United States

Department of

Agriculture

Forest Service

Genecology of

Longleaf Pine in Georgia

and Florida

Southeastern Forest

Experiment Station John F. Kraus

Earl R. Sluder
The Authors:

John F. Kraus is retired Project Leader and

Earl R. Skier is Principal Plant Geneticist

with the Southeastern Forest Experiment Station

at Dry Branch, GA.

May 1990

Southeastern Forest Experiment Station

PO. Box 2680

Asheville, North Carolina 28802
Genecology of Longleaf Pi

in Georgia and Florida

ne

Contents

‘age

Introduction . . . . . . . . . . . . . . . . . . . . . . . . .l

Past Work . . . . . . . . . . . .2

Survival ...........................2

Growth ...........................2

Disease. ........... ...............3

Other Traits ......... ...............3

Materials and Methods . . . .*. . . . . . . . . . . . .4

Seed Collections . . . . . .

Nursery Procedures. . . . .

Planting Location and Desig

Field Data Collection . . . .

Data Analysis . . . . . . . .

.4

.4

.6

.6

.6

..............

..............

in .............

..............

. . . . , . . . . . . . . .

Results . . . . . . . . . . . . . .7

Seed Wings . . . . . . . . . ...............7

Nursery Observations . . . ...............7

Seedling Needle Length . . ...............7

First-Year Survival. . . . . . ...............8

Second-Year Results. . . . ...............9

Fifth-Year Results. . . . . . ..............9

Tenth-Year Results . . . . . ..............IO

Fifteenth-Year Results . . . ..............11

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . 12

Acknowledgments . . . . . . . . . . . . . . . . . . . . 16

Literature Cited . . . . . . . . . . . . . . . . . . . . . * 17

ABSTRACT

Fourteen seed sources of Pinus palustris from Georgia, five from

Florida, and one from Alabama were grown at five locations in

Georgia and at two in Florida. Data collected through the 15th

year in the field show: (1) survival and early height growth were

best among the northern sources, possibly related to introgression

with P. faeda; (2) individual stand or seed source contributed

strongly to the components of variation, which could be due to

stand history; i.e.; past logging practices; (3) significant seed source

x plantation interaction for growth traits, interaction which possibly

could be useful in selecting a suitable seed source for a particular

site; and (4) a generally broad zone of adaptability in Georgia

precludes the necessity for conducting more than one breeding

program for longleaf pine for Georgia at this time.

Keywords: Pinus pa/ustris, geographic variation, survival and growth

traits, adaptability.

Introduction

Longleaf pine (Pinus palusrris Mill.) constituted much

of the pine forest in southeastern North America

when European settlers arrived. Extensive lumbering,

destructive fires, and difficulties with artificial regeneration have-resulted in conversion of much of the forest

acreage once occupied by longleaf pine to other

species, such as loblolly pine (P. raeda L.) and slash

pine (P. elliorrii Engelm.). Many years of forest

management experience have revealed, however,

that longleaf pine is better adapted to grow on deep

sand sites formerly occupied by the species than

are loblolly and slash pines. For this reason, plus the

highly favorable timber and wood qualities of longleaf

pine, commercial forest-land owners are showing

increasing interest in management of the species.

The range of longleaf pine, which extends from

southeastern Virginia to eastern Texas, includes over

two-thirds of the State of Georgia and more than half

of Florida (fig. 1). Approximately 40 percent of the

rangewide growing stock is found in these two States.

Longleaf pine is of major commercial importance

from the Piedmont-Coastal Plain boundary in Georgia

south to central Florida. Its commercial importance

requires that forest-land owners learn how to plant

and manage it successfully. A prerequisite to its

successful management is a knowledge of the species’

genetic variation.

The objective of this study was to determine the

pattern of racial variation in longleaf pine in Georgia,

and to supplement the Southwide Pine Seed Source

Study by including sources from a north-south transect

in Florida to as near the southern limit of the species’

%

250 --

I

Figure 1 -The natural range of longleaf pine.
range as possible. This study provides the muchneeded information previously lacking for this very

important portion of the range of longleaf pine.

In 1967, when this research was begun, only two

longleaf pine racial variation plantations were located

in Georgia. Both of these were in Dooly County and

were part of the Southwide Pine Seed Source Study’.

Each plantation contained only one Georgia seed

source, from Treutlen County. All the other known

studies containing Georgia sources of longleaf pine

were located outside of the State.

Past Work

Survival

Published reports from longleaf pine racial variation

studies have generally shown no strong patterns of

variation associated with seed source in planting

survival of seedlings, Significant differences in

individual plantations have been to some extent

associated with the performance of one or two seed

sources, usually from the extremes of the species’

range. A Hillsborough County, FL, seed source, for

example, had exceptionally poor survival in several

different plantations in the Southwide Study (Allen

1961; Shoulders 1965; Wakeley 1959, 1961, 1963;

Wells 1969; Wells and Wakeley 1970). At the other

extreme, in a longleaf pine seed source plantation in

Nansemond County, VA, the local seed source had

the best survival after 6 years (Allen 1961).

