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Research Note

Early Performance of CAMCORE Introductions


of Pinus patula in Brazil, Colombia and South Africa

w. S. Dvorak, J. K. Donahue, and J. A. Vasquez


Box 7626, Grinnells Laboratory, College ofForestResources, North Carolina State University, Rnleigh, NG. 27695 USA

SYNOPSIS

Seeds were collected from 13 provenances and 282 mother trees of Pinus patula Schiede & Deppe across a 900
km transect from Tamaulipas to Oaxaca, Mexico by the CAMCORE Cooperative, North Carolina State
University, USA. Thirty-three provenance/progeny tests were established in Brazil, Colombia, and South Africa
and assessed for height and diameter (DBH) at three years ofage. Sources from southern Oaxaca (Cuajimoloyas,
Manzanal, and Tlacuache) showed great susceptibility to cold atJessievale and Commonage, South Africa (26%
survival) versus the other P. patula sources (70% survival) . Average height growth at three years ofage exceeded
5 m in tests at Los Guadales, Colombia, and Maxwell, South Africa. The best provenances for height growth and
volume in both Brazil and South Africa were those from the central part ofthe P. patula range: Potrero de Monroy
(Veracruz), Zacualtipan, (Hidalgo) and Pinal de Amoles (Queretaro). In the Andean mountains of Colombia,
the southern Oaxacan sources of P. patula performed as well as the best sources from central Mexico. Some
CAMCORE open-pollinated families performed as well as or better in height growth than advanced generation
control lots of P. patula, P. taeda or P. elliottii in 29 of 33 tests.

INTRODUCTION of 4 provenances and 22 mother trees of P. patula from


the states of Puebla, Hidalgo and Oaxaca in 1969 and
Approximately one million hectares of Pinus patula
1970 (DarrowandCoetzee, 1983). The seeds from these
plantations have been established in the tropics and
collections were planted at Belfast,Jessievale, Mac-Mac,
subtropics for sawtimber and paper products (Wright,
Hebron, Spitskop and Tweefontein in South Africa. The
1994, Birks and Barnes, 1991). The majority of the P.
Center for Natural Resources Research (INTA) , Argen-
patula forests are located in southern Africa with South
tina, sponsored Dr. W. Barrett to collect 16 P. patula
Africa having more than 295 000 ha under operational
provenances and 110 mother trees in Mexico in 1972
management at the present time (S.A. Dept. Envn.
(Barrett, 1972; Barrett, 1978). This seed collection was
Affairs, 1993). Pinus patula is also an important commer-
unique because it covered the known range of P. patula
cial species in the Andes mountains in Colombia and
at that time and included the first sample from the
Ecuador. In southern Colombia, where approximately
northern state of Tamaulipas, at EI Cielo! (23 0 05' N
50000 hectares are planted, P. patula is used for news-
latitude). The seeds from this collection were planted in
print, sack paper, treated lumber and posts (Wright,
several genetic tests, most in the Jujuy state in northern
1994; Wright and Ladrach, 1994).
Argentina but one at Mac-Mac, South Africa (Picchi,
Even though P. patula is one of the most important
1988). In the early 1980s, the Food and Agriculture
commercial forest species in the tropics and subtropics,
Organization (FAO) distributed bulk seedlots from 17
the genetic base from which many plantations are de-
provenances of P. patula collected by the Mexican gov-
rived in southern Africa apparently originated from only
ernment forestry organization Instituto Nacional de
three commercial collections in Mexico made at the
Investigacion Forestal (INIF) (Forest Genetic Resources
beginning of the century (Burgers, 1975). It is thought
Information, 1982). Little information has been pub-
that these may have been collected in the Mexican states
lished from these studies but data is currently being
of Hidalgo and (northern) Oaxaca (Wormald, 1975).
analysed by several of the participants in both South
Therefore, the seed produced from seed orchards in
Africa and Zimbabwe.
southern Africa and used throughout the region and
In the mid 1980s, stafffrom the Central America and
also sold to Latin America presumably comes from a
Mexico Coniferous Resources Cooperative, (CAM-
relatively limited genetic base.
There were three major provenance collections of P.
1 Barrett referred to this collection as "Sierra de Guatemala" the
patula between the late 1960s and early 1980s to broaden name of the local mountain range. The CAMCORE "El Cielo"
the genetic base of the species. Coetzee and Fisk of the collection site, is part of the biosphere, "Rancho del Cielo" which
South African Department ofForestry made a collection is in the same general vicinity where Barrett presumably sampled.

