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-
fied several new stands of P. patula while making explo-
rations and seed collections for other pine species in
southern Mexico and believed that additional
populations existed that had never been sampled before
by other agencies. At the urging of the South African
Forestry Research Institute (SAFRI) 2 which was a full
member of the cooperative program at that time,
CAMCORE initiated range-wide explorations and seed
collections of P. patula in Mexico in 1986 to better
protect dwindling gene pools of the species and broaden
the existing genetic base being used by forestry agencies.
From 1986 to 1994 seeds were obtained from 22 prov-
enances and 510 mother trees from Tamaulipas to
Oaxaca, a distance of approximately 900 km (Dvorak
and Donahue, 1992). The collections included a series
of new provenances from the Sierra Madre del Sur in
southern Oaxaca not sampled by either the South Afri-
cans or Argentineans or included in the FAa distribu-
tions. Furthermore, the CAMCORE effort included the
most northern source of P. patula yet found (Conrado
Castillo, Tamaulipas 24° 00' N Lat.) approximately 100
km north of Barrett's EI Cie10 site, as well as new sites in
the state of Veracruz (Table 1). At present, seeds for 80
P.patula CAMCORE conservation plantings and genetic
tests have been distributed to members in Brazil, Chile,
Colombia, Mexico, South Africa and Zimbabwe.
This paper reports on three-year provenance/prog-
enyresults of33 tests established in Brazil, Colombia and
South Africa with seeds from the CAMCORE collections.
Provenance x site and genotype x site interactions were
examined to determine their effect on the structure of
breeding populations. Recommendations are made on
efficacy of continued seed collections for P. patula in
Mexico.
MATERIAL AND METHODS
In the initial distribution, seeds from 13 provenances
and 282 mother trees were sent to members of the
CAMCORE Cooperative in Brazil (Klabin), Colombia
(Smurfit Carton de Colombia) and South Africa
(MaNDl, SAFCOL and SAPPI) and established in 33
provenance/progeny tests (Figure 1 and Table 2). The
number of provenances and families per provenance
included in each test varied from one cooperator to
another. Each member established provenance/prog-
eny tests using the standard CAMCORE compact family
design (Wright 1978) with nine replications and six-tree
row plots. Also included in each test was a series of
common control lots. CAMCORE included 3 open-
pollinated families from the Potrero de Monroy,
Veracruz, provenance in 32 of the 33 tests. In the 24 tests
in South Africa, CAMCORE members also included 5
common controls of commercial and advanced genera-
tion P. patula and P. elliottii. In Brazil, Klabin included
two controls of commercial P. taeda and P. elliottii from
local seed stands. The best performing control lot in
each test is reported here.
Total shoot height (m) and diameter at breast height
(cm) were measured at three years of age and volume
was calculated using a formula for juvenile trees
0,00003d2h (Ladrach, 1986). A preliminary analysis of
our data showed that height-diameter ratios varied sig-
nificantly across sites. Therefore, even though the trials
were young, both height and volume were used in the
analyses ofprovenance and family productivity to obtain
• SAFRI membership eventually passed to the Department of Water
Affairs and Forestry (DWAF) and then to the South African
Forestry Company Ltd. (SAFCOL). MONDI and SAPPI became
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
basis in order to facilitate the computational perform-
ance.
To determine the practical importance of rank
changes of provenances and families across environ-
ments, stability analyses were conducted by computing
ecovalences using a modified SAS program from
Fernandez (1991). An ecovalence is simply the sum of
squares contribution of a genotype to the total sum of
squares. Significance tests were conducted on each prov-
enance and family to determine if their response paral-
leled the mean response of all genotypes in the trial (see
Lin et al., 1986). A stable genotype by this definition
implies stability only with respect to the other genotypes
in the test and can not be generalised (Lin and Binns,
1991). Since the number of individual trees represent-
ing the control lots was much less than those represent-
ing provenances and families across sites, control lot
data was excluded from multiple means and ecovalence
comparisons.
