s t t i wo f t v t J DISTRIBUiÇÃO, VARIAÇÃO E USOS DOS...

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DISTRIBUiÇÃO, VARIAÇÃO E USOS DOS RECURSOS GENETICOS DE ARAUcARIAS NO SUL DO BRASIL J.Y.Shimizu - Y.M.M.de Oliveira Unidade Regional de Pesquisa Florestal Centro-Sul Colombo, Paraná BRASIL Resumo Araucária angustifolia (Bert.) O. Ktze é a única espécie do seu gênero que ocorre no Brasil, sendo as maiores populações concentradas no estado do Sul e a grandes altitudes onde predominam temperaturas baixas e chuvas abundantes e bem dlstribufdas. Reservas naturais tem sido drasticamente reduzidas pela derrubada de árvores há mais de 1 século, tanto para exploração de madeira como p/ uso da terra para agricultura. Estudos genéticos tem revelado um padrão discontinuo de variação através da area natural de distribuição. As raças geográficas mais promissoras em termos de crescimento em regiões específicos, tem sido identificados através de testes de procedência. Os ma is recentes esforços para preservar o valor genético das araucarias esta sendo tentado através da formação de populações base de procedência as quais conterão de variabilidade genetica original representativa de cada raça geográfica. DISTRIBUTION, VARIATION ANO USES OF ARAUCARIA GENE RESOURCES IN SOUTHERN BRAZIL Summary Araucaria angustifol ia (Bert.) O.Ktze is t he only species o f its genu5 to occur in Brazil. The major populations are concen- trated in the southern States. on the highlands where cool tem - peratures and abundant and well distributed rainfall predominate. Natural stands have been drasticaly reduced in extension due to cont inuous fell In95 for aver a century for timber exploration as well as to make room for agricul tural crops. Genecological studies have revealed a discontinuous pattern of variation through the natural distributian range. Geagraphic reces o f mc s.t pramissing grawth for specific regians heve been identified through a number of provenance tests. lhe most recent effort to preserve the valuable genetlc re- s ou r c e s i s aimed at t he formation of provenance base pc p u l a t l on s , which will hopefully contain the original genetic variability. representative of eaeh geographic race. Introduction Known by other names such as "pinhei ro brasi lei ro" (Brazi - lian p t ne ) , "pinheiro-do-paranã" (Paraná pinelor just e r a uc a r La , Araucaria angustifol ia (Bert,) O. Ktze ls t he only Arauearia sp!, cl es to oecur in Brazil. It has played an extremely important role in the country' s economy. Straight t rees wi th almost eyl - indric boles, reaehing 30 to SO m in height and 2 m in diameter at breast height were commonplaee in the Southern States. lhe s pec i es p rovldes high qual i ty timber for general con- s t ruc t i on, frame wo rk, f u r nl t ure, v eneer, pu\p and many other use s. Araucaria wood in form of sawn timber and v eneer was, for '" long p er i od, o ne of the most important i tems of Brazi I ian ex- po~tation. lhe orrgin~1 arauearia forests covered an estimated area of 200000 km 2 J (GOLFARI. 1971) of whieh, part Is in t he provinceof Hisiones in Argentina. lhe major formatlons are in the States of Paraná and Sa~ta Catarina. where intensive 10gging contin- ues to present. days. Extensive areas of natural arauearia fo- rests were turned into endless fields of coffee, wheat. sova beans and other crops, as a result of t he timber exploration and the expansion of agrieulture. lhus, many of its valuable geogr~ phyc r eces are b e l ieved to be extermined. lhe need to preserve this valuable gene resouree has led many Brazilian forest r eseerch institutions into a joint pro gramme on gene conservation. Natural Distrfbution Araucaric:o angustifol ia occurs naturally through the south- ern and s ou t heas t ern a r az t j, extending from the latitudes of 19 0 30'S to 31°30'S and longitudes of ~1030'W to S~a30'W, inclu- ding el so part of the pravince of Hisiones in Argentina (GOLFA- RI, 1971). lhe species t hr Lv es on deep fertile soils within the cl imatic.types classified, according to the Ktlppen's system, as "Cfa" and "Cfbll(humid sub-tropical without dry season and mi 1 d to hot s umme r L. Such el imatic types are predominant through t he s ou t hern highlands where arauearia forests ar~ coneentrated. Within the given range of lati t udes, t he s e types of el imate are found from lower altitudes in the south and progressing to higher al t Lt u - des to t he north. Following a similar pattern, natural stands of arauearia are found as a general rule, from SOO m above sea levei in the south to over 2000 m as the populations extend northward to the States of são paulo, Rio de Janeiro and Minas Gerais. Exceptions to t he s e altitudinal limits are also found, especially t n the south, where the s pecles c ccurs a t altitudes of a bo u t 300 m t n São Hartinho and Tenente Portela (HATlOS,1972) and as low as 250 m (PITCHER, 1975) and 198 m (GURGEL FILHO PISANI, 1975) In Lauro MUIler, near the coast of Santa Catar i - na. As mentioned by HAACK (1968), ~. angustifolla ex t ends to altitudes lower than soa m only where cool air flows from the highlands towards the vai leys. Normal mean temperatures in the arauearia region are 20 to 21 0 C in the Summer and Ia to lloc in the v t oeer (OLlVEIRA.]9~8). Not only cool to mild temperatures but also plenty of rainfall through the year is required by the species. Annual rainfall figures ranging from 1270 to 2494 mm (GOLFARI. 1971)are usual bu t not less than 1000 mm (HUECK. 1972). ~.an9ustifo - ~ is lorol eran t to water defieit and this i s the maln reason for l t s restriction to altitudes e b ov e 800 m i n the northern seetion of its distribution range (GOLFARI, 1971). where they occur in smal1 spots scattered along the more humld portions of the mountain ranges. lhe distribution pattern of ~-. angustifolia ls well illus- trated by HUECK (1972)(Fig. I). lhe extensive forests. conce~ trated on the southern highlands, are frequently interrupted by spots of natural grasslands. As explained by KLEIN (1960), they are remnants af a vegetation type which predominated dur- ing a dryer period, previous to the formation of araucaria fo~ ests. Variatian Although not very noticeable at first sight, A. an qu s t l f c l ia 287