Suid-Afrikaanse Bosboutydskrif- nr. 174, November 1995 23


CORE), North Carolina State University, USA, identi- MATERIAL AND METHODS
fied several new stands of P. patula while making explo-
rations and seed collections for other pine species in In the initial distribution, seeds from 13 provenances
southern Mexico and believed that additional and 282 mother trees were sent to members of the
populations existed that had never been sampled before CAMCORE Cooperative in Brazil (Klabin), Colombia
by other agencies. At the urging of the South African (Smurfit Carton de Colombia) and South Africa
Forestry Research Institute (SAFRI) 2 which was a full (MaNDl, SAFCOL and SAPPI) and established in 33
member of the cooperative program at that time, provenance/progeny tests (Figure 1 and Table 2). The
CAMCORE initiated range-wide explorations and seed number of provenances and families per provenance
collections of P. patula in Mexico in 1986 to better included in each test varied from one cooperator to
protect dwindling gene pools of the species and broaden another. Each member established provenance/prog-
the existing genetic base being used by forestry agencies. eny tests using the standard CAMCORE compact family
From 1986 to 1994 seeds were obtained from 22 prov- design (Wright 1978) with nine replications and six-tree
enances and 510 mother trees from Tamaulipas to row plots. Also included in each test was a series of
Oaxaca, a distance of approximately 900 km (Dvorak common control lots. CAMCORE included 3 open-
and Donahue, 1992). The collections included a series pollinated families from the Potrero de Monroy,
of new provenances from the Sierra Madre del Sur in Veracruz, provenance in 32 of the 33 tests. In the 24 tests
southern Oaxaca not sampled by either the South Afri- in South Africa, CAMCORE members also included 5
cans or Argentineans or included in the FAa distribu- common controls of commercial and advanced genera-
tions. Furthermore, the CAMCORE effort included the tion P. patula and P. elliottii. In Brazil, Klabin included
most northern source of P. patula yet found (Conrado two controls of commercial P. taeda and P. elliottii from
Castillo, Tamaulipas 24° 00' N Lat.) approximately 100 local seed stands. The best performing control lot in
km north of Barrett's EI Cie10 site, as well as new sites in each test is reported here.
the state of Veracruz (Table 1). At present, seeds for 80 Total shoot height (m) and diameter at breast height
P.patula CAMCORE conservation plantings and genetic (cm) were measured at three years of age and volume
tests have been distributed to members in Brazil, Chile, was calculated using a formula for juvenile trees
Colombia, Mexico, South Africa and Zimbabwe. 0,00003d2h (Ladrach, 1986). A preliminary analysis of
This paper reports on three-year provenance/prog- our data showed that height-diameter ratios varied sig-
enyresults of33 tests established in Brazil, Colombia and nificantly across sites. Therefore, even though the trials
South Africa with seeds from the CAMCORE collections. were young, both height and volume were used in the
Provenance x site and genotype x site interactions were analyses ofprovenance and family productivity to obtain
examined to determine their effect on the structure of
breeding populations. Recommendations are made on • SAFRI membership eventually passed to the Department of Water
Affairs and Forestry (DWAF) and then to the South African
efficacy of continued seed collections for P. patula in Forestry Company Ltd. (SAFCOL). MONDI and SAPPI became
Mexico. members of CAMCORE in 1988.

TABLE 1. Summary of the CAMCOREPinus patula seed collections in Mexico, 1986-1994

Provenance State Latitude North- Elevation Annual No. of


Longitude South (m) Rainfall (mm) trees

Carrizal de Bravo Guerrero 17°35' - 99°51' 1980-2440 1209 10


Cumbre Muridores Hidalgo 20°19' - 98°21' 2380-2480 1869 30
La Cruz Hidalgo 20°17' - 98°18' 2300-2450 1869 24
La Encarnacion Hidalgo 20°45' - 99°13' 2400-2650 1200 18
Zacualtipan" Hidalgo 20°39' - 98°40' 1980-2200 2047 15
Cuajimoloyas!' Oaxaca 17°11' - 96°26' 2450-2770 1135 16
EI Manzanal" Oaxaca 16°06' - 96°33' 2350-2660 1348 28
EI Tlacuache" Oaxaca 16°44' - 97°09' 2300-2620 2000 28
Ixtlan'? Oaxaca 17°24' - 96°27' 2600-2870 1750 27
Juquila Oaxaca 16°13' - 97°14' 2090-2260 1325 21
Santa Maria Papale" Oaxaca 17°49' - 96°48' 2270-2720 1100 24
Pinal de Amoles' Queretaro 21°07' - 99°41' 2380-2550 1350 24
Llano Las Carmonas" Puebla 19°48' - 97°54' 2530-2880 1097 26
Conrado Castillo' Tamaulipas 23°56' - 99°28' 1500-2060 1012 25
EI Cielo Tamaulipas 23°04' - 99°14' 1600-1730 1200 23
Tlacotla" Tlaxcala 19°40' - 98°05' 2750-2915 1097 21
Corralitla" Veracruz 18°38' - 97°06' 2000-2230 1500 23
Cruz Blanca Veracruz 19°39' - 97°09' 2500 1347 8
Ingenio del Rosario' Veracruz 19°31' - 97°06' 2770-2870 1346 22
Petrero de Monroy" Veracruz 20°24' - 98°25' 2320-2480 1350 31
Manzanares Veracruz 19°39' - 97°08' 2400-2450 1295 30
CaIcahuaIco Veracruz 19°07' - 97°06' 2350-2400 2020 30
Magueyes Veracruz 18°53' - 97°16' 2250-2350 1200 30

1 Superscripts indicate map locations in Figure 1 of provenances referenced in this report.

24 South African Forestry Journal- No. 174 November 1995


26"N a more accurate measure of growth potential. Analyses
of variance for height and volume were conducted for
each individual test using the General Linear Model
Distribution of
". (GLM) procedure of SAS/STAT software (SAS Insti-
Pinus potuta tute, 1989). Provenance differences were assessed using
the Waller-Duncan test (Waller and Duncan, 1969).
22'
Family performance of the CAMCORE material was
statistically compared to the commercial and advanced
20'
generation control lots by Dunnett's test (Dunnett,
1980).
From the 33 tests, 6 trials (05L, 05El, 05E2, 05J, 05K
P<lciftc Ocecn
and 05Il) with the most balanced complement of prov-
enances and families were chosen for across-site analyses
in Brazil and South Africa. The objective of the multi-site
... - CAMCORE Prov"nonc" Location 16'
analysis was to determine if the best provenances and
families at one location were also the best at other sites.
Four provenances and 45 open-pollinated families were
FIGURE 1. Location ofthe Pinus patula provenances included common to the 6 test sites. An average of 11 open-
in the trials pollinated families represented each provenance. Across-
site analyses ofvariance were conducted on the trials for