Survival differences among the other seed sources

in these Southwide Study plantations generally

appeared to be random (Bethune and Roth 1960;

Collins 1964; Harms 1967; Wells and Snyder 1976).

There were, however, significant differences in survival

among 10 seed sources planted in Rapides Parish,

LA. These differences proved to have a significant

relationship with mean annual temperature of the

seed source, indicating that it might be safer to move

seed from cooler to warmer climates than the reverse

(Shoulders 1965; Shoulders and Terry 1968).

1 Working plan for cooperative study of geographic sources of

southern pine seed. Committee on Southern Forest Tree Improvement, Subcommittee on Geographic Source of Seed, Philip C.

Wakeley, Chairman. Southern Forest Experiment Station, New

Orleans, IA. 35 pp. illus. 1952.

In some cases, differences in planting survival among

seed sources may be associated with cultural

practices in the nursery. In other cases, the mortality

is an indication of inherent differences in resistance

to brown-spot needle blight, caused by Scirrhia acicola

(Dearn.) Sigg. (Snyder and Allen 1968).

Growth

The lo- and 15-year results from the Southwide Pine

Seed Source Study show that growth of seed sources

from the central Gulf Coast Region has been better

than that of other sources on most sites, ranging

from central Louisiana to central Georgia and north

to the Alabama Piedmont. Local sources grew best

in areas northeast of central Georgia and west of

central Louisiana (Collins 1964; Wells 1969, 1975;

Wells and Wakeley 1970). The seed source from

Hillsborough County, FL, was consistently slow

growing in two plantings, one in Virginia (Allen 1961)

and one in central Louisiana (Shoulders and Terry

1968).

In a study in South Carolina, after 10 years there

were no significant differences in growth among

eight seed sources from the Sandhills and Lower

Coastal Plain planted in the Sandhills in 3 consecutive

years (Harms 1967). When six coastal Mississippi

and six mountain Alabama sources were compared

in reciprocal plantings on sites in Mississippi and

Alabama, the local sources had the best volume

growth after 10 years (Snyder and Allen 1968). After

12 years in a south Mississippi planting, the familywithin-area differences were greater than the geographic differences for height, diameter at breast

height, and volume growth of longleaf pine progenies

from three trees in each of 20 areas in south Georgia,

2 in Alabama, and 1 each in Mississippi and Louisiana

(Wells and Snyder 1976).

A longleaf pine seedling seed orchard in central

Florida contained 34 families after roguing at 4 years.

Fifteen of these families (44 percent) were from the

Gulf Coast “optimum zone.” Only eight families from

the optimum zone were completely or heavily rogued,

The orchard originally contained 65 families. Indications were that the optimum zone should include

southeast Georgia and north Florida (Rockwood and

Kok 1977).

2
Disease

Most of the longleaf pine plantations in the Southwide

Pine Seed Source Study were sprayed during their

early years with Bordeaux mixture to prevent infection

by the brown-spot needle blight fungus. Four plantings

established in southern Mississippi with 16 seed

sources were left unsprayed to test for geographic

variation in resistance to that disease. After 5 years,

there were clear-cut differences: sources from the

western and eastern extremities of the range were

more heavily infected than those from the central

Gulf Coast. Even in the plantations that were sprayed

to protect them from brown spot, seed sources from

central Louisiana and east Texas were more heavily

infected than those from the central part of the range

(Bethune and Roth 1960; Wells 1969). Among the

seed sources from east of the Gulf Coast optimum

zone, there was no pattern to the variation in disease

resistance (Henry and Wells 1967). In the reciprocal

plantings of coastal Mississippi and Alabama mountain

sources, the Alabama mountain sources were

significantfy more heavily infected by brown spot

than the Mississippi sources, probably because

there has been little natural selection for resistance

to this disease in north-central Alabama where its

incidence is low (Snyder and Allen 1968).

Two tests of wind-pollinated progenies from more

than 750 individual-tree selections have produced

data showing geographic variation in resistance to

brown-spot needle blight. In one of the tests, which

included 227 selections, the most resistant progenies

were produced by parents from Alabama, South

Carolina, and Florida and the least resistant progenies

were from trees selected in Texas, Louisiana,and

North Carolina (Derr 1971). In the other test, 640

wind-pollinated longleaf pine progenies were grown

in two plantations in Gulfport, MS, and one in

Alexandria, LA. In those plantings, progenies from

southwestern Alabama were more resistant to

brown-spot needle blight than those from central or

southwest Georgia, north Florida, Mississippi, and

Louisiana (Snyder and Derr 1972).

Longleaf pine generally is considered to be relatively

resistant to the fusiform-rust fungus (Cronartium

quercuum f. sp. fusiforme). Data from three plantations

of the Southwide Pine Seed Source Study indicate

that sources from east Texas and central Louisiana

are more resistant to that disease than are sources

from Alabama and central Georgia (Wells and Wakeley

1970).

