Distribution of tropical American rain forest species in the light of successional processes"! ———-GERARDO BUDOWSKI= COMPENDIO Denso del barge biinede wopiea, I expecies gue integuan las diferenter ota par de fe sucesin, macsban chetar ediacteristicas definidar en sw dismibacton Las especies pioneran y lar recendeatas tevnfrates tienen ana deribucibn ame pia Lat espociss eeundensos daaias som deviduss » pueden alcanzay ny tana eench devable canada legen a una edad avarzets Ellas jor naan peste del climax en fos bas- (ques recon a deciditos Eat ln conmenidades climes te encucttia mucho exdemisno, all, ie enendo la beterogeneided de expaces ef la regla, pueden davre caves de soviinmeds de mao pct especies debide a la dnflnentia de cierios actors eddficar — FI autor Invodnetion ISTRIBUTION paterns of rain forest species are the sesult of many interacting factors smong vhich climate, soils, relief and historic geology hheve been treates in cletail- There ate also some studics tn the cole of wind, animals and water in seed dis- PoMowever, thee is sill another Importint factor, namely plant succession, which bears important impli= cations on distribution patterns and wisich, for the tr piel tain forest, as unjustly ceceived scanty attention The present’ study attemps to show certain cels- tionships between saccestional processes and distribution pattetes for the tropicel American lowland forests. "The areas studied cover Iatge extensions in northera tiopicat America and Cential America with a mean anoval tem peratuce over 22%C (71.G*F) and a mean annual sae. fall over 2,000 mm (approximately 80 inches) Tbe clusifying of connmuniter in their proper tacer sinned ate In ocdes to appreciate the relationship between suc- cessional patterns and the distribution of ain forest © esvel fr pobinion Janay 2%, 1968 17 Rest CGR funn at Capon Eee + RGU” revotry Program, tater American texte of Agoura! Sime aig Sed Rewind “Cee "Parsi Qin" Re species, itis essoncal that successional serl stages be recognized, This can be done by recording. curefully 1 series of crtial lors, physiognomic and stuctual featur which will give enough dacs for classifying the different setal stages These stages cin be clled —for convenience pioneer, eatly secondary, old sec ondary and climax, snd comprise the critical characters shown in Table 1, which were taken from 4 sees of plots of which age and past intervention were well knowa (2, 3) ‘The gineralizstions shown on the table have been simplified to indiate relative walnes rather than absolute sumerical ones, sine the latter would be very difficult to render in detail at this stage However, expericnce hae shown thar the secceding of these characters docs aot offer a serious problem to those who have a fiir scquaintence with topical sain forest vegetation. ‘Whea such dlastification is achieved, 2 distinc cla. Aionship between distribution patterns of some species and their peopet successional stats ean be found The disnibution patieme of ploneest and early secondary species These two groups, here lumped together for con. venience, are found over areas of very different clima. lic and edaphic conditions Under conditions of closed undisturbed forest the presence of species of these suce ‘essional stages is limited to openings or gaps made by falling trees landslide: or other accidents including, 40BUDOWSKI; FOREST SPECIES IN SUCCESSIONAT. PROCESS A Table 1-—Chatacteristcs of arboreal components of seral stages ia tropical American humid Forests Number of woody] few 1.5. J few nto x00 fap to 100 oF a Kittle more discern wih age Une «an Immossncous. dense | vette buane hing, thin hetrogeecous. incudes | many vatible sper of EE ee Lrothenat oers ery weds crow ows ewer sawn | dense, tnled dense, lage esbaccowe | ilatioly sence. incaes| sence. vith tolerant species Trequert Folesunt specie Spee Gromh won fies eay fat mina fist others | slow or vey slow sw Life span vesw short less than 10) | short, 1025 years luvs 40-100 yems, | ery Ag. 100-000, some Aces cs ome mete pesbasiy mote ‘Tolerance to sade] very intolerant ey imvleeant foterane i jatenie stage, | oleae. except i alt donnie Iter" Intoferiet stage Rexeneraien of | very saver practically ehnent stscok or sbardaet with] Tash, abun Aominest faege' meray ia exty Dissemination of | hinds bats. wind winds bis. bass svi. pincipally evsiy, mamas rodent fale dominanis vos Bias Wood and scm, | seny ight sot diametens} very Veh iemetess | aight to meio tard, | hard anal heaey: inches Sinan below 60 a Sime ven tage stems | large stems size of seed, or | smal all oath medi lan Frais" aisperel Viability of reads [Ioas. talent in ait fons. tent in sid J chore to medio show Leares of domi | everentes evergicen any deailocus cversieen piphvies atbsent fow toy nes bo foe] many apes ar i Vines slouralnt, besbuccous barf abundant, berhaccous but] abundant, but few of | abundant, inclndes very Tew specks fee apcies than ies inepe woody sects nubs rmunp. bu few gpecies | selatively sbandnt bat | foe few im number but many tse andine