverage).Dominguito(A-C2) (Coccoloba wrightii ) areaggregated.Inthe andpalmaboba(A-C2)areabundant.The herbaceouslayer,withacoverofhightovery herbaceouslayerisdense,80to100%witha high(80-100%)aredominguito,palma diversecomposition. boba,yareicillo,toad,yaicuaje, Eugenia Ecologyanddistribution.Soilsareshallow scaphophylla , copey, boniatillo, guárano, andwithrelativelyplasticclay,whichfavors canelónandcamaguilla.Theotherspeciesof moistureretention;exposuresaretothenorth theshrubandherbaceouslayers,ferns,lianas andtheinclinationis15to20degrees. andepiphytesareintheTable28. Weatherconditionsareavoidedastheyare Oecotopecharacteristics.Thiscommunity describedindominguitowithpalmaboba.The wasstudiedinthehighestpartsofSierradela typologyclassificationisconsideredasB2-3, GranPiedra,above1000masl;inclinationis 7a e,II-III,S.j.,P.a. variable,usuallygreaterthan20d egrees,the Forestmanagement.Pomarrosagallery exposureisdiverseandthemicroreliefisre gu - forestsexceptionallyprotectthebanksof lar.Weatherconditionsareavoided,asthey streamsandrivers,soitsreplacementmustbe aredescribedinpalmabobawithdominguito. gradual,insmallareas,andonlycontinueitafter Soilisleachateredferralitic(Hernán dez& thetreatedareaisconsolidated.Dominguito, al., 1994),poorandacid,usuallydeep,m ode- morurorojo, Tabebuia spp.,yaicuaje,palma ratelyfertile(B)andfresh(2),soitspotential boba,guárano,ramóncowandaguacatillocan productivecapacity isregular(III).Withered beusedasalternativespecies.Plantingshould leavesarewelldeveloped.TheLlayervaries bedenseinordertoachieverapidcoverage.The between1.5and4cm,mostlybetween2and areaswhereforestsarestablemustbealsopar - 2.5cm,whiletheFlayerfluctuatesbetween1 tiallysubstituted,bygroup,withspeciestypical and2cm,exceptionallymore.TheHlayer ofmountainrainforest,including:puriofangar, formsarootmatembeddedinamatrixof yaicuaje,toad,aguacatecimarrón,barril,boni - humus.Itisgreaterthan10cminthemost atillo,cuajaní,aguacatillo,robledeolor,ayúa, advancedareasandlittlteinsmallerareas.At jubilla,macagua,etc. 88 Reyes O.J. Lazaroa 37 2016: 43-103

Purio fangar with marañón de la maestra and1400masl(lowmountains),butitwas (Guatteria neglecta with Magnolia cubensis ; mainlystudiedbetween1200and1300masl Table 29) Thepredominantslopesvarybetween20and 45degreesandexposuresarevaried,the Physiognomyandfloristiccomposition.This microreliefisalsodiverse.Theclimateistrop - isaverycomplextypeofforest,sincetheyare icalmoistmountain(7a).Rainfallisaround lateandmaturecommunitiesofthismountain 2000mm(rainyweather),withfogandlow rainforest.Itusuallyhasbetween39and53 clouds(usuallydense)mostdaysoftheyear, speciesperinventory(speciesaverageof48.6). resultinginhighatmospherichumidity.Main Initsmaturestage,itusuallyhasacanopy dataaresimilartothosefromtheoecotopeof layerbetween18and20mhigh,oftenupto25 palmabobawithdominguito. morshowingemerginguntilthatpoint. ItdevelopsmainlyintheCobreGroup Coverageisdense(typically100%).Inthe rocks,mainlytuffs,andesitesandgranites. groupofconstantspecies,marañondela Thesoilisleachateredferrallitic(Renda, maestra (Magnolia cubensis subsp. cubensis ), 1989),oftendeep,verypoorandacidic,mod - yaicuaje,dominguito,puriofangar,toad, eratelyfertile(B)andfresh(2),soitspotential lenguadevaca( Ixora ferrea ),aguacatecimar - productive capacity isregular(III).Ithasgood rón( Persea anomala ),camaguilla,cletra, internalandexternaldrainage.TheLlayeris cordobán( Miconia pteroclada )and Viburnum between1.5and3cmthick,whiletheFlayer villosum aremainspecies.Inthegroupof generallyrangesfrom2to2.5cm.Rootmat accompanyingspecies,boniatillo,copeicillo rangesfrom6to13cm,sometimesupto20 (Clusia grisebachiana ),cuajaní andramon cminnanodepressions.Thetipologyclassifi - cowaresubdominantspeciesinthisstratum; - cationisconsideredasB2,7a e,III,Rm* **, whilebarril,aguacatilloandrobledeolor G.n.,M.c. (Tabebuia brooksiana )areaggregated(Table Distribution.Thistypeofforestgrowsin 29).Theshrub(thicketfrommediumtohigh, thehighlandsofWesternSierraMaestra. 40-70%)andherbaceous(mediumtohigh,40- Forestmanagement.Itmustbeprotected 80%)layersarerichintreespecies. fromallhumanactivity,eventheirsuccession - Sometimes,pinodelamaestra( Pinus alstages,becauseduetoitsgeomorphological maestrensis )formssuccessionalstages(fiera II-homeostasisII)ofthisrainforest.Inthese position.Suchpositiongivesitahugehydro - placesthepinetreeformsaclosedsub-layerof logicalimportance,andallowsittoprotect about30to35mtall.Inaddition,thereisa watersandanextremelyerodiblesoil.Only secondsub-layerbetween10and16min controlledobservationofitsvaluesandscien - heightandof50to60%coverage,exclusively tificresearchshouldbeallowed.Thisshould composedbybroadleaffromrainforests, beprioritizedespeciallyinareasoutsidepro - wherethemostabundantspeciesaredomin - tectedareas. guito(themostabundant),cordobán( Miconia dodecandra ),cordobán( M. pteroclada ),ca- maguilla,aguacatillo,boniatillo,lenguade Ferns with barril ( Cyathea furfuracea and vacaandbrunelia. C. parvula with Cyrilla silvae ) InthesuccessionalstagehomeostasisII, withexclusiveparticipationofthebroad-leaved Initialforeststhatmakeupthefernswith speciesfromthemountainrainforest,theyform barrilarethesuccessionalstageshomeostasis adensecanopylayer(100%)withaheight IandfieraIIfromthemountainrainforestin between15and18m.Relictemergingsarealso theWesternSierraMaestra. Cyrilla silvae and seenofabout25m.Althoughitisyoung,an thetreeferns Cyathea furfuracea and C. ecosystembalanceisobservedinthisstage,as parvula dominatetheshrub-canopylayer. thereisstabilityinthespeciesofthisforesttype. Eachstageshowssomestructuralandfloristic Themostabundantspeciesaredominguito, differences,whichallowseparatestudies,in puriofangar,yaicuaje,barril,copeicillo,bo- twosubtypes: niatillo,aguacatecimarrónandotherscattered Fernswithbarril, species.Atthisstagethereisagooddevelop - Fernswithbarrilandyagrumamacho. mentofthehumuslayers.TheLlayeris3cm, Oecotopecharacteristics.Itwasstudied Fis1.5cmandHlayerconstitutesarootmat between1000and1400maslTheecological embeddedinamatrixofhumusofabout3cm. conditionsareshowninpuriofangarwith Oecotope characteristics.Itoccupiesthe marañóndelamaestra( Guatteria neglecta “floorofmountainrainforests”,between800 with Magnolia cubensis ). Reyes O.J. Lazaroa 37 2016: 43-103 89