Transcript of s t t i wo f t v t J DISTRIBUiÇÃO, VARIAÇÃO E USOS DOS...

DISTRIBUiÇÃO, VARIAÇÃO EUSOS DOS RECURSOS GENETICOSDE ARAUcARIAS NO SUL DO BRASIL

J.Y.Shimizu - Y.M.M.de OliveiraUnidade Regional de Pesquisa Florestal Centro-SulColombo, ParanáBRASIL

Resumo

Araucária angustifolia (Bert.) O. Ktze é a única espéciedo seu gênero que ocorre no Brasil, sendo as maiorespopulações concentradas no estado do Sul e a grandesaltitudes onde predominam temperaturas baixas e chuvasabundantes e bem dlstribufdas.

Reservas naturais tem sido drasticamente reduzidaspela derrubada de árvores há mais de 1 século, tanto paraexploração de madeira como p/ uso da terra para agricultura.

Estudos genéticos tem revelado um padrão discontinuode variação através da area natural de distribuição. As raçasgeográficas mais promissoras em termos de crescimento emregiões específicos, tem sido identificados através detestes de procedência. Os ma is recentes esforços parapreservar o valor genético das araucarias esta sendo tentadoatravés da formação de populações base de procedênciaas quais conterão de variabilidade genetica originalrepresentativa de cada raça geográfica.

DISTRIBUTION, VARIATION ANOUSES OF ARAUCARIA GENE RESOURCESIN SOUTHERN BRAZIL

Summary

Araucaria angustifol ia (Bert.) O.Ktze is t h e only species o f

its genu5 to occur in Brazil. The major populations are concen-trated in the southern States. on the highlands where cool tem -peratures and abundant and well distributed rainfall predominate.

Natural stands have been drasticaly reduced in extension dueto cont inuous fell In95 for aver a century for timber exploration

as well as to make room for agricul tural crops.

Genecological studies have revealed a discontinuous patternof variation through the natural distributian range. Geagraphic

r e c e s o f mc s.t pramissing grawth for specific regians h e v e been

identified through a number of provenance tests.

lhe most recent effort to preserve the valuable genetlc re-

s ou r c e s i s aimed at t h e formation of p r ov e na n c e base pc p u l a t l on s ,

which will hopefully contain the original genetic variability.

representative of eaeh geographic race.