Other Traits

Although longleaf pine generally is considered to be

a straight, well-formed timber tree, it often shows a

tendency to fork in young plantations. In the Dooly

County, GA, planting of the Southwide Pine Seed

Source Study, forking showed significant differences

at 5 years but not at 10 years, with some indication

that the western sources from Louisiana and Texas

might have less tendency to fork than other sources

when grown in central Georgia (Bethune and Roth

1960; Collins 1964).

At two locations in southwest Alabama, inherent

geographic variation in the production of female

flowers was evident in lo-year-old longleaf pines

from 11 seed sources, ranging from Texas and west

Florida to North Carolina. Trees from a Richmond,

NC, seed source produced over 70 percent of the

female flowers counted at age 10 in the study (Boyer

and Evans 1967).

Variation in root morphology of longleaf pine seedlings

from 20 Georgia sources has been studied. Results

showed that seedlings from sources in eastern

Georgia had more fibrous root systems than those

from west of a line between Laurens and Echols

Counties (Snyder 1961).

Allen and McGregor (1962) detected no differences

in growth under long and short photo-periods among

longleaf pine seed sources in a greenhouse study.

The seed sources used did not cover a wide range

of latitudes, however, and since the study was

discontinued after 6 months, the results may not be

considered conclusive.

This review of the literature reveals a rather sketchy

picture of the geographic and local genetic variation

in longleaf pine. Particularly noticeable is the paucity

of information for Georgia. The purpose of this study

is to more completely fill in the picture for this very

significant portion of the range of the species.
Materials and Methods

Seed Collections

Seed collections in Georgia were by physiographic

province. Three collections were made in each of the

Flatwoods, Lower Coastal Plain, Upper Coastal Plain,

and Sandhills Provinces; two collections were made

in the western Piedmont Province. Five collections

were made in Florida to sample the latitudinal variation

of the species in that State. The Baldwin County, AL,

source was included for a further test of its performance. In total, there were 20 seed sources (fig. 2).

Cones were obtained at each location from at least

15 representative trees and from up to as many as

26 trees at one location. Cones were kept separate

by individual tree. Usually, 15 to 20 cones per tree

were collected.

The seeds were extracted by hand, kept separate by

individual tree, and stored at -7 “C until used. Samples

of winged seeds were saved, on which the seed

wing lengths and widths were measured. Data were

obtained from an average of 8 wings per tree from

20 trees in each of 16 seed sources and from 21

and 26 trees in two of the other sources.

For the major part of the study, bulk seedlots were

made up for each source. Except for the Pasco County,

FL, source, bulk seedlots were cornposited from 42

grams of seed from each of 20 individual trees per

seed source. Due to relatively low seed yields from

the Pasco County collection, almost all available

seed from those 25 trees was used in making up

that bulk seedlot.

Each of the bulk seedlots was divided into two parts.

The Florida Forest Service received one part, consisting of 225 grams per seed source, for sowing in the

Andrews Nursery near Chiefland, FL The Georgia

Forestry Commission received the other part of each

lot for sowing in the Morgan Nursery near Byron,

GA.

Nursery Procedures

The bulk lots from 20 seed sources were sown in

the Morgan Nursery in November 1968. Three

seedbeds that had been fumigated with methyl

bromide were used, with each seedbed serving as a

nursery replication. The seed sources were randomized within each replication, and each source was

broadcast-sown on a 15-foot section of nursery bed.

In addition to the bulk lots, seeds from 313 of the

individual-tree lots were sown in two replications. In

the first replication, which occupied one bed, each

individual-tree seedlot was sown in a single row

across a 4-foot bed, with the seed source locations

randomized and the individual-tree lots randomized

within locations. The second replication of the

individual-tree lots occupied two nursery beds; the

same randomization was used, but, instead of single

rows, the seeds of each seedlot were broadcast

over an area equivalent to two rows in the first

replication.

Watering, weeding, and nursery culture generally

followed standard nursery practices. In the late spring

of 1969, the bulk seedlots were hand-thinned to a

density of approximately 15 seedlings per square

foot.

In both bulk and individual-tree lots, seedlings having

normal green cotyledons but yellow primary foliage

were tallied and marked after germination was

complete in the spring. Root injury caused by a root

parasitic nematode, Tylenchorhynchus claytoni, was

severe in some seedlings. Badly injured seedlings

were discarded during lifting, but some mortality

following planting may have been due to this pest.

Sonderegger seedlings (the longleaf x loblolly F-l

hybrid) were tallied in September 1969 in the

individual-tree progenies. They were not discarded

during lifting and planting but were treated as part of

the population being studied.

In the fall of 1969, differences in seedling needle

length in the nursery were obvious. Consequently,

4
I I

,‘_ . -1 \ 2758@

----

*

Natural range of longleaf pine

Days in growing season

0

Seed Origins

Number

2753

2754

2755

2756

2757

2758

2759

2760

2761

2762

2763

2764

2765

2766

2767

2768

2769

3065

3066

3085

County State

Baldwin AL

Bryan GA

Colquitt GA

Jefferson GA

Jenkins GA

Meriwether GA

Paulding GA

Pulaski GA

Richmond GA

Taylor GA

Telfair GA

Terre11 GA

Ware GA

Washington GA

Wayne GA

Glades FL

Jackson FL

Marion FL

Taylor FL

Pasco FL

A

Plantations

Number County State

109 Polk GA

110 Crawford GA

111 Houston GA

112 Laurens GA

113 Ware GA

114 Baker FL

115 Lake FL

Figure 2-Location of longleaf pine seed sources and planting

locations.