Because future seed collections of P. patula were
planned by the CAMCORE Cooperative, the mean height
and volume of provenances at three years of age were
correlated with the latitude and altitude of the collec-
tion site in Mexico to determine if either would serve as
a good predictor ofseed source performance. Five of the
33 tests were used in the correlation analyses. The tests
chosen (OIL, 02D, 05L, 06E, and 07D) were established
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-
enous, replication means for all five tests were standard-
ized using the common controls before the correlation
analyses were run.
RESULTS
Survival of the tests ranged from 34% to 96% and
averaged 83% (Table 3). Two provenances from south-
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,
Jessievale and Commonage, South Africa, where freez-
ing temperatures are common during winter, the aver-
age survival of these three Oaxacan sources was 26%
versus 70% for the other remaining provenances. At
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
exceeded 5 m in tests at Los Guadales, Colombia (test
02C) and Maxwell, South Africa (test 06E). The poorest
growth was on dry and cold sites with sandy soils at
J essievale, South Africa (test 02B). The best provenances
in South Africa and Brazil were Potrero de Monroy,
Zacualtipan and Pinal de Amoles from the central part
ofthe P. patularange in Veracruz, Hidalgo and Queretaro.
In more tropical areas like the Andean highlands of
26
Colombia, the southern Oaxaca sources of P. patula
showed as much promise as the best central Mexico
sources. Provenance differences were statistically signifi-
cant in 23 of 33 tests for height and 26 of 33 tests for
volume.
The correlation between height or volume of trees at
Maxwell, South Africa and latitude of the collection site
in Mexico was not significant (r=-0,42). There was also
no significant correlation between growth or productiv-
ity and elevation of the collection site (r= 0,01). How-
ever, when only six provenances from central Mexico
were included in the analysis (Pinal de Amoles,
Zacualtipan, Llano Carmonas, Tlacotla, Ingenio Rosario
and Corralitla), the correlation between height or vol-
ume with latitude was -0,85 and -0,88, respectively and
highly significant (Figure 2).
In approximately 60% ofthe tests, the best CAMCORE
provenance was as good as or better in height/volume
than the most productive controls (Tables 3 and 4). On
average, 23% of the CAMCORE unimproved families
performed as well as or better in height growth than
advanced generation controls in 29 trials (Table 5). In
the remaining four trials (20-Q7-07D, Maxwell, and 20-
Volume, dm-
7..5 r - - - - - - - - - - - - - - - - - - - - - - - ,
• Potrero
de
r =- 0.88 Monroy
7,0 Ingenio
del
Roaario
Corrallta
Llano de
laa Carmonaa
Tlacotla
Pinal de
Amolea
Zacualtlpan
5,5 '------''--..--J._----'-_--'-_--'--_-'-_-'--_l....-----'_..--J._---J
18~5 19..0 20..0 20,5 21,0
Latitude N
FIGURE 2. Correlation between mean volume peiformance of
Pinus patula provenances at Maxwell, South Africa, and the
latitude ofseedcollectionsites in Mexico. The Potrerode Monroy
proenance was not used in the analysis.
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-
07C4, Campina Grande, Brazil) the local controls were
superior to CAMCORE introductions. Several families
from the Potrero de Monroy provenance were especially
outstanding.
Significant provenance x site and family x site interac-
tions were found in the across-site analysis (Tables 6a
and 6b). Despite the large environmental differences
among sites, 25 of the 51 families were stable and did not
contribute significantly to the interaction (Table 7). All
families in the Llano Las Carmonas, Puebla provenance
showed no interactive tendencies.