Table29. Characteristiccombinationofpuriofangarwithmarañóndelamaestra (Guatteria neglecta with Magnolia cubensis ,9plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer

E3,2,1 Magnoliacubensis subsp. cubensis P1 V Guatterianeglecta D1 V Mataybaoppositifolia D1 V Chionanthusdomingensis D1 V Ocotealeucoxylon D1 V Ixoraferrea D1 V Perseaanomala D1 V Myrsinecoriacea I1 V Clethracubensis I1 V

E3,2 Miconiapteroclada I1 V Viburnumvillosum I1 V

E3,2,1 Dendropanaxarboreus C1 IV Cinnamommumelongatum C1 IV Prunusoccidentalis C1 IV

E3,1 Clusiagrisebachiana I1 IV E3,2,1- Cyrilla silvae C2 III E3,1 Alchornealatifolia C1 III E3,2 Tabebuiabrooksiana I1 III Shrublayer

E2,1 Merianialeucantha var. nana -1 V Graffenriedarufescens -1 V Palicoureaalpina -1 V Psychotriagrandis -1 V Eugenialaeteviridis -1 V Garryafadyena -1 III Weinmanniapinnata -1 III Clidemiaumbellata -1 III Herbaceouslayer

E1 Zeugitesamericana -1 V Arthrostylidiummultispicatum -1 V Hedyosmumgrisebachii -1 III Panicumglutinosum -1 III Peperomiamaculosa -2 III Begoniacubensis -1 III Ferns

E3- Cyatheaparvula I1 V E3,2 Cyatheafurfuracea I1 IV E2- Alsophyllamajor -1 V L- Odontosoriaaculeata -1 IV Ep- Hymenophyllum sp. - 1 IV Polybotriaosmundacea -1 III Campiloneurumangustifolium -1 III Polypodiumloriceum -1 III Epiphytes Guzmania sp. - 1 V Guzmaniamonostachya -1 IV Tillandsiafasciculata -1 IV Isochiloslinearis -1 IV

Scatteredspeciesinthecanopylayer: Ocotea spathulata , macaguey(toad), yagruma,ayúa,macagua,yareicillo, Sapium erythrospermum ,almendrillo,guárano,jubilla,cordobán( Miconia dodecandra ),pinodelamaestra,palma boba,brunelia, Pimenta sp.,ramonhorse,achotillo( Hieronyma nipensis ), Manilkara valenzuelana ,bone,jiquí( Pera bumeliifolia ), Ditta myricoides , Coccoloba sp.andvigueta. 90 Reyes O.J. Lazaroa 37 2016: 43-103

Ferns with barril ( Cyathea furfuracea and lobata ),rompezaraguey( Chromolaena odorata ), C. parvula with Cyrilla silvae ; Table 30) etc,begintodisappear.Inthegroupofcon - stantspecies, barril, Cyathea parvula and C. Physiognomyandfloristiccomposition.As furfuracea aremainspecies.Lenguadevaca, stageofhomeostasisIinoecotopesofthis camaguilla,copey,cletraandcordobán mountainrainforest,barrilandtreeferns (Miconia dodecandra )aretreesubdominantin Cyathea parvula and C. furfuracea dominate shrubandherbaceouslayers. Aguacatillo,bru - theshrubcanopylayer,withaheightbetween nelia,toad,marañondelamaestraandyaicua - 5and8m;occasionallyreachingupto12m. Theyusuallyclosethecanopy,with100% jeareagregated .Otherspeciesareshownin coverageandmore(superdense).Thiscreates Table30. umbrophylleconditionsandtheoriginalforest Humusdiversificationisclearlyobserved speciesenterwithinthem.Theyreachthis atthisstage;anLlayerof1.5to3cmispre - layerorarealreadyinthebelowones. sented,aswellasanFlayerof0.5to1.5cm. Likewise,thepioneerspecies:cucaracha TherearebigdifferencesintheHlayer,from (Pteridium aquilinum var. arachnoideum ), tracesuptoawell-developedrootmat10cm Dicranopteris pectinata ,malvablanca( Urena thick.