TABLE 2. Summary of the CAMCORE Pinus patula test sites

Company/Test code Location Lattitude Evaluation Annual Rainfall


(m) (mm)

KLABIN BRAZIL
20-26-0511 Imbauzinho 24°16'S 780 1473
20-26-05J Marica 27°08'S 920 1464
20-26-06Cl Imbauzinho 24°16'S 780 1473
20-26-06Cl Campina Grande 27°45'S 830 1463
20-26-07Cl Imbauzinho 24°16'S 780 1473
20-26-07C4 Campina Grande 27°43'S 780 1473

CARTON DE COLOMBIA COLOMBIA


20-02-02C Los Guadales 02°25'N 1750 2046
20-02-03B Sombreros 02°15'N 2450 2177
20-02-04B Sombreros 02°15'N 2450 2177

MONDI FORESTS SOUTH AFRICA


20-18-01J Commonage 31°02'S 1480 757
20-1-02E Commonage 31°02'S 1480 757
20-18-05K Commonage 31°02'S 1480 757
20-18-06D Commonage 31°02'S 1480 757
20-18-07A Commonage 31°02'S 1480 757

SAFCOL SOUTH AFRICA


20-10-01A Tweefontein 25°03'S 1150 1274
20-10-01B Jessievale 26°14'S 1730 921
20-10-02A Tweefontein 25°03'S 1150 1274
20-10-02B Jessievale 26°14'S 1730 921
20-10-03Al Tweefontein 25°03'S 1150 1274
20-10-03A2 Jessievale 26°14'S 1730 921
20-10-04Al Tweefontein 25°03'S 1150 1274
20-10-04A2 Jessievale 26°14'S 1730 921
20-10-05E1 Tweefontein 25°03'S 1150 1274
20-10-05E2 Jessievale 26°14'S 1730 921
20-10-06B1 Tweefontein 25°03'S 1150 1274
20-10-06B2 Jessievale 26°14'S 1730 921
20-10-07Bl Tweefontein 25°03'S 1150 1274
20-10-07B2 Jessievale 26°14'S 1730 921

SAPPI FORESTS
20-07-01L Maxwell 30 002'S
1350 817
20-07-02D Maxwell 30 0 0 2 ' S 1350 817
20-07-05L Maxwell 30 0 0 2 ' S 1350 817
20-07-06E Maxwell 30 0 0 2 ' S 1350 817
20-07-07D Maxwell 30 0 0 2 ' S 1350 817

Suid-Afrikaanse Bosboutydskrif- nr. 174, November 1995 25


height and volume. Analysis was done on a plot mean Colombia, the southern Oaxaca sources of P. patula
basis in order to facilitate the computational perform- showed as much promise as the best central Mexico
ance. sources. Provenance differences were statistically signifi-
To determine the practical importance of rank cant in 23 of 33 tests for height and 26 of 33 tests for
changes of provenances and families across environ- volume.
ments, stability analyses were conducted by computing The correlation between height or volume of trees at
ecovalences using a modified SAS program from Maxwell, South Africa and latitude of the collection site
Fernandez (1991). An ecovalence is simply the sum of in Mexico was not significant (r=-0,42). There was also
squares contribution of a genotype to the total sum of no significant correlation between growth or productiv-
squares. Significance tests were conducted on each prov- ity and elevation of the collection site (r= 0,01). How-
enance and family to determine if their response paral- ever, when only six provenances from central Mexico
leled the mean response of all genotypes in the trial (see were included in the analysis (Pinal de Amoles,
Lin et al., 1986). A stable genotype by this definition Zacualtipan, Llano Carmonas, Tlacotla, Ingenio Rosario
implies stability only with respect to the other genotypes and Corralitla), the correlation between height or vol-
in the test and can not be generalised (Lin and Binns, ume with latitude was -0,85 and -0,88, respectively and
1991). Since the number of individual trees represent- highly significant (Figure 2).
ing the control lots was much less than those represent- In approximately 60% ofthe tests, the best CAMCORE
ing provenances and families across sites, control lot provenance was as good as or better in height/volume
data was excluded from multiple means and ecovalence than the most productive controls (Tables 3 and 4). On
comparisons. average, 23% of the CAMCORE unimproved families
Because future seed collections of P. patula were performed as well as or better in height growth than
planned by the CAMCORE Cooperative, the mean height advanced generation controls in 29 trials (Table 5). In
and volume of provenances at three years of age were the remaining four trials (20-Q7-07D, Maxwell, and 20-
correlated with the latitude and altitude of the collec-
tion site in Mexico to determine if either would serve as Volume, dm-
a good predictor ofseed source performance. Five of the
7..5 r - - - - - - - - - - - - - - - - - - - - - - - ,
33 tests were used in the correlation analyses. The tests • Potrero
chosen (OIL, 02D, 05L, 06E, and 07D) were established de
by SAPPI on the same site at Maxwell, South Africa (30 0
S. Lat.). The five trials contained all 13 provenances
shown in Figure 1. Even though the site is rather homog-
r =- 0.88 Monroy

enous, replication means for all five tests were standard-


ized using the common controls before the correlation 7,0 Ingenio
analyses were run. del
Roaario