5
needle-length data were collected by measuring 15

needles in each plot of the bulk seed-source lots

and 10 needles per plot of 10 individual-tree progenies

in each seed source. In addition, seedlings were

obtained in the spring of 1970 from the other sowing

at the Andrews Nursery, and needle length was

measured on an average of 20 seedlings per seedlot

from each of the three replications there.

Only seedlings from the Georgia nursery were used

for outplanting. Seedlings for each plantation were

lifted separately, a day or two before planting.

Seedlings from each seedlot were tied in plot-size

bundles, tagged, assembled into field replications,

and wrapped with wet moss in moisture-barrier paper.

Planting Locatlon and Deslgn

Seedlings from the bulked seedlots were outplanted

December-February 1969-l 970 at seven locations -

five in Georgia and two in Florida. In Georgia, one

planting was established in each of the five physiographic provinces. In Florida, the plantings were

located in the northern and central portions of the

species’ range (fig. 2). Each planting was a randomized complete-block design with six replications of

25-tree plots planted in squares of 5 x 5 trees. Spacing

between trees was 2.5 x 2.5 meters. A two-row isolation

border was planted around each plantation. There

were not enough seedlings of seedlot 3085 (Pasco

County, FL) to plant this source at all locations, so it

was used only in the Georgia Upper Coastal Plain

and central Florida plantings. The most nearly local

seed source was substituted for seedlot 3085 in the

six remaining plantings.

County (Upper Coastal Plain), and Laurens County

(Lower Coastal Plain), GA, and in Baker County,

northern Florida.

Data recorded for all plantations at 5 years after

planting were: survival, total height, number of years

out of the ‘grass stage’ (0.2 foot or taller) for the

trees that had begun height growth, and percentage

of brown-spot infection. The average annual height

growth rate was calculated for all trees that had

begun height growth.

At 10 years, total-tree height and diameter at breast

height were measured; fusiform-rust infections per

tree were counted; and trees having forks, ramicorn

branches, and broken tops were recorded. Average

annual height growth rate between the 5th and 10th

year; tree volume; and percentages of trees with

fusiform rust, forks, and ramicorn branches were

calculated. Field data collected at 15 years were on

the same traits as at 10 years.

Data Analysis

All percentage data were analyzed after transformation

to the arc sine ,/% Differences among seed source

means and among province means were tested for

significance by analysis of variance (ANOVA) and

Duncan’s multiple range test. Principal components

analyses were used to identify seed sources that

stood out from the general population.

Field Data Collection

The first year after planting, survival was counted at

all plantations. The second year after planting, the

number of living trees and the number of trees that

had begun height growth were counted in four of

the plantations: Crawford County (Sandhills), Houston

6
Results

Seed Wings

Harlow and Harrar (1958) in their dendrology textbook

described the seed wing of longleaf pine as ‘striped

and used such a wing in their illustration. In the

collections made for this study, trees with striped

seed wings occurred with a much lower frequency

than those having solid-colored wings (table 1). There

appeared to be no latitudinal or other pattern to the

variation in wing pigmentation.

ANOVA of the seed wing measurements showed

highly significant differences among individual trees

within source for seed wing length, width, and the

width/length ratio for all 18 seed sources evaluated

for the traits. ANOVA among sources showed highly

significant differences for the width/length ratio only,

and principal components analysis indicated that

most of the variation was due to the relatively short

seed wings of the Glades County, FL, source (fig. 3).

Nursery Bbservations

Xanthescens - Seedlings having normal green

cotyledons but yellow primary foliage (xanthescens)

were tallied in the spring after germination was

complete. There was no clear geographic pattern to

their occurrence (table 2). The relatively higher

0.8 -

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z

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p 0.4 -

In

; Glades County, FL

= .

c

z O5

E

:

3 -0.4 -

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frequency of xanthescens seedlings in the Taylor

and Washington Counties, GA, sources was probably

due to individual-tree carriers.

Hybrids-Although there was considerable variation

in their frequency within seedlots, Sonderegger

seedlings were found only in Georgia progenies

from the Piedmont, Sandhills, and Upper Coastal

Plain (table 3). Presumably,the possibilities for

hybridization of P. palustris with P. taeda are greater

in these areas of species range overlap combined

with possible overlap in flowering time (Boyer 1978).

Seedling Needle Length

ANOVA’s of all three sets of needle-length data

showed highly significant differences among seedsource means. The average needle lengths of

seedlings from the Piedmont, Sandhills, and Upper

Coastal Plain sources were shorter than those of the

Lower Coastal Plains, Flatwoods, and Florida sources

(table 4).

.

.

.

.

.

. .

.

.

.

. =

.

. .

.