Introduction

Known by other names s u c h as "pinhei ro brasi lei ro" (Brazi -

lian p t ne ) , "pinheiro-do-paranã" (Paraná pinelor just e r a uc a r La ,

Araucaria angustifol ia (Bert,) O. Ktze l s t h e only Arauearia sp!,

cl e s to oecur in Brazil. It has played an extremely important

role in the country' s economy. Straight t r e e s wi th almost eyl -

indric boles, reaehing 30 to SO m in height and 2 m in diameter

at breast height were commonplaee in the Southern States.

lhe s p e c i e s p r o v l d e s high qual i ty timber for general con-

s t r u c t i o n , frame wo r k , f u r nl t u r e , v e n e e r , pu\p and many other

use s.

Araucaria wood in form of sawn timber and v e n e e r was, for '"

long p e r i o d , o n e of the most important i tems of Brazi I ian ex-

po~tation.

lhe orrgin~1 arauearia forests covered an estimated area of

200000 km2J

(GOLFARI. 1971) of whieh, part Is in t h e provinceof

Hisiones in Argentina. lhe major formatlons are in the States

of Paraná and Sa~ta Catarina. where intensive 10gging contin-

ues to present. days. Extensive areas of natural arauearia fo-

rests were turned into endless fields of coffee, wheat. sova

beans and other crops, as a result of t he timber exploration and

the expansion of agrieulture. lhus, many of its valuable geogr~

phyc r e c e s are b e l ieved to be extermined.

lhe need to preserve this valuable gene resouree has led

many Brazilian forest r e s e e r c h institutions into a joint pro

gramme on gene conservation.

Natural Distrfbution

Araucaric:o angustifol ia o c c u r s naturally through the south-

ern and so u t h e a s t e r n a r az t j , extending from the latitudes of

19030'S to 31°30'S and longitudes of ~1030'W to S~a30'W, inclu-

ding el s o part of the pravince of Hisiones in Argentina (GOLFA-

RI, 1971). lhe species t h r Lv e s on deep fertile soils within

the cl imatic.types classified, according to the Ktlppen's system,

as "Cfa" and "Cfbll(humid sub-tropical without dry season and

mi 1 d to hot s umme r L.Such el imatic types are predominant through t he s ou t h e r n

highlands where arauearia forests ar~ coneentrated. Within the

given range of lati t u d e s , t he s e types of el imate are found from

lower altitudes in the south and progressing to higher al t Lt u -

des to t h e north. Following a similar pattern, natural stands

of arauearia are found as a general rule, from SOO m above sea

levei in the south to over 2000 m as the populations extend

northward to the States of são paulo, Rio de Janeiro and Minas

Gerais. Exceptions to t he s e altitudinal limits are also found,

especially t n the south, where the s p e c l e s c c c u r s a t altitudes

of a bo u t 300 m t n São Hartinho and Tenente Portela (HATlOS,1972)

and as low as 250 m (PITCHER, 1975) and 198 m (GURGEL FILHO

PISANI, 1975) In Lauro MUIler, near the coast of Santa Catar i -

na. As mentioned by HAACK (1968), ~. angustifolla e x t e n d s to

altitudes lower than soa m only where cool air flows from the

highlands towards the vai leys.

Normal mean temperatures in the arauearia region are 20 to

210C in the Summer and Ia to lloc in the v t o e e r (OLlVEIRA.]9~8).

Not only cool to mild temperatures but also plenty of

rainfall through the year is required by the species. Annual

rainfall figures ranging from 1270 to 2494 mm (GOLFARI. 1971)are

usual b u t not l e s s than 1000 mm (HUECK. 1972). ~.an9ustifo -

~ is l o r ol e r a n t to water defieit and this i s the maln reason

for l t s restriction to altitudes e b ov e 800 m i n the northern

seetion of its distribution range (GOLFARI, 1971). where they

o c c u r in smal1 spots scattered along the more humld portions of

the mountain ranges.

lhe distribution pattern of ~-. angustifolia l s well illus-

trated by HUECK (1972)(Fig. I). lhe extensive forests. conce~

trated on the southern highlands, are frequently interrupted

by spots of natural grasslands. As explained by KLEIN (1960),

they are remnants af a vegetation type which predominated dur-

ing a dryer period, previous to the formation of araucaria fo~

ests.

Variatian

Although not very noticeable at first sight, A. an qu s t l f c l ia

•287

presents some morphological variation which might be, to some

extent, associated to the environmental conditions. Tr~es grow -

ing on high and cold flatlands in Santa r a t a r t e a , o n- shallow I i-

thosol near São Joaquim, were r e pc r t e d by COLFARI (1971)as being

larger in diameter and disproportionately s ho r t e r {e r ou n d 10 m in

he Lq h t Lt h a n t he t r e e s on warmer s t t e s with deeper soil, wh e r e

they reach over 30 m in height.