Corrallta
RESULTS

Survival of the tests ranged from 34% to 96% and Llano de


averaged 83% (Table 3). Two provenances from south- laa Carmonaa
ern Oaxaca (Manzanal and Tlacuache) and one from
central Oaxaca (Cuajimoloyas) were extremelysuscepti-
ble to freezing temperatures relative to the other P.
patula provenances tested. At two of the coldest sites, Tlacotla
Jessievale and Commonage, South Africa, where freez-
ing temperatures are common during winter, the aver-
age survival of these three Oaxacan sources was 26% Pinal de
Amolea
versus 70% for the other remaining provenances. At
Zacualtlpan
Maxwell, South Africa, where freezing temperatures are
infrequent, all sources had approximately the same
survival (90%).
Height and volume of provenances are shown in
Tables 3 and 4. Average height growth equalled or
5,5 '------''--..--J._----'-_--'-_--'--_-'-_-'--_l....-----'_..--J._---J
exceeded 5 m in tests at Los Guadales, Colombia (test
18~5 19..0 20..0 20,5 21,0
02C) and Maxwell, South Africa (test 06E). The poorest
growth was on dry and cold sites with sandy soils at Latitude N
J essievale, South Africa (test 02B). The best provenances
in South Africa and Brazil were Potrero de Monroy, FIGURE 2. Correlation between mean volume peiformance of
Zacualtipan and Pinal de Amoles from the central part Pinus patula provenances at Maxwell, South Africa, and the
ofthe P. patularange in Veracruz, Hidalgo and Queretaro. latitude ofseedcollectionsites in Mexico. The Potrerode Monroy
In more tropical areas like the Andean highlands of proenance was not used in the analysis.

26 South African ForestryJournal- No. 174 November 1995


~ TABLE 3. Mean height (m) for Pinus patula assesed at 3 years of age in Brazil, Colombia and South Africa
~
~ Test Survival Conrado Pinal Zacual- Potrero Llano Tlacotla Ingenio Corralitla Santa Ixtlan Cuajimo- El El Test Control
~ Code % Castilo de tipan de de las del Maria loyas Tla- Man- Mean
~
~ Amoles Monroy Carmonas Rosario Papalo cuache zanal
~
20-02-02C 83 5,4 5,6 5,4 5,7 5,6 5,5 5,4CC
~ 20-02-03B 86 3,4 3,3 3,3 2,9 3,4 3,3 3,4CC
w
~
20-02-o4B 94 3,5 3,5 3,7 3,6 3,7CC
e- 20-07-olL 96 4,9 4,8 4,8 4,8 5,0 P7
;0,-
20-07-02D 90 4,8 4,3 4,2 4,1 4.3 4,3 4,8P9
~
I 20-07-o5L 94 4,3 4,3 4,7 4,8 4,5 4,3 4,5 4,9 P9
;:l
;-! 20-07-o6E 94 4,7 4,7 4,8 5,4 5,1 5,1 5,0 5,5 P7
...... 4,4 4,4 4,5 4,8 4,5 4,7 4,5 4,5 5,0P9
20-07-o7D 92
-~ 20-07-olA 92 4,1 3,8 4,0 4,0 3,9 P4
~ 20-10-01B 68 4.1 4,0 4,1 4,1 4,2 P9
~ 20-10-02A 75 3,7 3,1 2,8 2,7 2,7 3,0 3,5CC
~ 20-10-02B 34 2,8 2,3 2,1 2,1 1,9 2,2 2,7 P7
...... 20-10-03Al 83 4,9 4,4 4,3 4,4 4,0 4,3 4,4 4,8P9
\0
~ 20-10-03A2 38 4,1 3,5 3,6 3,4 3,2 3,0 3,5 4,1 P9
20-l0-04A1 94 3.4 3,3 3,3 3,3 3,4P7
20-10-04A2 60 3,9 3,9 3,9 3,9 4,OP4
20-10-05E1 93 3,6 3,4 3,7 3,9 3,5 3,3 3,6 4,OP9
20-1O-o5E2 78 2,7 2,7 2,7 2,9 2,7 2,6 2,7 2,8CC
20-10-06B1 90 2,8 2,9 2,9 2,7 2,7 2,4 2,7 2,9 P9
20-10-06B2 80 2,7 3,1 2,8 3,0 2,8 2,9 2,9 2,8P4
20-10-07Bl 88 3,5 3,7 3,7 3,9 3,4 3,6 3,2 3,6 3,9P9
20-10-07B2 64 2,7 2,7 2,9 2,8 2,7 2,2 2,7 3,OP4
20-18-01J 95 4,5 4,6 4,5 4,6 4,6CC
20-18-02E 64 4,7 4,2 4,0 3,5 3,5 3,9 4,7P7
20-18-05K 89 4,4 4,7 4,8 4,8 4,6 4,5 4,6 4,9 P4
20-18-06D 89 4,3 4,6 4,4 4,9 4,5 4,6 4,5 4,8 P4
20-18-07A 87 4,6 4,7 4,6 5.1 4,6 5,0 4,4 4,7 4,9P9
20-26-0511 99 4,3 4,4 4,8 4,9 4,5 4,3 4,5 4,6FT
20-26-05J 95 3,8 3,7 3,9 3,8 3,6 3,7 3,8 3,9FT
20-26-06C1 96 4,1 4,3 4,3 4.3 4,2 4,1 4,2 3,7FT
20-26-06C4 92 3,4 3,5 3,5 3,4 3.4 3,5 3,5 3,7FT
20-26-07C1 90 4,2 4,2 4,3 4.5 4,2 4,1 4,6 4,3 4,2FT
20-26-07C4 85 3,3 3,2 3,7 3,3 2,7 3.2 2,9 3,2 3,5FT