.

.

-0.8’ I I I 1 I I 1

-2.0 -1.6 -1.2 -0.8 -0.4 0 0.4 0.8

Principal component I (58.1 percent)

Figure 3-Principal components analysis of seed wing measurements of longleaf pine.
Figure 4-Seedlot mean first-year

survival averaged over all

plantations and plotted as deviations

from the grand mean (percent).

--__ Natural range of longleaf pine

* Days in growing season

n Plbdmont

.

.

Al.b.m.-Florid. n

I . FI.twood.

37 36 39 40 41 42 43 44 45 46

Mean seedling needle length (cm)

Figure SRegression of mean first-year survival on mean seedling

needle length. A = mean of physiographic province.

First-Year Survival

Analysis of first-year survival showed the greatest

source of variation to be that among plantations.

Among seed sources, the best survival over ail

plantations was in sources from the Upper Coastal

Plain and northward (table 5). A small but highly

significant source x plantation interaction probably

was due to the poor survival of Lower Coastal Plain

sources in the plantation in central Florida. The

deviations of the individual seediots from the survival

grand mean are shown in figure 4, plotted at the

seed source locations; the minus deviations are

predominantly from the southern seed sources. The

regression of first-year survival on seedling needle

length was significant at the 5percent level of

probability (f=O.30; fig. 5).

8
Figure 6-Seedlot mean

second-year percentage survival

averaged over the plantations

in Crawford, Houston, and

Laurens Counties, GA,

and Baker County, FL.

240*

\ 63.29’-. 65.1 . \

-___ Natural range of longleaf pine

l Days in growing season

Second-Year Results

Survival at 2 years is summarized in table 6. Differences

among seedlots in the individual plantations were

highly significant except in the Baker County, FL,

planting. Combined analysis showed highly significant

differences among the seedlots averaged over all

plantations, among the means of the physiographic

provinces, and among the means of the individual

outplantings. As both table 6 and figure 6 indicate,

there is a definite tendency for seed sources from

the Piedmont, Sandhills, and Upper Coastal Plain to

survive better than seed sources from farther south.

The percentage of trees out of the ‘grass stage” 2

years after planting showed highly significant differences among the seedlots and significant differences

among the means of the physiographic provinces

when averaged over the four plantations in which

this trait was studied (table 7). Differences among

seedlots were significant within three of the four

plantations. The tendency for seedlots from the Upper

Coastal Plain, Sandhills, and Piedmont to have a

higher percentage of seedlings out of the ‘grass

stage” at this early age was rather distinct (fig. 7).

Fifth-Year Results

Fifth-year data summaries for survival, number of

years out of the ‘grass stage,’ total height, and height

growth rate are shown in tables 8-l 1. Differences

among outplanting locations were highly significant

for’all four traits, Variation among seedlots within

province was highly significant for all traits, while that

among province means was significant only for

survival.

For survival, there were highly significant differences

among the seedlots and among the means of the

physiographic provinces averaged over all plantations

(table 8). The same trend existing after the first and

second years was still evident: the Upper Coastal

Plain, Sandhills, and Piedmont sources had higher

survival than seed sources from farther south.

Most of the seed sources apparently came out of

the ‘grass stage’ during their third year in the field.

Based on averages over all plantations, the differences

among seed sources were highly significant, but the

differences among the means of physiographic

provinces were not significant (table 9).

9
_--_ Natural range of longleaf pine

* Days in growing season

Differences in total height among seed sources within

province were significant within three plantations

(table 10). When averaged over all plantations they

showed highly significant differences. Physiographic

province means, however, were not significantly

variable.

Apparently, annual height growth rate was fairly

uniform within a given plantation once the trees came

out of the ‘grass stage,’ as indicated by the lack of

statistically significant differences among the seed

sources at any of the individual plantations (table

11). Based on the means over all plantations, however,

differences among seed sources were highly significant but did not show any significant relationship

with physiographic province.

No significant interactions were noted between

plantations and physiographic provinces for any

traits at age 5 years, indicating that the relative

performance of the provinces was similar at the

different plantation locations. Interaction between

plantation and seed source within province for total

height and survival was highly significant, however.

,270*

Figure 7-Seedlot mean percentage

of trees out of the ‘grass stage’

2 years after planting averaged

over the plantations in Crawford,

Houston, and Laurens Counties, GA,

and Baker County, FL.

During the 5th-year assessment, 15 trees were tallied

as putative hybrids, based on needle length and

crown form; 13 of these were from Piedmont, Sandhills,

or Upper Coastal Plain seed sources and were

probably natural longleaf x loblolly crosses. Of the

other two hybrids, one was from Marion County and

the other from Glades County, FL. The hybrid from

Marion County could be longleaf x loblolly since

both species occur in that area, but the Glades County

hybrid is probably a natural cross between longleaf

pine and South Florida slash pine (P. elliottii var.

densa Liile & Dorman).