DISCUSSION
The preliminary results on provenance performance
support findings of Falkenhagen (1979) who assessed
the performance of four P. patula provenances at seven
years of age in South Africa that were collected by
Coetzee and Fisk. He found that sources from Puebla
and Hidalgo in central Mexico were superior to those
from more southern (Oaxaca) areas ofthe species range
on most locations that the trials were planted in South
Africa, Sources from Puebla also did well at Mac-Mac,
South Africa that were included in Barrett's original
introduction (Eric Kietzka, personal communication,
1995). Our results are also consistent with provenance
performance of Barrett's collections established in Ar-
gentina between 1400 to 2100 m elevation and assessed
at 11 to 14 years that were reported on by Picchi (1988).
He found that the provenances ofZacualtipan, (Hidalgo),
Pinal de Amoles, (Queretaro), and Zacatlan, (Puebla)
(not collected by CAMCORE) were good across all sites.
However, the best source in Barrett's studies in jujuy
state was EI Cielo, Tamaulipas. This source has also been
sampled by CAMCORE but not included in this series of
tests. The other Tamaulipas provenance that was in-
cluded, Conrado Castillo, showed average height and
volume growth but with an occasionally outstanding
family. Interestingly, the needle morphology and branch-
ing pattern exhibited by trees in the Conrado Castillo
source were noticeably different from other P. patula
provenances. Needles from this source were thicker and
stiffer than typical P. patula and the branching pattern
more narrow and compact.
Barrett's studies (Picchi, 1988) like Coetzee's and
Fisk's (Falkenhagen, 1979) also indicated that the
Oaxacan sources were generally inferior to other sources.
However, in more tropical climates in Colombia the
southern and central Oaxacan sources (Manzanal,
Tlacuache, and Cuajimoloyas) performed well. These
same sources have also done well in the mild subtropical
climates of Swaziland (Kanzler, 1994). When studied in
natural stands in Mexico, these three provenances were
found to have internal and external foliage and cone
morphology intermediate between typical P. patula and
P. tecunumanii and were classified as "variant forms" of P.
patula because taxonomically they grouped with neither
species (Dvorak and Raymond 1991). Their taxonomy
is now being further studied at the genomic level at
North Carolina State University. A more applied ques-
Suid-Afrikaanse Bosboutydskrif - nr. 174, November 1995
tion of particular interest is whether these southern
sources of ''variant'' P. patulawill have better tolerance to
stem breakage than P. tecunumanii (Dvorak et al., 1993;
Parfitt and van der Sijde, 1993). It were these same
Oaxacan sources with atypical morphology that were
extremely susceptible to cold in the CAMCORE tests.
Correlations between height or volume in test
plantings and latitude of the collection site were weak
when all 13 provenances were used in the analysis be-
cause sources at the species northern and southern
geographic extremes performed more poorly relative to
those in Hidalgo, Puebla, and Veracruz in central Mexico
thus masking any range-wide trends. The correlations
improved once the extreme northern and southern
populations were excluded. The significant negative
correlation obtained in using the sources of P. patula
suggests that the further south one goes within the
central part of the country the better the productivity.
The implication is that latitude may be a good indicator
of provenance performance for central Mexico sources
of P. patula planted in South Africa, but not northern or
southern Mexico sources. However, the use of latitude
of the collection site in central Mexico as a predictor of
performance in South Africa should be viewed with
some caution. Potrero de Monroy (Veracruz) performed
much better in South Africa relative to the other prov-
enances from central Mexico, and therefore, its growth
was not well correlated to latitude of the collection site.
It was so much an outlier that when included in the
correlation analysis for central Mexican provenances,
the correlation dropped from -0,85 to -0,50 and was not
significant.