Provenances are ordered from north (left) and south (right). Bold numbers indicate largest provenance mean in each test.
Test Codes: 20-02 corresponds to Carton de Colombia; 20-07 to SAPPI, South Africa; 20-10 to SAFCOL, South Africa; 20-18 to MONDI, South Africa; and 0-26 to Klabin, Brazil. All controls are P. patula unless
Nl
~ otherwise stated: CC = Commercial Control (South Mrica); P4 = M3004 (2nd generation); P9 = M2999 (2nd generation); P7 = M2997 (1st generation); FT = Pinus taeda.
sg TABLE 4. Mean individual tree volume (dm3) for Pinus patula assessed at 3 years ofage in Brazil, Colombia and South Africa

Test Survival Conrado Pinal Zacual- Potrero Llano T1acotla Ingenio Corralitla Santa Ixtlan Cuajimo- EI EI Test Control
Code % Castilo de tipan de de las del Maria loyas Tla- Man- Mean
Amoles Monroy Carmonas Rosario Papalo cuache zanal

2()..()2-C2C 83 13,05 14,20 11,71 12,35 11,96 12,97 11,16 CC


2()..()2-C3B 86 3,01 2,77 2,54 1,71 2,66 2,47 2,44 CC
2()..()2-C4B 94 3,13 3,14 3,65 3,29 2,95 CC
2o..07-CIL 96 7,42 6,86 6,86 7,06 5,31 P7
2()..()7-C2D 90 6,18 4,55 4,62 4,47 4,69 4,67 4,38 P9
2()..()7-C5L 94 4,66 4,97 5,84 6,09 5,01 4,52 5,03 6,59 P9
2()..()7-C6E 94 7,13 7,84 7,29 10,59 8,59 8,05 8,13 8,31 P7
2()..()7-C7D 92 5,41 5,81 5,70 7,00 5,81 6,25 5,70 5,95 5,83 P9
2o..07-CIA 92 2,67 2,44 27,8 2,65 1,96 P4
2o..l()"()IB 68 3,37 3,04 3,28 3,26 3,95 P9
2o..l()"()2A 75 1,78 1,18 0,83 0,89 0,84 1,01 1,48CC
2o..l()"()2B 34 0,54 0,34 0,22 0,17 0,18 0,30 0,63 P7
20..1()..()3Al 83 5,54 4,45 4,18 4,69 3,30 3,81 4,12 5,69 P9
2o..I()"()3A2 38 3,56 2,27 2,63 2,03 2,16 1,39 2,40 3,54 P9
2o..l()"()4Al 94 1,72 1,40 1,62 1,59 2,16 P7
2o..l()"()4A2 60 3,05 2,83 3,07 3,10 3,12 P4
2o..I0-C5El 93 2,03 1,67 1,90 2,50 1,72 1,46 1,84 2,54 P9
2o..Io..05E2 78 0,66 0,69 0,61 0,73 6,21 0,60 0,67 O,66CC
~ 2o..I0-o6Bl 90 0,86 0,97 0,82 0,66 0,71 0,48 0,73 0,78 P9
~
So 20..1()..()6B2 80 0,73 1,10 0,83 0,99 0,85 0,99 0,89 0,73 P4
20..1()"()7B1 88 2,08 2,31 2,11 2,78 1,59 1,91 1,54 1,93 2,47 P9
~ 20..1 ()"()7B2 64 0,73 0,89 0,81 0,79 0,71 0,51 0,78 0,98 P4
Sl~
20..18-011 95 6,32 6,41 6,14 6,28 6,20CC
~ 2o..l8-02E 64 6,57 4,70 4,20 2,45 3,12 4,28 7,97 P7
1Jl
3 2o..l8-05K 89 5,99 7,15 7,23 7,49 5,82 6,50 6,61 8,32 P4
~ 2o..l8-06D 89 5,13 6,53 5,34 7,45 5,76 6,07 6,08 7,67 P4
; 2o..l8-07A 87 7,05 7,87 7,06 9,66 7,41 8,80 6,87 7,71 8,94P9
\:l 5,31
.... 20..26-0511 99 5,63 6,88 7,66 8,91 6,16 6,50 5,35 PT
I
20..26-051 95 3,10 3,28 3,70 3,53 3,05 3,37 3,29 4,91 PT
~ 2o..26-06CI 96 3,90 4,94 4,61 5,01 4,32 3,76 4,27 3,57PT
..... 1,94 2,12 1,95 1,99 1,95 2,15 2,06 2,73PT
2o..26-06C4 92
~
2o..26-07Cl 90 4,20 4,23 4,83 5,49 4,05 4,03 6,51 4,61 5,99PT
~ 2o..26-07C4 85 1,83 1,63 2,43 1,75 1,42 1,76 1,61 1,72 2,84PT

~..... Provenances are ordered from north (left) and south (right). Bold numbers indicate largest provenance mean in each test.
10 Test Codes: 2()"()2 corresponds to Carton de Colombia; 2()"()7 to SAPPI, South Africa; 20-10 to SAFCOL, South Africa; 20..18 to MONDI, South Africa; and 0-26 to Klabin, Brazil. All controls are P. patula unless
~ otherwise stated: CC = Commercial Control (South Africa); P4 = M3004 (2nd generation); P9 = M2999 (2nd generation); P7 = M2997 (1st generation); PT = Pinus taeda.
TABLE 5. Percentage offamiliesperformingequaltoorbetter than themostproductivecontrolin thetest (H), a 1stgeneration control
(P7) and a 2nd generation (P4) controlin differentPinus patula tests in Brazil, Colombia and South Africa