Tenth-Year Results

When the lOth-year data were combined over all

plantations, percentage of survival was the only trait

showing a significant difference among provinces

(table 12). The three northern provinces (Piedmont,

Sandhills, Upper Coastal Plain) all had better survival

10
than the overall mean. Based on the combined data,

differences among seed sources within province

were highly significant for survival, average height,

average annual height growth, average individual-tree

volume, and percentage of fusiform-rust infection.

Also significant were differences among sources for

percentage of trees with forks.

The tallest seed sources were from the Georgia

Lower Coastal Plain and northwest Florida. If survival

and average tree volume also were considered, the

best seed sources were those from the Upper Coastal

Plain of Georgia.

A principal components analysis based on survival,

height, height growth rate, diameter, and volume

over all plantations showed the most northern seed

source from Paulding County, GA, and the two most

southern seed sources from Pasco and Glades

Counties, FL, to be farthest removed from the general

grouping of the other seed sources (fig. 8).

In the individual plantations, variation in growth rate

was quite evident among the seed sources, with the

fastest growing sources at any one planting location

seldom including the local source. There was,

however, a slight indication of a north-south trend in

growth rate. The eight fastest growing seed sources

in the Piedmont plantation originated at an average

e 0.6

E

$ 0.4

ki

P

0.2

2

c 0

+Ik -0.2

z -0.4

2

0 -0.6 0

i5a -0.6

77 t

.

Pauldinb County, GA *

.

.

Glades County, FL

latitude of 32”58’N, while the average latitude of the

seven fastest growing seed sources in the central

Florida plantation was 30”51’N.

Fusiform-rust infection was relatively heavy in the

Upper and Lower Coastal Plain plantations, averaging

56 percent in the former and 12 percent in the latter.

Both locations showed highly significant differences

among the seed sources but not among physiographic

provinces (Kraus 1985). The three most southerly

sources from Marion, Pasco, and Glades Counties in

Florida had the lowest fusiform-rust infection.

Fffteenth-Year Results

The survival data combined from all the plantations

showed significant differences among the physiographic provinces of seed origin (table 13). The

Piedmont, Sandhills, and Upper Coastal Plain seed

sources had the best survival. Differences among

the individual seed sources were highly significant.

The seed source from Glades County, FL, was

significantly lower in survival than any of the others.

No significant differences were associated with the

physiographic province of seed origin for any of the

other traits.

Differences among individual seed sources for height,

average annual height growth, and average individualtree volume (table 13) were highly significant. The

.

.

.

.

Pasco County, FL

Lt -1.2 -0.8 -0.4 0 0.4 0.6 1.2

Principal component I (60.8 percent)

Figure 8-Principal components analysis of geographic sources of

longleaf pine based on lOth-year survival, height, height growth,

diameter, and volume over all plantations.

11
Pasco County, FL, seed source was significantly

taller than any of the others. Average annual height

growth was slowest in the Paulding and Meriwether

Counties, GA, sources and in the Glades County,

FL, source. The Paulding and Meriwether Counties

sources also had the lowest average individual-tree

volumes.

Fusiform-rust infection was severe in the Upper

Coastal Plain plantation, with an average of 70 percent

of the trees having stem and/or branch infections. It

also was a problem in the Lower Coastal Plain

plantation, with an average infection of 24 percent.

Over all plantations combined, there were highly

significant differences among the individual seed

sources in fusiform-rust infection. The seed sources

having the lowest rust infection were from Glades

and Pasco Counties, FL.

Significant interaction was observed between plantation location and physiographic province of seed

source for height, diameter breast height, and volume

(table 14). This interaction is illustrated by the

differential growth of the seed sources from the

Piedmont vs. those from Florida (fig. 9). Also, there

was highly significant interaction between plantations

and individual seed sources for survival, height,

average annual height growth, and volume (table

15). Again, the interaction was associated with the

differential response of the northern vs. the southern

seed sources when planted at the northern and

southern extremes of the species’ range.

The occurrence of forking also showed significant

interaction between plantation and source province.

As at 10 years, seed sources from the northern

provinces tended to have the highest percentage of

forking overall, but the interaction occurred in the

sources from the Upper Coastal Plain, the Flatwoods,

and the Sandhills.

Discussion

What sort of genetic differences should we have

expected among populations of longleaf pine? The

species occupies a range without great extremes of

temperature. In the northern part of Georgia longleaf

pine is subject to freezing temperatures yearly, and

9.

P-115

8- P-109

7-

';;iS

t

j5

E4

WI .-

;3

0'

5

I I

10 15

Age (years)

Figure 9-Comparison of the most southern (2768) and most

northern (2759) seed sources at the most southern ( ) and

most northern (-) sites. At both locations the local seed source

has grown better than the others.

light snow or ice is not uncommon. Except at its

most southerly occurrence, freezing temperatures

are likely to occur at some time during the normal

lifetime of a tree as shown by the recent (1984-l 985)

repeated devastating cold damage to the citrus groves

as far south as Orlando, FL. Throughout the longleaf

pine range east of the Mississippi River, rainfall seldom

is limiting and soil moisture would only be a factor

on sites so wet that longleaf could not compete with

slash pine or hardwoods, or on the most extreme

sandy soils occupied by sand pine (P. clausa (Chapm.

ex Engelm.) Vasey ex Sarg.). Most of the longleaf

pine range is below 500 feet (152 m) elevation in a

region in which changes in elevation are more likely

to be associated with differences in soil type than

with climate. Undoubtedly the combined effects of

temperature, rainfall, soils, and elevation exert some

genetic selection pressure on longleaf pine, but it

has not likely produced extremes of variation among

seed sources.