Table30. Characteristiccombinationoffernswithbarril (Cyathea parvula and C. furfuracea with Cyrilla silvae ,7plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Cyrillasilvae D3 V Cyatheaparvula D3 V Cyatheafurfuracea D2 V Shrublayer Ixoraferrea -1 V Myrsinecoriacea -1 V Chromolaena sp. - 1 V Palicoureaalpina -1 V Shrubsandherbaceouslayers Alchornea latifolia -1 III Brunelliacomocladifolia -1 III Ocotealeucoxylon -1 III Magnoliacubensis subsp. cubensis -1 III Ossaeagranulata -1 III Mataybeoppositifolia -1 III Viburnumvillosum -1 III Herbaceouslayer Clidemiaumbellata -1 V Clusiagrisebachiana -1 IV Clethracubensis -1 IV Miconia dodecandra -1 IV Panicumglutinosum -1 IV Coccocypselumlanceolatum -1 IV Zeugitesamericana -1 III Sclerialithosperma -1 III Ferns Hypolepis nigrescens -1 IV Sticherusbifidus -1 III Dicranopterispectinata -1 III Epiphytes Guzmaniamonostachya -1 III

Scatteredspeciesinthecanopylayer: robledeolor,cordobán( Miconia pteroclada ), Eugenia laeteviridis ,jubilla. Reyes O.J. Lazaroa 37 2016: 43-103 91

Ferns with barril and yagruma macho groupofconstantspeciesfromthetreeand (Cyathea furfuracea and C. parvula with shrublayers.Brunelia,aguacatillo,camaguilla Cyrilla silvae and Schefflera morotottoni ; aresubdominant,aswellaslenguadevaca Table 31) andtoadintheshrublayer.Inthegroupof accompanyingspecies,copeicilloissubdominant Asitisknown,thefieraIIphasebegins andyagrumamacho( Schefflera morotottoni ), whentreespeciessurpassthehomeostasisIstra - cletra,cordobán( Miconia pteroclada ), tum,andcompetitivelyinteractamongthem. marañondelamaestra and Eugenia laete - Theyformacanopylayerof8to12m, viridis areaggregatedspecies. withisolatedindividualsof15mandcoverage Theshrubandherbaceouslayershavea fromsparsetodense(50-100%).barril,cor - highdensity,60to90%and50to70%, dobán( Miconia dodecandra )andthe Cyathea respectively,withvarioustreesprouts.Other furfuracea and C. parvula fernsareinthe speciesarefoundinTable31.

Table31. Fernswithbarrilandyagrumamacho (Cyathea furfuracea and C. parvula with Cyrilla silvae and Schefflera morotottoni ,7plots)

Species Vertical Abundance- PresenceDegree structure Coverage Canopyandshrublayers Cyrilla silvae D3 V Miconia dodecandra D2 V Cyatheafurfuracea D2 V Cyatheaparvula D2 V Brunelliacomocladifolia C1 V Alchornealatifolia C1 V Myrsinecoriacea C1 V Clethracubensis I1 III Miconiapteroclada I1 III Shrublayer Palicoureaalpina -2 V Ixoraferrea -1 V Clidemiaumbellata -1 V Ocotealeucoxylon -1 V Clusiagrisebachiana -1 IV Chromolaena sp. -1 IV Magnoliacubensis subsp. cubensis -1 III Herbaceouslayer Panicumglutinosum -1 V Coccocypselumherbaceum -1 IV Eugenialaeteviridis -1 III Sclerialithosperma -1 III Ferns Hypolepisnigrescens -1 V Sticherusbifidus -1 III Dicranopteris pectinata -1 III Epiphytes Guzmaniamonostachya -1 III Differentialcombinationfromfernswithbarrilandyagrumamacho Arthrostylidiummultispicatum -1 .V Passiflorasexflora -1 V Blechnumfragile -1 V Schefflera morotottoni C1 III Callicarpaferruginea -1 III Oplismenussetarius -1 III Alsophyllamajor 1 III Odontosoriajenmanii -1 III Elaphoglossumchartaceum -1 III

Scatteredspeciesinthecanopylayer:yaicuaje,canelón,robledeolor,oreganillo( Weinmannia pinnata ),dominguito andramoncow. 92 Reyes O.J. Lazaroa 37 2016: 43-103