Three-year assessments have generally provided reli-
able information about what provenances will be poor
performers at half-rotation age when final selections are
made in the CAMCORE program. These early results do
not necessarily indicate which of the remaining sources
are best. However, major rank changes after three years
of age in provenance performance are unlikely in the
future. Picchi (1988) reported few important rank
changes for P. patula diameters between ages 2 and 11
years in jujuy, Argentina. Because of statistically differ-
ent height-diameter ratios across sites, volume and taper
equations will have to be developed for each site to
better estimate productivity in future assessmen ts, There
were no significant correlations between height-diam-
eter ratios and the elevation and latitude ofplanting site
in this study. At present we have no explanation for this
phenomenon. Observations of changes in height-diam-
eter ratio of P. tecunumanii and P. oocarpain CAMCORE
studies in Brazil by the senior author were thought to be
related to altitudinal effects. When planted below their
normal elevational range, trees had a tendency to de-
velop more rapidly in shoot height at the expense of
diameter increment.
The provenances included in the across-site analysis
were planted on locations that varied in elevation by
1000 m and rainfall by 700 mm per annum. Some
changes in rank across site were expected. Furthermore,
a severe drought in southern Africa in 1992 may have
had a confounding effect on results, altering growth
31
more on the sandier, drier sites than on the moister
clayey sites. However, the changes in rank observed at
the provenance level represent only small differences
and are not of great biological significance.
Families changed rank in height and volume over
short geographic distances within South Africa in the
CAMCORE trials. Family x site interactions for P. patula
have been observed in other studies. Falkenhagen (1979)
also saw significant family x site interactions in P. patula
at seven years of age on several sites in South Africa.
Barnes et al., (1992) found that family x site interactions
assessed for volume at 8 years of age across three sites in
Zimbabwe were of practical significance. There may be
several explanations for the interactions in the
CAMCORE studies that are not related to genetic causes.
First, as mentioned previously, the 1992 drought may
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
tests and contributed to the magnitude of the interac-
tions.
One of the most promising aspects of these compre-
hensive studies is the excellent performance ofa number
of the unimproved CAMCORE families relative to the
improved controls which came from bulked seed or-
chard lots. For example, in nine of the South African
tests, 20% or more of the CAMCORE families were
superior in height growth to the second generation
control M3004. It appears that some of the best selec-
tions from the CAMCORE material may be incorpo-
rated directly into advanced second generation breed-
ing programs without any loss of genetic gain for pro-
ductivity. However, it is apparent that the unimproved
introductions are generally inferior for stem form rela-
tive to advanced generation selections in South Africa.
Another positive and exciting aspect of the CAMCORE
testing program is the opportunity to cross these new P.
patula introductions with the excellent selections of P.
tecunumanii or P. greggii to develop hybrids that grow
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
the world outside of their native ranges.
All CAMCORE tests will be remeasured at 5 and 8
years of age and selection of plus trees will commence at
that time. Additional P. patula tests are being established
with the remaining provenances in South Africa and
other countries.
Based on these three-year results, additional seed
collections are planned by the CAMCORE Cooperative
in the center of the species geographical range in the
states ( C 4idalgo, Puebla, Queretaro and Veracruz to
samp.t: genotypes with good growth for areas with sea-
sonal frosts, The southern sources in Oaxaca should be
targeted for further collections for the more tropical
areas. Populations on the fringes of the species geo-
graphic rang:> should be sampled, possibly less inten-
sively, to broaden adaptability and preserve rare genes.
Northern Mexi co in particular may offer unique oppor-
tunities for further conservation ofgene pools. The area
surrounding the Rancho del Cie10 nature reserve in
32
Tamaulipas contains remnan t populations ofboth tropi-
cal and temperate flora (Hernandez et aL, 1951) that
may hold much biodiversity. It is presumed that more P.
patulastands exist in the area. Most interestingly, records
indicate that a botanical sample of P. patula was col-
lected 10 km south of Monterrey (Nuevo Leon) at
Canon del Diente (260 N Iat.) in 1939'. If the stand still
exists, this population would be 190 km further north
than Conrado Castillo, the northernmost known stand
sampled by CAMCORE. Cooperative staff are now ex-
ploring this region to see if they can locate any P. patula
stands in the area. Also of particular interest and a top
priority for the CAMCORE program are explorations in
the remote areas of south-western and western Mexico
in Guerrero, Michoacan andJalisco where "patula-like"
populations have been reported (Donahue, 1990).