Test ID H P7 P4

KLABIN BRAZIL
20-26-0511 26PT
20-26-05J 13PT
20-16-06Cl 29YT
20-26-06C4 OPT
20-26-07Cl 8PT
20-26-07C4 OPT

CARTON DE COLOMBIA COLUMBIA


20-02-02C 70CC
20-02-03B 25CC
20-02-04B 66CC

MONDI FORESTS SOUTH AFRICA


20-18-01J 84CC 26 47
20-18-02E 2 P7 2 5
20-18-05K 10P4 22 10
20-18-06D 3 P4 21 3
20-18-07A 12 P9 33 12

SAFCOL SOUTH AFRICA


20-10-01A 64P4 64 64
2Q-IQ-OlB 19 P9 71 71
20-1Q-02A 3CC 8 3
20-1Q-02B 3P7 3 26
20-1Q-03Al 2P9 10 6
20-1Q-03A2 OP9 10 5
20-1Q-04Al 40P7 40 44
20-1Q-04A2 23P4 96 23
20-1Q-05E1 3 P9 15 17
20-1Q-05E2 18CC 53 41
2Q-IQ-06Bl 21 P9 69 69
20-1Q-06B2 42 P4 80 42
20-1Q-07Bl 11 P9 20 17
20-1Q-07B2 4P4 60 4

SAPPI FORESTS SOUTH AFRICA


20-07-01L 27P7 27 95
20-07-02D 6P9 13 6
20-07-05L 5 P9 19 7
20-07-06E 3 P7 3 16
20-07-07D OP9 6 0

TABLE 6A. Mean height (m)for Pinus patula prooenanes in the across-site analysisat three yearsof agel. Forty-fivefamilies were
common to the six tests

Provenance Maxwell Tweefontein Jessievale Commonage Imbauzinho Marica


S.Mrica S.Mrica S.Mrica S. Arica Brazil Brazil
(05L) (05El) (05E2) (05K) (05IL) (05J)

Conrado 4,26a 3,58ab 2,67a 4,33a 4,31a 3,79ab


Casillo
Pinal de 4,29a 3,37b 2,65a 4,63b 4,41a 3,69b
Amoles
Zacualtipan 4,72b 3,65a 2,71b 4,73b 4,78b 3,87a

Llano de las 4,46a 3,50b 2,72b 4,39a 4,4a 3,64b


Carmonas

Suid-AfrikaanseBosboutydskrif- nr. 174, November 1995 29


TABLE 6B. Mean individual treevolume (dm 3) for Pinus patula provenances in the across-siteanalysis at threeyears ofage', Forty-
five families were common to the six tests

Provenanc Maxwell Tweenfontein Jessievale Commonage Imbauzinho Marica


S. Africa S. Africa S.Mrica S. Africa Brazil Brazil
(05L) (05E1) (05E2) (05K) (0511) (05J)

Conrado 4,66b 2,02a 0,65a 5,99b 5,63a 3,10b


Castillo
Pinal de 4,97ab 1,67b 0,72a 7,15a 6,88a 3,27ab
Amoles
Zacualtipan 5,84a 1,90ab 0,62a 7,23a 7,66b 3,70a
Llano de las 5,00ab 1,71ab 0,63a 5,82b 6,16a 3,05b
Carmonas
I
Means in a column with the same letter are not significantly different.
TABLE 7. Family volume means (dm 3) , ecovalences and p-valuesforPinus patula

Proenance/ Mean Ecovalence Ecovalence PoF


Family Absolute %
Pinal de Amoles
199 3,17 49,38 0,72 0,5147
202 3,37 89,66 1,32 0,1773
205 3,02 106,75 1,57 0,1062
208 3,01 71,85 1,05 0,2928
209 2,83 119,63 1,76 0,0710
210 2,43 383,59 5,64 0,0001
211 3,33 100,14 1,47 0,1299
212 3,13 161,36 2,37 0,0181
214 3,77 502,85 7,39 0,0001
217 3,14 170,62 2,51 0,0132
218 3,34 49,54 0,73 0,5129
219 3,13 170,88 2,51 0,0131
220 2,87 21,77 0,32 0,8629
222 3,64 181,37 2,67 0,0091
Conrado Castillo
225 3,71 191,63 2,82 0,0064
227 3,74 146,79 2,16 0,0294
228 2,66 335,16 4,93 0,0001
230 3,19 88,01 1,29 0,1860
231 2,81 44,61 0,65 0,5725
232 2,46 491,43 7,22 0,0001
233 2,91 56,93 0,84 0,4303
234 2,14 259,23 3,81 0,0006
237 2,63 129,40 1,90 0,0520
238 2,77 53,19 0,78 0,4710
239 2,47 97,35 1,43 0,1412
240 2,54 142,30 2,09 0,0341
241 3,56 108,37 1,59 0,1010
242 2,30 236,99 3,48 0,0013
243 2,93 38,26 0,56 0,6534
245 2,53 160,26 2,35 0,0188
Zacualtipan
251 4,06 393,58 5,78 0,0001
252 3,41 85,81 1,26 0,1982
254 3,63 216,60 3,18 0,0026
256 3,92 267,44 3,93 0,0004
257 2,49 170,25 2,50 0,0134
258 3,53 131,62 1,93 0,0484
261 2,84 147,96 2,17 0,0283
262 3,79 52,92 0,78 0,4740
Llano de las Carmonas
264 2,66 115,45 1,69 0,0816
270 2,81 34,68 0,51 0,7002
272 2,80 154,01 2,26 0,0231
280 3,69 43,34 0,64 0,5884
284 3,39 51,63 0,76 0,4886
285 2,25 61,81 0,91 0,3810
286 2,88 114,12 1,68 0,0845