Barril with marañon de la maestra ( Cyrilla low,generallyrangingbetween1.44and5.02 silvae with Magnolia cubensis ; Table 32) cmol(+).kg -1 .CIC(Tvalue)isfrom2.5to 18.75cmol(+).kg -1 .Thisshowsthelow Physiognomyandfloristiccomposition. degreeofsaturationandthatthecomplexhas Aszonalformationitisacloudforest(Samek, littlecationduetointenseweatheringprocess. 1975;Reyes,2006,2011-2012),alsoknownas Therefore,thisedatopeshouldberegardedas freshforest(León,1946;Samek,1974)and littlefertile(A),fresh(2)andwithapotential elfinforest(Borhidi,1996).Inthisforest,the productivecapacitybad(IV). canopylayerwith10to15minheightis Adistinctivefeatureofthisforestisits dense(80-100%).Barrilandmarañondela well-definedhumuslayers(sensuHerrera& maestraaremainspeciesandthemostabun - Rodríguez,1988).TheLlayerreachesabout2 dantinthegroupofconstantspecies.Cletrais cm,occasionallymoreorless;Flayerfluctu - subdominantinthegroupofaccompanying atesbetween1and2cm,andHlayerisawell- species,while Henriettea ekmanii isaggregated . developedrootmatembeddedinamatrixof Otherspeciesfromtheshrub(mediatohigh humus,wherethefinerootsandrootletsare, coverage,40-70%)andherbaceous(high, andinwhichthenutrientcycleofthiscon - 50-80%)layersarefoundinTable32. spicuousecosystemoccurs(Reyes& al ., Oecotopecharacteristics.Itusuallyoccurs 2011).Thismeans,thatthispoor,acidicsoil,is insteeptopographicpositions,mostly onlyusedforplantfixation(thickroots)and between40and45degrees(Reyes&Acosta, virtuallydoesnottakepartintheirenergy 2005)andwithvariousexposures.Intherange source. ofaltitudinalvariationofthistypeofforest, Thetipologyclassficationsisconsideredas theaveragetemperaturesat1500ma.s.lvary A2,7b e,IV,Rm**,C.s.,M.c. between15.2ºCinthenorthslopeand15.5ºC Distribution.Thiscloudforestdevelopsin onthesouthside.Itcanalsoreacharound13 thehighestpartsofWesternSierraMaestra, ºCat1972masl(Montenegro,1991b). fromabout1500mabovesealeveltothetop Althoughtherearenotdirectmeasurementsin ofPicoRealdelTurquino.Itsgreatestextentis theareaoccupiedbythisforest,itfallsaround observedinthemassifsofTurquinoandLa 2000mmofrain.Thewettestperiodisfrom MaestricadelosLibertadores–LaBayamesa. aprilormaytooctoberandthelessrainype- Inotherplaces,thereareonlyisolatedfrag - riodisfromnovembertomarchorapril ments. (Trusov& al .,1983).Themeanannualrela - Forestmanagement.Becauseoftheecolo - tivehumidityisbetween85and90% gyconditions(Reyes&Fornaris,2011),the (Montenegro,1991c).Orographiccloudsor vegetationgrowthisextremelyslowandtrees mistsoccuralmosteverydayoftheyear,and havesmallthicketandsize.Duetoitstopo - alongwiththerain,theybringlotsofmoisture graphicandgeomorphologicposition,the intotheforest(eveninthedryseason),thecli - highestvalueofthistypeofforestisthepro - mateishighmountaintropical(7b). tectionofsoilandwater.Thatiswhy;itmust Fromthegeologicalpointofviewitoccu - bepreservedagainstallformsofhumaninter - piesrocksfromtheCobreGroup,tuffs,tuffa - vention.AnareafromhighLaBayamesa ceoussandstones,etc.(Méndez& al .,1994). (1572masl)wasstudied.Thisareawas Thepredominantsoilisleachedyellowferral - deforestedin1962duetothetriangulationof litic,whichdominatesabove1500masl SierraMaestra.Fortyfiveyearslater, AccordingtoRenda& al. (1981b)andRenda regenerationhavenotexceedthreemeters,so (1989),thepHintheupperhorizonsisfrom itisconsideredthatitsrecoverytomature acidtoveryacidicanditreachesvaluesof forestcanlastfrom150to200years;eventhe 4.45to5.3inwater,and3.5to4.15inClK. restorationoftherootmat(basisforits Theassimilablenitrogenvariesfrom0.80to nutrientcycling)isexceptionallyslow.Its 11.08mg/100gandtendstosharplydecrease protectingvaluewidelyexceedstheoneitmay indepth.Phosphorusisabsentorintraces; haveaswoodproducer;thereforeonly assimilablepotassiumfluctuatesbetween3.0 controlledobservationandscientificresearch and10mg/100g.TheCCB(Svalue)isvery shouldbeallowed. Reyes O.J. Lazaroa 37 2016: 43-103 93

Table32. Characteristiccombinationofbarrilwithmarañondelamaestra (Cyrilla silvae with Magnolia cubensis ,6plots) Species Vertical Abundance- PresenceDegree structure Coverage Canopylayer Cyrilla silvae D3 V Magnoliacubensis subsp. cubensis D2 V Clusiagrisebachiana C1 V Dittamyricoides C1 V Ixoraferrea C1 V Chionanthusdomingensis C1 V Weinmanniapinnata C1 V Brunelliacomocladifolia I1 V Myrsinecoriacea I1 V Clethracubensis I1 IV Henrietteaekmanii I1 III Shrublayer Graffenriedarufescens -3 V Purdiaeastenopetala var. stenopetala -1 V Palicoureaalpina -1 V Ilexmacfadyenii -1 V Viburnumvillosum -1 V Lyoniaelliptica -1 V Eugenialaeteviridis -1 V Guapiraobtusata -1 IV Herbaceouslayer Hedyosmumgrisebachii -1 V Lisianthiusglandulosus -1 V Sclerialithosperma -1 V Zeugitesamericana -1 V Ocoteaspathulata -1 IV Ilex sp. - 1 IV Callicarpaferruginea -1 IV Sapiumerythrospermum -1 III Phaiustankervilliae -1 III Peperomiatenella -1 III Ferns Alsophyllamajor -1 V Cyatheaparvula -1 V Odontosoria scandens -1 V Sticherusbifidus -1 V Trichomanesrobustum -1 V Cyatheafurfuracea I1 IV Elaphoglossumchartaceum -1 IV Campiloneurumphyllitidis -1 III Politrichum sp. - 1 III Polybotriaosmundacea -1 III Epiphytes Isochiluslinearis -1 V Catopsis cf. floribunda -1 IV