CONCLUSIONS
The CAMCORE studies demonstrate the great value for
continued explorations and seed collections in natural
stands ofpines in Mexico and Central America. The best
provenances of 10 years ago are not necessarily the best
ones today. Provenances of P. patula from central
Mexico like Potrero de Monroy, Zacualtipan, and Pinal
de Amoles show much promise in the tropics and sub-
tropics. Although the far northern source from
Tamaulipas state, Conrado Castillo, is average in growth
it will contribute several outstanding families to the base
breeding population. Many individual families from
selections in natural populations in Mexico were supe-
rior in growth to advanced generation local controls.
Even though more time is needed to determine if this
superiority will be maintained, initial performance is
exceptionally promising. The success ofthe CAMCORE
P. patula program has been due to the strong coopera-
tive support received from forestry organizations on 4
continents and 14 countries.
ACKNOWLEDGEMENTS
We would like to thank Mr. Eric Kietzka (MONDI), Mr.
William Ladrach (Zobel Forestry Associates), Mr. Terry
Stanger, (SAPPI) Mr. Louis van der Merwe (SAFCOL)
and Dr. Jeffery Wright (Smurfit Carton de Colombia)
for their editorial comments on the manuscript. The
authors would also like to thank the research staff of
Smurfit Carton de Colombia, Klabin (Brazil), and
MONDI, SAFCOL, SAPPI for establishing, maintaining
and measuring the CAMCORE trials. Furthermore, we
would like to thank the Mexican institutions Instituto
Nacional de Investigaciones Forestales and Agrope-
cuarias (INIFAP), Centro de Genetica Forestal (CGF)
and Universidad Veracruzana for their assistance and
collaboration in the seed collections.
'Sample 2653, collected by C. H. Muller, Aug. 11, 1939, held at the
herbarium of the University of Texas, Austin, Texas USA. The
sample was verified as P. patula by Dr. B. T. Styles, Oxford Forestry
Institute, UK, Aug. 1986.
South African Forestry Journal- No. 174 November 1995
UTERATURE CITED
BARNES,RD, MULLIN, L.]. and BATfLE G., 1992. Genetic control
of eighth-year traits in Pinus patula Schiede and Deppe. Silvae
Genetica 41 (6): 318-326.
BARREIT, W. H. G., 1972. Variacion de caracteres morfologicos en
poblaciones naturales de Pinus patula Schlecht. et Cham. en
Mexico. IDlA, Suplemento Forestal No.7. 9-35.
BARREIT, W. H. G., 1978. Some early observations on the variation
of height growth of PinuspatulaSchiede and Deppe. In: Proc.Joint
IUFRO Workshop, S2.02-08 & S2.03-01. Brisbane, Australia, 1977.
pp 722-723.
BIRKS,]. S. and BARNES, RD., 1991. Genetic control ofwood quality
in Pinus patula. Final Report. ODA Research Scheme R4616,
Oxford Forestry Institute, University of Oxford, UK. 29 pp.
BURGERS, T. F., 1975. Mexican origins of Pinus patula seeds
introduced in South Africa. Forestry in South Africa 16. pp. 31-43.
DARROW, W. K. and COETZEE, H., 1983. Potentially valuable
Mexican pines for the summer rainfall region of southern Africa,
South African Forestry [oumal; No. 124. pp. 23-36.
DONAHUE,]. K., 1990. Closed-cone pine explorations in the state of
Guerrero, Mexico. CAMCORE Technical Note No.4, College of
Forest Resources, North Carolina State University, Raleigh, NC.
USA. 1 p.
DUNNEIT, C. N., 1980. Pairwise multiple comparisons in the homog-
enous variance, unequal sample size case. Journal of the American
Statistical Association. 75: 789-795.