The size of the seeds also varies remarkably. CASTRO (1959)

Fig. I. Natural distribution a r e a s of Araucaria

angustifolia {f r o m HUECK, 1972).

has shown t h e t , seed size, when measured by its length, is asso-

ciated to t he altitude of its s ou r c e between 500 and 1580 m.

Smaller seeds were ~roduced by stands from higher altitudes. On

the other hand, the latitude of the seed source was shown by

BANOEL (1966), to b e more fmportant than altitude to the vari -

ation of the seed weight. In his study, seed weight increased

with l n c r e a s e in the lati tude of t h e sources.

Other types Df variation exist whose pattern of distributon

or the association to specific environment is not yet cl e a r .

REITZ & KLEIN (1966) described them as 9 varieties and one form

of the specles as follow:

a) ~. angustifolia varo elegans {Ho r t . } Reitz & Klein.

(Branches slender and numerous; needles smaller and closely

i n s e r t e d} .

b} ~. angustifolia varo sancti josephi Reitz & Klein.

{Kernels ripe In February and Harch}

c) ~. angustifol ia varo caiava Reitz " Klein.

(Kernels ripe in June and July)

d) ~. angustifol ia varo Indehiscens Mattos.

(Kernels..smal1 e r , ripe from September to January bu t do no t

fali by themselves off the cone).

e) ~. angusti foi ia varo angusti foi ia

(Ke r n e I s r ed , r i p e I n Ap r i 1 a n d Ma y)

Uses of Araucaria angustifol Ia Cene Resources

~. angustifolia varo ~ Reltz " Klein

(Kernels dark-red almost black before drylng)

g) ~. angustifol ia varo striata Reitz & Klein

(Kernels r e d with darker l o n q t t u d i n a l s t r f p e s )

Of ali these variants, the type variety seems to be by far

t he most frequent. For practical purposes, ~. angustifol ia is

expected to mature between Harch and Hay.

Several experiments for the study of the species racial va~

iation h a v e been establ ished in Brazil and Argentina. A t e s t

planted in Santa Rita do Passa Quatro, SP, (CURCEL " CURCEL FI-

LHO, 1973) showed remarkable differences between 3 widely dis -

tributed s~ed sources. At 12.5 years of age. volume production

of Capão Bonito provenance was 49% and 68% greater than Campos

do Jordão and Lages provenances respectively. In a larger ex -

periment involving 23 p r o v en e n c e s planted in Tres Barras, SC

(GURCEL & CURCEL FILHO, 1968), better growth was obtained with

seeds from São Paulo and southern Hinas Cerais than from Para

ni. Santa Catarina and Rio Crande do Sul. Campos do Jordio

provenance produced larger number of branches and was more sus-

ceptible to frost, while Irati p r c v en e n c e showed a pecul i a r ye!

lowish-green needles.

Crowth differences among provenances have been quite disco~

tinuous through the sampled area. Correlations of this ch~rac-

ter with the environmental factors at the seed sources have not

been conslstent. Crowth was stgntflcantly correlated with lon-

gitude and annual mean t emp e r e r u r e but not with annual rainfall,

el t Lt u d e or latitude. ln a t e s t planted t n Puerto Libertad, Ar-

gentIna (FAHLER & DI LUCCA, 1979), while In southern são Paulo,

Brazi 1 (SHIHIZU & HrCA, 1979). only the lati t u de at s e e d source

was correlate~ to height growth up to stx years of age. Thls e~

periment showed further that growth dlfferences among provenan-

ces and the correlation of height groHth with latitude of seed

source tend to decrease with age.

Only in some cases (BALOANZI e t a l • , 1973; FAHLER & 01 LUC-

CA, 1979) h a v e the local provenances performed as well as the

best in the group; in others (CURCEL & CURCEL FILHO, 1968; SHI-

HIZU " HIGA, 1979; ~ONTEIRO & SPELTZ, 1979) they have not shown

outstanding growth. Thus, t h e optimal ity of t h e local r e c e for

early growth can not be generallzed for the species.

Negative correlations between altitude at seed source and

groW'th at 3.5 years (KACEYAHA & JACOB, 1979) and at 2 years of

age (SHIHIZU " HICA. 1979) we r e statistically significant. How~

ver, It i s I ikely to reflect mostly t he effect of seed size on

e e r f v growth; low altitude s cu r c e s havtng larger seeds (CASTRO.