Scatteredspeciesinthecanopylayer: Guatteria moralesii ,macurije,cordobán( Miconia dodecandra ),pinodelamae - straand Wallenia sp. Localities:highestaltitudesofSierraMaestra. 94 Reyes O.J. Lazaroa 37 2016: 43-103

Conclusions masl,andforpoornessthoseclosedtomarine influence.It’soutstandingthat,theprimary Thirtyfivebroadleafforesttypesand/or montanetypesarelianaspoorestandepipytes subtypeswerestudied.Theforesttypesand/ andfernsarerichestthanthetypesatlowalti - orsubtypesmoreabundantarethosebelon- tude.Foresttypesabove800maslandman - gingtothesemideciduousmicrophyllforest grovesareprotectionforests. (12),followedbythemangrovesandmoun - Thetropicalclimatewith5to6months little tainrainforestwithanequalamount(7). rainy(4bTh)isthemostprevalentinSierra The27foresttypesand/orsubtypeswith - Maestra. outmarineinfluence,withcoastalandlow altitude,63%,haveapotentialproductive capacity(PPC)goingfrombad(44.4%)tovery Acknowledgments bad(18.5%),becausetheerosiondecreasethe effectivedeepandincreasedthestonyinthe Mostofthesamplesweretakenwithinthe soilhorizon.TheregularPPC(III)isprevail - frameworkoftwoprojects:“Biologicaldiver - inginthemittelandmountainzonebecauseof sityofSierraMaestraandNipeSaguaBaracoa thegreatprofundityofsoil.Inthehighest mountainranges”and“Environmentaleduca - foresttypethePPCdecreaseagainowingto tionalinitiativestoencouragesustainable thepoorestTandSvalue. localdevelopmentinprotectedareasofrain - Initscharacteristiccombinationthe pri - forestsoftheEasternCubanRegion”. maryforeststypestherearerichestintreespecies TranslationtoEnglishwasmadebyHaydee (18.4)thanthesecondary(8.4); outstandingfor NoemiVidalCarrillo.Wehighlyvalueyour itsrichnessthepresentsfrom500to800 support.

References

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Appendix1.Scientificnames(genus,specieandauthor)andcommonnamesfromspeciesused inthetables1-32.

Scientificnames Commonnames Abaremaglauca (Urb.)Barneby&J.W.Grimes abey Abrusprecatorius L. peonía Abutilon sp. - Acaciamacracantha H.&B.ex Willd. guatapaná Acrostichumdanaefolium Lang. &Fisch acróstico Adeliaricinella L. jíablanca Adiantumtrapeziforme L. culantrillodemonte Agaveunderwoodii Trelease maguey Agave sp. maguey Alchornealatifolia Sw. aguacatillo Allophylluscominia (L.)Sw. palodecaja Alsophylla major Caluff&Shelton - Alternantheramaritima (Mart.) Standl. - Alvaradoaamorphoides Lieb. aromablanca Alvaradoaarborescens Griseb. carmín Amyriselemifera L. cuabilla,.cuaba Anthirearadiata (Griseb.)Urb. vera Anthuriumcubense Engler anturio Aristolochialindeniana Duch. - Arthrostylidiumangustifolium Nash tibisí Arthrostylidiummultispicatum Pilger tibisí Auerodendroncubensis Britt.& Wils. cocuyodecosta Avicenniagerminans (L.)L. blackmangrove Bacopamonnieri (L.)Pennullvar. monnieri verdolagadecosta Begoniacubensis Hassk. begonia Beilschmiediapendula (Sw.)Benth.&Hook. aceitunillo Bidenspilosa L. romerillo Blechnumfragile (Liebm.)Morton&Lellinger - Blechnumoccidentale L. - Bothriochloapertusa (L.) A.Camus camagueyana Bourreriasetoso-hispida O.E.Schulz - Bourreriavirgata (Sw.)G.Don cafecillo Bourreria sp. - Brunelliacomocladifolia Bonpl. brunelia Burserasimaruba (L.)Sargent. almácigo Caesalpiniavesicaria L. brasil Callicarpaoblanceolata Urb. filigrana Calophyllumantillanum Britt. ocuje Calyptronomaplumeriana (Martius)Lourtier manaca Campiloneurumangustifolium (Sw.)Fée - Campyloneurumphyllitidis (L.)C.Presl pasadenegro Canellawinterana (L.)Gaertn. cúrbana Cappariscynophallophora L. aceitunillo Capparisflexuosa L. pichadeperro Capparisgrisebachii Eichl. olivo Capsicumfrutescens L. ajígaguao Carpodipteracubensis Griseb. majaguilla Caseariahirsuta Sw. raspalengua Caseariasylvestris Sw.var.myricoides sarnilla cimarrona Caseariasylvestris Sw.var.sylvestris agueditadulce Castelaspinosa Cronq. abrojodecosta Catopsis cf. floribunda L.B.Sm. curujey Cecropia peltata L. yagruma Cedrelacubensis Bisse redcedar Cedrelaodorata L. cedar 98 Reyes O.J. Lazaroa 37 2016: 43-103