A significant factor with which longleaf pine must

contend for survival is brown-spot needle blight, and

there is evidence for marked differentiation in susceptibility to this disease among longleaf pine seed

sources. Wells and Wakeley (1970) reported high

12
infection in Texas and Louisiana sources. The Glades

County, FL, source was highly susceptible in this

study (Kraus 1986).

Another potentially significant source of genetic

variation in longleaf pine is its ability to hybridize

naturally with both loblolly pine (Chapman 1922) and

slash pine (Perry and Wang 1957). In general the

separation of flowering times is greater between

longleaf pine and slash pine than between it and

loblolly pine, so the possibility of introgression with

slash pine is much less than with loblolly. The natural

hybrid with loblolly is much more common than with

slash. There was some evidence of hybridization

with loblolly in the cones collected for this study. At

least one tree from Meriwether County (2758) had

typical longleaf foliage and buds but smaller than

normal cones with spinosity and color resembling

loblolly pine, and collections from two trees in Pulaski

County (2766) had cones which were not typical for

longleaf pine. Concrete evidence for longleaf x loblolly

pine hybridization was found in the nursery (table 3).

It should be noted that all of the Georgia hybrids

were in sources from Piedmont, Sandhills, or Upper

Coastal Plain collections. Early descriptions of the

southern pine forests emphasize that the ‘pine-barren

flats’ (Flatwoods) and the ‘rolling pine hills’ (Lower

Coastal Plain) were dominated by Savannah-like

stands of longleaf pine (Mohr 1897, Plate l), while in

the ‘belt of mixed growth’ (Upper Coastal Plain)

loblolly pine and shortleaf pine (P. echinafa Mill.)

were the dominant species. Given their interspecific

crossabilii and the mixed character of the virgin

stands, it is easy to suggest that the needle lengths

measured on the seedlings (table 4) reflect past and

present introgression of longleaf and loblolly pines.

Another potential source of genetic differentiation in

longleaf pine is past logging history. One of the

larger sources of variation in this study was the

differences among seed sources within physiographic

province. Such differences could be the result of

timber harvests conducted in such a way that the

subsequent regeneration was largely from trees

genetically inferior for growth and timber characteristics (dysgenic selection) on some areas and from

average or better trees on other areas.

One might expect to find evidence of longleaf x loblolly

pine introgression in the seed wing data, but such

was not the case. Seed wing striping was obviously

an individual-tree characteristic not associated with

geographic seed source. The seed wing measurements showed no evidence of shorter or slimmer

seed wings resembling loblolly pine among the seed

sources from the Piedmont, Sandhills, or Upper

Coastal Plain.

Additional evidence of possible longleaf x loblolly

pine introgression comes from the nursery measurements of seedling needle lengths (table 4) where

the more southerly seed sources show a distinct

tendency to have longer needles than the sources

from the part of the range where loblolly pine is the

dominant species. The major exception was the

Pasco County, FL, source-which had germinated

poorly in the Morgan nursery and produced only

enough seedlings to be used at two planting locations.

The survival data for the first 5 years (tables 5, 6, 8;

figs. 4, 6) correlate well with the nursery needle-length

data (fig. 6) and tend to support the idea that the

northerly sources of longleaf pine contain some

loblolly pine genetic material. With survival, there

could also be an element of climatic selection involved.

Longleaf pine seeds generally germinate soon after

seedfall and overwinter as seedlings, growing a

taproot when temperatures are favorable. It is

conceivable that natural selection for cold and drought

tolerance could take place during the seedling stage,

and that seedlings having some level of interspecific

hybridization would be more likely to survive.

The ability to tolerate adverse conditions early in life

may carry over for a year or two. Although most of

the differences in number of years out of the ‘grass

stage’ (YOG) at age 5 were among seed sources

within province, the more northerly seed sources

tended to begin height growth younger than those

from farther south. This ability to get an early start in

height growth is important and has long-lasting effects.

There is a highly significant correlation (R = 0.66)

between YOG and average 15th-year height, but not

between YOG and growth rate between 10 and 15

years.

It will probably be several more assessment intervals

before the results of this study will be conclusive.

Some of the seed sources are still growing well at

13
the most northern site (fig. IO), but the height growth

of both the fastest and slowest growing sources has

apparently culminated (fig. 11) at the most southern

site. The strong effect of planting site is evident in

the growth of the most southern and most northern

seed sources, both of which have slowed down in

their height growth rates at the very sandy, welldrained Sandhills and central Florida locations (figs.

12, 13). Other seed sources also seem to be slowing

down; at the most fertile site (Upper Coastal Plain),

and at the poorest site (Sandhills), both the fastest

and slowest growing seed source at each location

appear to be slowing in their height growth (fig. 14).