dant or scarce weatce suatce2 TURRIALBA: VOL 15 NUM 1, TRIMESTRE ENFRO.MARZO 1965 cf course, iminsmade clearings These are the biole ics! nomads as they have been rightlp called by van) Steenis (4, 6) However, these species are also found in ther stiches such as the river banks, the edges of swomps and, i some cases, they are also. components ff the early tages of succession of the drier Formation, ie. the deciddous forest, although they are not deci uous themselves ‘The obvious implicaion is that thete species are well adapted to drought, be it lack of rainfall o° physiological. through flooding Very poct seils or racky outcrops also constitute niches where these species take reluge Their seed. dissemination mechanism is seny efficient ‘The small seeds are not only produced profusely but can remain dormant for a considerable period wader the shade of high canopy species until full. sunlight triggers their germination The distribution of pioneers anu erly secondary spscies his of course, considerably increased over the lise 50 years as result of mans impact on the vege tation The disitbetinn pattern of le secandeny species The mos sinking characteristic of these species is heir deciducusness, even in ateas of very heavy rainfall The most significant factor in connection with distribu. tion is that many of the species are also found in drier Tubitas, mainly the deciduous forest oc che very day forest In some of those formations they aaally re roduce well, ane hence may be considered as members ‘8 the imax This possibility hes been discussed by Tool ane! Richards (as comments on pape by Aubert dels Rae (1) Weis defintely true for many American species, such as Geetlalta seivirla, Biases somasnbs Lichen seomani, Condia alliadove, svong others Pos sibly Cache fentondsa, of panteopical distbution, enters into that extegory too Seaie of these older secondity species may actually remain in place for centuries and stain great size This has aso been pointed out by san Stenis (56), among others Curicuy, thea, Somme af the leegest trees oF the fpical rain Focest may ‘aften be old secondary species wich have remained in the arcs fora long period, bur do not regenctte. ‘This is certainly the case for some of the valsable Meliacene, Such as Sieeievia macrepbylla. Cediele atesieane sod some Bombiciceie. sich as Bonbucoptis sepinm snd Coiba pentandna tn Paoawa and Colombia: Cunanle~ Sur plauaiflie is anether oustanding example Dhe ditabntine af elintas spociee For conenience of definition, 2 climax community is the end product of a successional sere when rel tively stable —although certainly noe static— com: ‘munity has been reached and when changes of floristic composition, structure and physiogacimy over the age ‘yuan of the dominants became insiggaficant The sale in climax communities is a thorough mix: tuce of species whenever drainage is not extreme of impeded This has been sressed, among others by Richards (4) As scon as one single of a few species Fecome dominant, some edaphic factor, usually related to excessive water —at least duting patt of the yese— can be suspected bul, generally speaking, endemisan is frecuent from the floristic standpoint In conclusion it i felt that in order to understand Uistibution patterns of rain Forest species much consi: eration should be given to successional patterns The slassfieation of communities into theie proper succes sloral sequence, 1 matter which caa be achieved by carefully recording indicative flosstic, physiognomic and sructural characteristics of the communities, seems 10 be an essential previous step A large propoition of pioneers, early secondary, old secondary and climax spe ies appears to display 2 distribution pattern which is Iypieal of thefe successional status Sunmay Within the tropical sain forest dhe species that inte- urate different seral stages display characteristic disti bution patterns Pioneers and early: secanelary species have a wide distribution secondasy species are Aeciduous and ray attain considersble sive when they surow (0 old aye They are fact of the climax in the rier or deciduous forest Much endemism is Found communities; while 2 mixture of species is there may be edaphic factors which favor the dominance of one or a few species Literstare ented 1 AUBERT DE LARUE, © Mons inftucice on topiesl scavion fa Prncadings Nigth Pasic Scene Co tres. 1987. ‘Benghok Secretarnt Ninth Pace Seen fe Congras 1858 pp RTT Studies om fore succeeton in Casts Doctor of Paitonuphy thes Rew YYale Universi Sehool of Forestry 1980 5 Forest succession i tempi aweed Ta Wealby BAYH 198) 1 RICHARDS PW Wat the tepies cam contribute to Beclogy S12) 280201 198s 5 VAN STEENIS CG GJ Rejucenition ao juleing the statue of vegstition ten the Bi bona theony 1H Stody cf Teepe! Veseaxion Pro feedings of the Kandy Sjenposiom. Ceylon 18-21 Meech 1984 pp 212315 1958 Tropical lowland. vegetation; the character Tarcs of im pes. ata tir relation fo clinate dy Brovestings Nit Pactie scence” Cengtes, 1837 anstedeSecttaiat Ninth Baifc Science Contes peasar 19st