Celtistrinervia Lam. aguedita Cenchrusciliaris L. guizazo Centrosemavirginianum (L.)Benth. cricadenegra Cinnamomumelongatum (Nees)Kosterm. boniatillo Cissampelospareira L. bejucoprieto Cissustrifoliata L. bejucouví Cissusverticillata (L.)D.H.Nicolson&C.Jarvis bejucouví Citharexylumcaudatum L. penda Citharexylum sp. - Clethracubensis A.Rich. cletra Clidemiaumbellata (Mill.)L.O. Wms. cordobán Clusiarosea Jacq. cupey Clusiatetrastigma Vesque cupey Coccocypselumlanceolatum Aubl - Coccolobadiversifolia Jacq. guayacanejo Coccolobauvifera Jacq. uvacaleta Coccolobawrightii Lindau uverillo Coccoloba sp. - Coccothrinaxelegans Borhidi&Mz. yuraguana Coccothrinaxfragans Burret yuraguana Coccothrinaxgundlachii León yuraguana Coffeaarabica L. cafeto Cojobaarborea (L.)Britt.&Rose morurorojo Colubrinaelliptica (Sw.) Brizicki&Stearn. carbonero Columneacubensis (Urb.)Britt. - Commelinaelegans K. canutillo1 Commelinaerecta L. canutillo Commicarpusscandens (L.)Standl. bejucodepurgación Comocladiadentata Jacq. guao Conocarpuserectus L. yana Corchorus sp. - Cordiaalba Jacq. uvita Cordiacollococca L. ateje Cordiagerascanthus L. baría Crossopetalum sp. - Crotonlucidus L. cuabadeingenio Cupaniaamericana L. guárano Cupaniaglabra L. guara Cupaniaglabra Sw.var. glabra guara Cyatheaaspera (L.)Sw. - Cyatheafurfuracea Baker helecho Cyatheaparvula (Jenm.)Domin helecho Cynanchum sp.1 - Cynanchum sp.2 - Cyperusalternifolius L. paraguita Cyperusgiganteus Vahl - Cyrillasilvae Berazaín barril Cyrtopodiumpunctatum (L.)Ldl. cañuela Chaptalia sp. - Chiococcaalba (L.)Hitchc. bejucodeberraco. Chionanthusdomingensis Lam. dominguito Chromolaenaodorata (L.) King&Robins. rompezaraguey Chromolaena sp. - Chrysophylumoliviforme L. caimitillo Dalechampiascandens L. - Davilarugosa Poit. bejucocolorado Desmanthusvirgatus (L.) Willd. adormidera Desmodiumincanum DC. amorseco Reyes O.J. Lazaroa 37 2016: 43-103 99

Desmodium sp. - Dichaeahystricina Rchb. - Diclipteravahliana Nees gallitos Dicranopterispectinata (Willd.) Und. patadegallina Diospyrusgrisebachii (Hiern.) Standl. blackebony Diplazium sp. - Distictisrhynchocarpa Urb. bejucologarte Dittamyricoides Griseb. - Drypetesalba Poit. bone Drypetesmucronata Griseb. chicharrón Drypetes sp. - Echitesumbellata Jacq. huevodegallo Elaphoglossumchartaceum (BakerexJenm.)C.Christ. - Eleocharis elegans (K.)R.&S. junco Eleocharismutata (L.)R.&S. junco Encyclia sp. - Epidendrumnocturnum Jacq. flordesanpedro Epidendrumpygmaeum Hook. - Erithalisfruticosa L. cuabaprieta Erythroxylumareolatum L. arabocarbonero Erythroxylumhavanense Jacq. jibá Erythroxylumrotundifolium Lunan yarúadecosta Eugeniaasperifolia Berg. guairaje Eugeniaaxillaris (Sw.) Willd. guairaje Eugeniafloribunda West. mije Eugeniamaleolens Poir. guairaje Eugeniascaphophylla Wr. - Eugenia sp. guairaje Eupatorium sp. - Euphorbiaheterophylla L. lechosa Exostemacaribaeum (Jacq.)R. &S. liriosantana Exotheapaniculata Radlk. yaicuaje Farameaoccidentalis (L.) A.Rich. - Ficuslaevigata Vahl jaguey Ficusmembranacea C. Wr. jagueymacho Ficussuffucans Griseb. jaguey Galactia sp. - Garryafadyena Hook. - Geoffroeainermis Wr. yaba Gesneriaheterochroa Urb. - Gesneriaviridiflora (Dcne.)Kuntze. - Gesneria sp. - Gouanialupuloides (L.)Urb. jaboncillo Graffenriedarufescens Britt.& Wils. - Grisebachianthusplucheoides (Griseb.)King&Robins. - Guapiraobtusata (Jacq.)Little macaguey(toad) Guareaguidonia (L.)Sleumer yamagua Guatterianeglecta (Griseb.)P. Wils. puriofangar Guazumaulmifolia Lam. guásima Guettardacalyptrata A.Rich. guayabillo Guettardaelliptica Sw. ciguilla Guettardavalenzuelana A. Rich. vigueta Guettarda sp. - Guzmaniamonostachya (L.)RusbyexMez curujey Guzmania sp. curujey Gymnantheslucida Sw. yaití Harrisiaeriophora (Pfeiff.)Britt. jijira Hedyosmungrisebachii Solms - 100 Reyes O.J. Lazaroa 37 2016: 43-103