30 South African ForestryJournal- No. 174 November 1995


lQ-03A2jessievale, South Africa; 20-26-06C4 and 20-26- tion of particular interest is whether these southern
07C4, Campina Grande, Brazil) the local controls were sources of ''variant'' P. patulawill have better tolerance to
superior to CAMCORE introductions. Several families stem breakage than P. tecunumanii (Dvorak et al., 1993;
from the Potrero de Monroy provenance were especially Parfitt and van der Sijde, 1993). It were these same
outstanding. Oaxacan sources with atypical morphology that were
Significant provenance x site and family x site interac- extremely susceptible to cold in the CAMCORE tests.
tions were found in the across-site analysis (Tables 6a Correlations between height or volume in test
and 6b). Despite the large environmental differences plantings and latitude of the collection site were weak
among sites, 25 of the 51 families were stable and did not when all 13 provenances were used in the analysis be-
contribute significantly to the interaction (Table 7). All cause sources at the species northern and southern
families in the Llano Las Carmonas, Puebla provenance geographic extremes performed more poorly relative to
showed no interactive tendencies. those in Hidalgo, Puebla, and Veracruz in central Mexico
thus masking any range-wide trends. The correlations
DISCUSSION improved once the extreme northern and southern
populations were excluded. The significant negative
The preliminary results on provenance performance correlation obtained in using the sources of P. patula
support findings of Falkenhagen (1979) who assessed suggests that the further south one goes within the
the performance of four P. patula provenances at seven central part of the country the better the productivity.
years of age in South Africa that were collected by The implication is that latitude may be a good indicator
Coetzee and Fisk. He found that sources from Puebla of provenance performance for central Mexico sources
and Hidalgo in central Mexico were superior to those of P. patula planted in South Africa, but not northern or
from more southern (Oaxaca) areas ofthe species range southern Mexico sources. However, the use of latitude
on most locations that the trials were planted in South of the collection site in central Mexico as a predictor of
Africa, Sources from Puebla also did well at Mac-Mac, performance in South Africa should be viewed with
South Africa that were included in Barrett's original some caution. Potrero de Monroy (Veracruz) performed
introduction (Eric Kietzka, personal communication, much better in South Africa relative to the other prov-
1995). Our results are also consistent with provenance enances from central Mexico, and therefore, its growth
performance of Barrett's collections established in Ar- was not well correlated to latitude of the collection site.
gentina between 1400 to 2100 m elevation and assessed It was so much an outlier that when included in the
at 11 to 14 years that were reported on by Picchi (1988). correlation analysis for central Mexican provenances,
He found that the provenances ofZacualtipan, (Hidalgo), the correlation dropped from -0,85 to -0,50 and was not
Pinal de Amoles, (Queretaro), and Zacatlan, (Puebla) significant.
(not collected by CAMCORE) were good across all sites. Three-year assessments have generally provided reli-
However, the best source in Barrett's studies in jujuy able information about what provenances will be poor
state was EI Cielo, Tamaulipas. This source has also been performers at half-rotation age when final selections are
sampled by CAMCORE but not included in this series of made in the CAMCORE program. These early results do
tests. The other Tamaulipas provenance that was in- not necessarily indicate which of the remaining sources
cluded, Conrado Castillo, showed average height and are best. However, major rank changes after three years
volume growth but with an occasionally outstanding of age in provenance performance are unlikely in the
family. Interestingly, the needle morphology and branch- future. Picchi (1988) reported few important rank
ing pattern exhibited by trees in the Conrado Castillo changes for P. patula diameters between ages 2 and 11
source were noticeably different from other P. patula years in jujuy, Argentina. Because of statistically differ-
provenances. Needles from this source were thicker and ent height-diameter ratios across sites, volume and taper
stiffer than typical P. patula and the branching pattern equations will have to be developed for each site to
more narrow and compact. better estimate productivity in future assessmen ts, There
Barrett's studies (Picchi, 1988) like Coetzee's and were no significant correlations between height-diam-
Fisk's (Falkenhagen, 1979) also indicated that the eter ratios and the elevation and latitude ofplanting site
Oaxacan sources were generally inferior to other sources. in this study. At present we have no explanation for this
However, in more tropical climates in Colombia the phenomenon. Observations of changes in height-diam-
southern and central Oaxacan sources (Manzanal, eter ratio of P. tecunumanii and P. oocarpain CAMCORE
Tlacuache, and Cuajimoloyas) performed well. These studies in Brazil by the senior author were thought to be
same sources have also done well in the mild subtropical related to altitudinal effects. When planted below their
climates of Swaziland (Kanzler, 1994). When studied in normal elevational range, trees had a tendency to de-
natural stands in Mexico, these three provenances were velop more rapidly in shoot height at the expense of
found to have internal and external foliage and cone diameter increment.
morphology intermediate between typical P. patula and The provenances included in the across-site analysis
P. tecunumanii and were classified as "variant forms" of P. were planted on locations that varied in elevation by
patula because taxonomically they grouped with neither 1000 m and rainfall by 700 mm per annum. Some
species (Dvorak and Raymond 1991). Their taxonomy changes in rank across site were expected. Furthermore,
is now being further studied at the genomic level at a severe drought in southern Africa in 1992 may have
North Carolina State University. A more applied ques- had a confounding effect on results, altering growth