1959), would grow more than those from higher altitudes. The

effect of seed size on t h e growth r a t e of~. angustifolia was

shown by CLASER " OLIVEIRA (1972). Seeds larger than 6 cm In

lenght resulted in plants growing 18% more than those de r l v e d

from seeds smaller than 3~9 em in length and 16% more than tbo se

derlved from the mixture of ali sized seeds, at the fourth year

a f t e r plantin.g.

Araucaria forests have been the main source of high qual i ty

timber for construction, frame work, furniture, v e n e e r , pulp

and other uses.

Sawn timber and veneer have been exported to severa I

tries for a 10ng period; at times they were rated second only Wc o f f e e in exported v al u e .

The araucaria reserves in the State of sio Paulo were

hausted 10 the period between 1930 and 1940 (KRUG, 1964).

h) ~. angustifol ia varo semi-alba Reitz &

(Kernels with white tip b e f o r e drying)

Klein

As estimated by MAACK (1968), 7·4 million hectars of natural

araucaria forest existed In the State of Paraná, of whlch only

34% was left by 1950 and 28% by 1960. The remnants l n the State

were f u r t he r reduced to 433 580 ha (MACHADO, 1975) in 1967 and

316620 ha (FUPEF/IBOF, 1978) in 1977, the l e t t e r being only

4.3% of the original area.

Araucarla tlmber production In ali States l nc r e a s e d from

1.5 million c v bl c me t r e s In 1945 to 3.3 mtllton tn 1950. From

that year to 1966, t h e output was steady around 2.8 mi 11 ion cu-

bic me t r e s a year. Then untll 1972, l t d e c r e a s ed to an annual

i) ~. angustifo! ia varo alba Reitz & Klein

(Kernels white ar yellowish before d r vínç I

j) ~. angusti foi ia for. monoica

(Strobili mate and female on the same tree)

288

output of. 1.8 mi l l ion cubic m e t r e s (PARANA. 1976).

Reforestation with arãucaria was done at first by a few co~

p e nl e s as wel l as by the federal government. Close to 90 000

ha had be e n planted by 1979, according to the estimates from

severa I s o u r c e s .

~' angustifolia is nç t c o n s l d e r ed a fast growth species in

Brazil. Volume in c r eme n t s ln mature natural stands we r e e s tl mq

ted to a maximum of 2.11 m3/ha/year (HUNIZ, 1966), while in

planted stands, it varied greatly depending on the site quali -

ty. Inside be r k volume increment in planted stands varied from

2.56 m3/ha/year in the poorest sites to 14.48 in the best sites

l n a su r v e v by HOOGH (1978). The same variable compared in

different sites by $PELTZ (1973) r e v e a l ed growth ln c r em e n t

varying from 1.1 m3/ha/year on grassland area to 10.8 m3/ha/year

on f o r e s t s o l I.

Oue to such growth be ha vl o u r , plantatlon r e t e s decl ined

sharply in the last years to not more than 300 ha a year, giv-

ing place to t h e l n t r od u c e d Pinus ell10ttii,!.. taeda and~-

~ which are less site d erne nd Lnq and have higher growth ra-

tes.

Population Improvement

As investments in reforestation wi th ~' angusti foi ia

become less attractive due to its slower growth rate as compa -

red to Pinus and Eucalyptus s p e c i e s , efforts for population g!

netic improvement of the s p ec l e s have been lnslpient.

A few planted or natural stands have been maintained for

seed production areas by t he Brazilian Institute for Forestry

Oevelopment (IBOF) and some compani'es as well.

Further experiments for the study of provenance and proge-

ny variations are under way, in connection to the formation of

genetlc base populations for l mp r o v erne n t wo r k , u nd e r t he c o o r r-

dinatlon of the Brazilian Working Group on Forest lree Impro -

vement. Lhe s e wlll hopefully lead to a more precise mapping

of t he seed z on e s , l f important differences among provenances

are ma Ln t a l n e d through the rotation age. Also other important

facts on population structure and patterns of variation at

dlfferent levels may be known.

Seed collections from natural stands hav e been organized

through joint effort by the main forest research institutions

in the araucaria region.