Helicteressemitriloba Bert. tapaculo Helicteres sp. - Heliotropiumindicum L. alacrancillo Henrietteaekmanii (Urb.) Alain - Hibiscusbrasiliensis L. paloperegrino Hippocrateavolubilis L. bejucodecastaña Hymenophyllum sp. - Hyperbaenapaucinervis Urb. pichajutía Hyperbaena sp. - Hypolepisnigrescens Hook. - Ichnanthusmayarensis (Wr.)Hitchc. - Ilexmacfadyenii (Walp.) Rehder acebocubano Ilexrepanda Griseb. viguetanaranjo Ilex sp. - Ipomoeaalba L. florde Y Ipomoea sp. - Isidoreapolyneura (Urb.) Aiello. - Isochiluslinearis (Jacq.)R.Br. - Ixoraferrea (Jacq.)Benth. lenguadevaca Jatrophagossipifolia L. frailecillo Lagunculariaracemosa (L.) Gaertn. whitemangrove Lantanacamara L. filigrana Lantanamontevidensis (Spreng.)Briq. filigranacimarrona Lantanareticulata Pers. oréganocimarrón Lasiacisdivaricata (L.)Hitchc. tibisíchico Leucaenaleucocephala (Lam.)De Wit ipilipil Licariajamaicensis (Nees)Kost. leviza Lisianthiusglandulosus A.Rich. - Lithacnepauciflora (Sw.) Beauv. pitoenano Lonchocarpusblainii Wr. guamáhediondo Lonchocarpusdomingensis Urb. guamá Ludwigiaerecta (L.)H.Hara - Lygodiumvolubile Sw. - Lyoniaelliptica (Small) Alain clavellina Lyonialatifolia (A.Rich.) Griseb.subsp. calycosa clavellina Lysylomasabicu Benth. sabicú Magnoliacubensis Urb.subsp. cubensis marañóndelamaestra Malpighiaapiculata Urb. - Malpighiacnide Spreng. palobronco Malpighiasuberosa Small. palobronco Malvastrumcorchorifolium (Desv.) Britt. - Mataybaoppositifolia (A.Rich.) Britt. yaicuaje Maytenusbuxifolia (A.Rich.)Griseb. carnedevaca Melicoccusbijugatus Jacq. mamoncillo Melochianodiflora Sw. malvacolorada Merianialeucantha Sw.var. nana Triana - Merreniabisecta (Jacq.)Hallf. aguinaldodealmendra Miconia dodecandra (Desv.)Cogn. cordobán Miconiapteroclada Urb. cordobán Mikaniamicrantha K. guaco Morindaroyoc L. piñipiñi Myrciafenzliana O.Berg.(=Gomidesialindeniana Berg.) yareicillo Myrica cerifera L. arraigán Myrsinecoriacea (Sw.)R.Br.exRoem. camaguilla Ocoteacuneata (Griseb.)Urb. canelón Ocoteafloribunda (Sw.)Mez boniatolaurel Ocoteaglobosa Schlecht. & Cham. - Ocotealeucoxylon (Sw.)Laness. toad Reyes O.J. Lazaroa 37 2016: 43-103 101

Octomeriatridentata Lindl. - Odontosoriajenmanii Maxon - Odontosoriascandens (Desv.)C.Christ. tembladera Oeceocladesmaculata (Ldl.)Ldl. - Olyralatifolia L. tibisí Oplismenussetarius (Lam.)R.&S. cañamazo Oploniatetrasticha (Wr.exGriseb.) Stearn. nometoques Ossaeagranulata Urb. - Oxandralanceolata (Sw.)Baill. yaya Palicoureaalpina (Sw.)DC. tapacamino Panicumglutinosum Sw. pegapega Panicum sp. - Paspalumbreve Chase cañamazo Passiflorafoetida L. pasionariahedionda Passiflorapenduliflora Bert. bejucomanteca Passiflorasexflora Juss. pasionaria decerca Passiflorasuberosa L. bejucomantequilla Pavoniafructicosa (Mill.)Fawcett & Rendle tábano Pavoniaspinifex (L.)Cav. malaguilladecosta Peperomiamaculosa (L.)Hook. - Peperomiamagnoliaefolia (Jacq.) A.Dietr. - Peperomiarotundifolia (L.)K. lenguadelasmujeres Peperomiatenella A.Dietr. - Peperomia sp. - Peperomia sp.1 - Peperomia sp.2 - Perseaanomala Britt.& Wils. aguacatecimarrón Petiveriaalliacea L. anamú Phaiustankervilliae (Banks)Blume - Pharusglaber K. pelode perro Philodendronlacerum (Jacq.) Schott bejucolombricero Phylanodiflora (L.)Greene oroazul Phyllanthusepiphyllanthus L. panetela Phyllostylonbrasiliensis Cap. jatía Picramniapentandra Sw. aguedita Picrodendronbaccatum Kruge&Urb. aceituna Pictetiamucronata (Griseb.)Beyra&Lavin - Pilosocereusbrooksianus (Vamp.)Bol.&Row. jijira Pinusmaestrensis Bisse pinodelamaestra Piper sp. - Piscidiapiscipula (L.)Sargent. guamácandelón Pisoniaaculeata L. zarza Pithecocteniumechinatum (Aubl.)Schum. huevodetoro Platygynadentata Alain - Pleurothallisgelida Ldl. flordellantén Pleurothallissertularioides (Sw.)Spreng. - Plumbagoscandens L. malacara Plumeriaemarginata Griseb. lirioamarillo,suchel Poeppigiaprocera Presl. tengue Politrichum sp. - Polybotriaosmundacea H.&B.ex Willd. - Polypodiumaureum L. - Polypodiumloriceum L. - Polypodiumpolypodioides (L.) A.S.Hith. - Prestoeaacuminata (Willd.)H.E.Moorevar. montana (Graham) An. palmaboba Hend. & Galeano Protiumcubense (Rose)Urb. copal Prunusmyrtifolia (L.)Urb. almendrillo 102 Reyes O.J. Lazaroa 37 2016: 43-103