Suid-Afrikaanse Bosboutydskrif - nr. 174, November 1995 31


more on the sandier, drier sites than on the moister Tamaulipas contains remnan t populations ofboth tropi-
clayey sites. However, the changes in rank observed at cal and temperate flora (Hernandez et aL, 1951) that
the provenance level represent only small differences may hold much biodiversity. It is presumed that more P.
and are not of great biological significance. patulastands exist in the area. Most interestingly, records
Families changed rank in height and volume over indicate that a botanical sample of P. patula was col-
short geographic distances within South Africa in the lected 10 km south of Monterrey (Nuevo Leon) at
CAMCORE trials. Family x site interactions for P. patula Canon del Diente (260 N Iat.) in 1939'. If the stand still
have been observed in other studies. Falkenhagen (1979) exists, this population would be 190 km further north
also saw significant family x site interactions in P. patula than Conrado Castillo, the northernmost known stand
at seven years of age on several sites in South Africa. sampled by CAMCORE. Cooperative staff are now ex-
Barnes et al., (1992) found that family x site interactions ploring this region to see if they can locate any P. patula
assessed for volume at 8 years of age across three sites in stands in the area. Also of particular interest and a top
Zimbabwe were of practical significance. There may be priority for the CAMCORE program are explorations in
several explanations for the interactions in the the remote areas of south-western and western Mexico
CAMCORE studies that are not related to genetic causes. in Guerrero, Michoacan andJalisco where "patula-like"
First, as mentioned previously, the 1992 drought may populations have been reported (Donahue, 1990).
have accentuated rank changes. Second, site prepara-
tion may have not been the same at all locations. Third,
weed control may not have been consistent across all CONCLUSIONS
tests and contributed to the magnitude of the interac-
tions. The CAMCORE studies demonstrate the great value for
One of the most promising aspects of these compre- continued explorations and seed collections in natural
hensive studies is the excellent performance ofa number stands ofpines in Mexico and Central America. The best
of the unimproved CAMCORE families relative to the provenances of 10 years ago are not necessarily the best
improved controls which came from bulked seed or- ones today. Provenances of P. patula from central
chard lots. For example, in nine of the South African Mexico like Potrero de Monroy, Zacualtipan, and Pinal
tests, 20% or more of the CAMCORE families were de Amoles show much promise in the tropics and sub-
superior in height growth to the second generation tropics. Although the far northern source from
control M3004. It appears that some of the best selec- Tamaulipas state, Conrado Castillo, is average in growth
tions from the CAMCORE material may be incorpo- it will contribute several outstanding families to the base
rated directly into advanced second generation breed- breeding population. Many individual families from
ing programs without any loss of genetic gain for pro- selections in natural populations in Mexico were supe-
ductivity. However, it is apparent that the unimproved rior in growth to advanced generation local controls.
introductions are generally inferior for stem form rela- Even though more time is needed to determine if this
tive to advanced generation selections in South Africa. superiority will be maintained, initial performance is
Another positive and exciting aspect of the CAMCORE exceptionally promising. The success ofthe CAMCORE
testing program is the opportunity to cross these new P. P. patula program has been due to the strong coopera-
patula introductions with the excellent selections of P. tive support received from forestry organizations on 4
tecunumanii or P. greggii to develop hybrids that grow continents and 14 countries.
well on sites not well suited for P. patula. Through the
cooperative activities, CAMCORE members have the
largest genetic base of the Mexican closed-cone pines in ACKNOWLEDGEMENTS
the world outside of their native ranges.
All CAMCORE tests will be remeasured at 5 and 8 We would like to thank Mr. Eric Kietzka (MONDI), Mr.
years of age and selection of plus trees will commence at William Ladrach (Zobel Forestry Associates), Mr. Terry
that time. Additional P. patula tests are being established Stanger, (SAPPI) Mr. Louis van der Merwe (SAFCOL)
with the remaining provenances in South Africa and and Dr. Jeffery Wright (Smurfit Carton de Colombia)
other countries. for their editorial comments on the manuscript. The
Based on these three-year results, additional seed authors would also like to thank the research staff of
collections are planned by the CAMCORE Cooperative Smurfit Carton de Colombia, Klabin (Brazil), and
in the center of the species geographical range in the MONDI, SAFCOL, SAPPI for establishing, maintaining
states ( C 4idalgo, Puebla, Queretaro and Veracruz to and measuring the CAMCORE trials. Furthermore, we
samp.t: genotypes with good growth for areas with sea- would like to thank the Mexican institutions Instituto
sonal frosts, The southern sources in Oaxaca should be Nacional de Investigaciones Forestales and Agrope-
targeted for further collections for the more tropical cuarias (INIFAP), Centro de Genetica Forestal (CGF)
areas. Populations on the fringes of the species geo- and Universidad Veracruzana for their assistance and
graphic rang:> should be sampled, possibly less inten- collaboration in the seed collections.
sively, to broaden adaptability and preserve rare genes.
Northern Mexi co in particular may offer unique oppor- 'Sample 2653, collected by C. H. Muller, Aug. 11, 1939, held at the
herbarium of the University of Texas, Austin, Texas USA. The
tunities for further conservation ofgene pools. The area sample was verified as P. patula by Dr. B. T. Styles, Oxford Forestry
surrounding the Rancho del Cie10 nature reserve in Institute, UK, Aug. 1986.

32 South African Forestry Journal- No. 174 November 1995


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Suid-Afrikaanse Bosboutydskrif- nT. 174, November 1995 33

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