Seed procurement, their sources and storage problems

Procurement of araucaria seed has become harder in the l e s t

years, due to continuous fell ings of ma t u r e , seed bearing natu

r a l stands. Horeover, bumper seed crops occur a t times only in

1 l mI ted a r ee s , Huch o f t he s e e d fa II en off the t r e e s are eaten

by wlld and domestic animais and people as well.

Large scale production of planting stock has, therefore,

frequently depended on s u p pl i e s from distant natural s t a n d s .

Seed s u p p l y from a bumper crop to be sown in the followi ng

years has not been feasible due to storage problems. Seed vi~

bility d e cl l n e s. sharply from 100% to almost 0% in 3 to 4months

l f adequa te storage facility were not available. By keeping

t h e seeds in plastic bags at low temperatures (o to SOC), via-

bil ity was maintained between 70 and 90% after 4 month storage

(PRANGE, 1964).

Vegetative propagation

Several methods of vegetative propagation have been tried

with~. angustifo.lia. Neither air-Iayering no r rooting of hoI.

mon treated cuttings, tried by BANDEL (1966). did lead to any

promisslng result. Certainly, a number of factors must be

taken lnto account and caréfully controlled •. Rooting sucesses

of 2S to 26t was r e po r t e d by TESDORFF (1969) wi th cuttings ta-

ha 5

ken from 12 ve e r r-ol d t r e e s and treated with 0.5t, 1.0~ and

2.0% s ol u ti o n s of indol-butyric-acid.

Among severa! grafting methods tested by GURGEL & GURGEL

FILHO (1967), the best result (27t) was obtained with veneer

s id e graft.

Similarly to t he growth habit of other Araucaria species,

grafting of lateral s ho o t s of~. angustifol1a results l n pl e -'

giotropic growth ando if straight growth were desired, ortho-

tropic shoots must he grafted.

The serious limitation of most grafting m e t ho d s l s the

s ma l ! number of o r t h o t r o p !c shoots that each tree usual1y pr~

duces.

Lhe "p a t c h graft" method, as described by NIKLES (1973),

appl ied sucesfully to ~' angustifoJ ia by KAGEYAMA & FER -

REIRA (1975). lhe main advantage of t hl s method l s that many

buds can be obtained from each apical shoot and therefore,the

number of grafts per tree can be greatly increased.

Constraints on Progress in Populatibn Genetic Improvement

lhe main difficulties in promoting genetic improvement in

this species arises when breeding l s considered. lhe trees

usually produce seed only after 15 or 20 years of age (BANOEL

& GURGEL, 196]).

Controlled polI ination work on t h e trees is almost impossl

ble due to the height and targe span of the crown. The repro-

ductive organs are produced at the tips of the branches, in

the midst of densely packed tufts of prickly needles.

Large amounts of seed is annually l o s t to cone and s e e d in

sects whose behaviour and control methods should be better

understood.

Vegetative propagation techniques must be further studied

e n d directed toward a more efficient method.

Very rarely has ~' angustifol ia been planted in large sc~

le i n recent years. Being a very site demanding species, it

Is rapiddly loosing ground to the introduced Pinus and Eucaly~

~ species which have higher growth rates. Extensive

in the araucaria region have e l s c been turned into endless

fields of coffee, wheat, sova bean and o t h e r c r o p s .

L'l t e r e t u r e Clted

BALOANZI, G.; RITTERS'HOFER, F.O. & REISSMANN, C.B., 1973, En -

saio comparativo de procedências de Araucaria angustifo -

1 ia {Be r t .") O. Ktze. l n : CONGRESSO FLORESTAL BRASILEI -

RO, 2., Curitiba. Anais. FIEP, p p . 123-124.

BANOEL, G., 1966: O pinheiro brasileiro Araucaria angustifo -

I ia (Bert.) O. Ktze. ESALQ, Piracicaba. 67pp.

BANOEL, G. & GURGEL, J.T.A., 1967: Proporção do sexo em Arau-

caria angustifolia. Silvicultura em São Paulo, v ol , 6,

p p • 209-220.

CASTRO, Y.G.P., 1959: Variação do tamanho de sementes de ~

caria angustifolia {s e r t . ) O. Ktze. Anuário Brasileiro

de Economia Florestal, n? 11, pp. 124-133.

FAHLER, J.C. & DI lUCCA, C.M., 1979: Variaciõn geográfica de

Ar e u c a r La angustifol ia (Bert.) O. Ktze. 26pp. (Presented

in t he first IUFRO Meeting on the Araucarias, Curitiba -

unpubl ished).

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