Prunusoccidentalis Sw. cuajaní Pseudocarpidiumavicennioides (A.Rich.)Millsp. chicharrón Pseudolmediaspuria Griseb. macagua Psidiumguajava L. guayaba Psychotria grandis Sw. tapacamino Psychotriauliginosa Sw. tapacamino Psychotria sp. - Purdiaeastenopetala Griseb.var. stenopetala - Randiaaculeata L. agalladecosta Randiaspinifex (R.&S.)Standl. espuela Rauvolfiatetraphylla L. alelí Ravenialeonis M. Vict. arraiján Rhipsaliscassutha Gaertn. disciplinilla Rhizophoramangle L. redmangrove Rivinahumilis L. coralitos Roystonearegia (K.)O.F.Cook. royalpalm Salacianipensis Britt. - Samaneasaman (Jacq.)Merr. algarrobo Sapiumerythrospermum Muell. Arg. - Sapiumjamaicense Sw. dairy Saviabahamensis Britt. hicaquillo Schefflera morotottoni (Aubl.)Mag.,Stey.&Frodin yagrumamacho Scirpusolneyi A.Gray juncodetresfilos Sclerialithosperma (L.)Sw. cortadera Selenicerusgrandiflorus (L.)Britt.&Rose flordelcáliz,pitahaya Sennaatomaria (L.)Irwin.&Barneby. frijolillo Serjaniadiversifolia (Jacq.)Radlk. bejucoangarilla Serjaniasubdentata Juss. bejucoesquinado Sidarhombifolia L. malva,malvadecochino Sideroxylonfoetidissimum Jacq.subsp. foetidissimum jocuma Sideroxylonsalicifolium(L.)C.F.Gaertn.(= Dipholissalicifolia A.DC.) cuyá Smilaxhavanensis Jacq. alambrillo Smilaxlanceolata L. zarzaparrilla Solandralongiflora Tuss. paloguaco Spermacoce assurgens Ruiz&Pavón garro Spermacocelaevis Lam. garromorado Spirotecoma sp. - Spondiasmombin L. jobo Sticherusbifidus (Willd.) Ching patadegallina Stigmaphyllonlineare Wr. bejucosanpedro Stigmaphyllonsagreanum A.Juss. bejucosanpedro Syzygiumjambos (L.) Alstonin Trimen pomarrosa Tabebuiabibracteolata Griseb. - Tabebuiabrooksiana Britt. robledeolor Tabebuiamyrtifolia (Griseb.)Britt. - Talaumaorbiculata Britt. marañóndecosta Thouiniapatentinervis Radlk. negracuba Tillandsiabalbisiana Schult. curujey Tillandsiafasciculata Sw. curujey Tillandsiapruinosa Sw. curujey Tillandsiarecurvata L. curujey Tillandsiaschiediana Stud. curujey Tillandsiausneoides L. barbadeindio Tournefortiaglabra L. nigua Tournefortiahirsutissima L. nigua Tournefortiavolubilis L. nigua Tragiahexandra Jacq. amansaguapo Reyes O.J. Lazaroa 37 2016: 43-103 103

Tragiavolubilis L. - Tremamicrantha (L.)Blume almezdeflorespequeñas Trichiliahirta L. cabodehacha Trichocentrumundulatum (Sw.) Ackerman&M.W. Chace guatacadeburro Trichomanesrobustum Fourn. - Trichostigmaoctandrum (L.)Britt. bejucodecanasta Triopterisrigida Sw. amansaguapo Trophisracemosa (L.)Urb. ramonhorse Turbinacorymbosa (L.)Raf. aguinaldodepascuas Turneraulmifolia L. marilope Typhadomingensis (Pers.)Kunth macío Urerabaccifera (L.)Gaud. chichicate Urochloahumidicola (Rendle)Morrone&Zuloaga yerbadeguinea Vanilla sp. vainilla Varroniaglobosa L. - Varroniaglobosa L.subsp. humilis - Varronialineata L. - Vernoniahieracioides Griseb. rompezaraguey Vernonia sp. - Viburnumvillosum Sw. - Vitistiliaefolia H.&B. uvacimarrona Wallenialaurifolia Sw. casmagua Waltheriaindica L. malvablanca Weinmanniapinnata L. oreganillo Wissadulaamplissima (L.)R.E.Fries. - Wissadulaperiplocifolia (L.)Presl. - Zamiaangustifolia Jacq. guáyara Zanthoxylumcubense P. Wils. ayúadesierra Zanthoxylumelephantiasis Macfd. bayúa Zanthoxylumfagara (L.)Sargent. amoroso Zanthoxylummartinicense (Lam.)DC. ayúa Zapotecagracilis (Griseb.)Bässler - Zeugitesamericana Willd. - Zuelaniaguidonia (Sw.)Britt.&Millsp. guaguasí