DVORAK,W. S. and DONAHUE,]. K., 1992. CAMCORE Cooperative
Research Review 1980-1992. College of Forest Resources, North
Carolina State University, USA. 93 pp.
DVORAK,W. S., LAMBETH, C. C. and LI, B., 1993. Genetic and site
effects on stem breakage in Pinus tecunumanii. New Forests7: 237-
253.
DVORAK, W. S. and RAYMOND, R. H., 1991. The taxonomic status
ofclosely related c1osed-cone pines in Mexico and CentralAmerica.
New Forests 4: 291-307.
FALKENHAGEN, E. R, 1979. Early assessment offour half-sib family
trials of Pinus patula (Schiede et Deppe in Sclechtendal et
Chamisso). South African ForestryJournalNo. 109. pp. 7-13.
FERNANDEZ, G. C.]., 1991. Analysis of genotype x environment
interactions by stability parameters. Hort. Science 26: 947-950.
FOREST GENETIC RESOURCES INFORMATION. 1982. Interna-
Suid-Afrikaanse Bosboutydskrif- nT. 174, November 1995
tional provenance trials of Pinus patula. No. 11. FAO, Rome, Italy.
54 pp.
HERNANDEZ, E, W. H., CRUM, H., FOX, W. B. and SHARP, A.].
1951. A unique vegetational area in Tamaulipas. Bulletin of the
Torrey Botanical Club. Vol. 78, No.6, pp. 458-463.
KANZLER,A. 1994. Interim report on trial 146: results from a thirty-
three month old Pinus patula provenance/progeny test. Forest
Research Document 6/94. Usutu Pulp Company, Forest Research,
Swaziland. 8 pp.
LADRACH, W. E. 1986. Comparaciones entre procedencias de siete
coniferas en las Zona Andina al finalizar los ocho mos. Smurfit
Carton de Colombia. Informe de Investigacion No. 105. 8 pp.
LIN, C. S. and BINNS, M. R. 1991. Genetic properties of four types of
stability parameters. Theor. Applied Genetics.-82: 505-509.
PARFIIT, E. and VAN DERSljDE,]. H. R 1993. Stem breakage of P.
tecunumanii in South Africa-A preliminary report. South African
ForestryJourna~ No. 167.51-53.
PICCHI, C. G. 1988. Comportamiento de origenes y progenies de
Pinus patula en el Departamento Capital, Provincia de Jujuy. En:
Convenio con la Dir. de Est. Experimentales. Instituto Forestal
Nacional, San Pablo de Reyes,Jujuy, Argentina. 12 pp.
SASINSTITUTE 1989. SAS/STAT User's Guide, Version 6, Fourth
Edition, Volume 2. Cary NC.p.891-996.
SOUTH AFRICA DEPARTMENT OF ENVIRONMENTAL AFFAIRS.
1993. Directorate National Forestry Planning and Extension.
Report on commercial timber resources and primary roundwood
processing in South Africa, Department ofEnvironmental Affairs,
Pretoria. 129 pp.
WALLER RA. and DUNCAN, D.B., 1969. A Bayes rule for symmetric
multiple comparison problems. Journal of the American Statistical
Association. 64: 1484-1499.
WORMALD, T.]., 1975. Pinus Patula. Tropical Forestry Papers No.
7. Oxford Forestry Institute, University of Oxford, UK. 172 pp.
WRIGHT, ]. A. 1994. Utilization of Pinus patula: An annotated
bibliography. O. F. I. Occasional Papers No. 45. Oxford Forestry
Institute, University of Oxford, UK. 73 pp.
WRIGHT,]. A. and LADRACH, W. E., 1994. Ten-year evaluation of
a field demonstration in Colombia ofpine, cypress and eucalyptus
transmission poles and fence posts treated with CCA. Forest Prod-
uctsJournal44( 4):69-71.
WRIGHT,J. W. 1978. A simplified design for combined provenance
and progeny testing. Silvae Genet. 27:68-70.
33