To better define the interaction between seed source

and plantation for height growth, the average latitude

of the tallest seed source in each plantation was

regressed on the plantation latitude. This regression

was done for ages 10 and 15 years. The results

show that the farther north of latitude 31’36 N. a

plantation was located, the farther south of the planting

site was the average latitude of the best seed source

(fig. 15). The inverse applied to plantations south of

latitude 31”36’ N., with the average latitude of the

best seed sources in the most southerly plantation

being almost 2” north of the planting location. Based

on these results, an optimum seed collection zone

for longleaf pine to be grown in Georgia would lie

between 30”39’ N. and 32”30’ N. (fig. 16).

Based on these growth data, it is possible to use

figure 17 to locate an approximate area from which

to obtain seed for a given planting location. For

example, with a proposed planting site at latitude

30” N., one would want to go 1 .o” north for seed,

always keeping in mind the strong effect of individual

stand variation. For a planting site at 33” N., one

would look about 0.9” south for seed, remembering

that survival might be somewhat lower with a southerly

seed source.

One of the objectives of this research was to determine

the necessity of establishing separate longleaf pine

breeding programs for different parts of Georgia. At

this time, that does not seem necessary. The optimum

zone shown in figure 16 is quite broad, and the

variation among seed sources within province is so

great that a longleaf pine breeding population for

Georgia can include trees from good stands anywhere

in the lower half of the State.

5 10 15

Age (years)

Figure lo-Comparison of fastest and slowest growing seed sources

at the most northern site (P-109). Both sources are from Florida,

and there is no indication that the rate of height growth has

culminated.

1 0

Age (years)

Figure 11 -Comparison of fastest and slowest growing.seed sources

at the most southern site (P-115). Rate of height growth has

apparently culminated for both sources.

’ 14
P-111

P-113

P-112

P-109

P-114

P-115

P-110

“5 10

Age (years)

Figure 12-Comparison of height growth of the most northern

seed source (2759) at all sites. Rate of growth has slowed down at

the Sandhilk (P-l 10) and central Florida locations (P-l 15) but

increased in north Florida (P-l 14).

12r P-l,,

P-113

P-116

P-100

P-114

P-l,2

P-110

3

2

1

01

5 10

Age (years)

Figure 13-Comparison of height growth of the most southern

seed source (2759) at all sites. Rate of growth has apparently

culminated at the Sandhills (P-l 10) and central Florida (P-l 15)

locations but increased at the Piedmont (P-109) and north Florida

(P-l 14) sites.

14

13

12 F /

P-111

P-110

0

5

,

10

Age (years)

15

Figure 14-Comparison of fastest and slowest growing seed sources

at the best and poorest sites. Growth is apparently beginning to

slow down.

QI 34

r

i! I

‘0

g 33-

(D

f j “““““‘~15~ear’, , ,

5 &-

2 29 30 31 32 33 34

Latitude of plantation

Figure 15 -Regression of average latitude of best seed sources on

plantation latitude at 10 and 15 years. The slopes of the regressions

are almost identical, but the average latftude of the fastest growing

sources has shifted slightly south.

15
Figure 16--Zone of best general

of longleaf pine in Georgia.

North 1

- - - - Natural range of longleaf pine

* Days in growing season

29 3 0 31 3 2

Latitude of planting site

Figure 17-Regression on planting site latiiude of deviation of

latitude of fastest growing seed source from planting latitude. This

can be used to locate an appropriate area from which to obtain

seed for a given latitude.

Acknowledgments

This research was funded in part by the Georgia

Forestry Commission. Its personnel helped locate

stands for cone collections and assisted with the

assessment of the study plantations. Some of the

cone collections were provided by the Florida Division

of Forestry; the Brunswick Pulp Land Co.; the Buckeye

Cellulose Corp.; the Forestry Department of the

University of Florida; the Southern Institute of Forest

Genetics of the Southern Forest Experiment Station;

270*

and the Southeastern Forest Experiment Station

staffs at Fort Myers, Olustee, and Marianna, FL. Land

and fire protection were generously provided by the

Georgia Kraft Company in Polk and Houston Counties,

GA.; by the Armstrong Cork Co. in Crawford County,

GA.; Union Camp Corp. in Laurens County, GA.;

Brunswick Pulp Land Co. in Ware County, GA.; the

National Forests in Florida in Lake County, FL., and

the Naval Stores and Timber Production Laboratory

of the Southeastern Forest Experiment Station in

Baker County, FL

16
Literature Cited

Allen, Peter Ii. 1981. Florida longleaf pine fail in Virginia. Journal

of Forestry 591453454.

Allen, R.M.; MacGregor, W.H.D. 1982. Seedling growth of three

southern pine species under long and short days. Silvae

Genetica 114345.

Bethune, James E.; Roth, Elmer R. 1980. Source of seed affects

growth of longleaf pine-fifth year results. Res. Notes 148.

Asheville, NC: U.S. Department of Agriculture, Forest Service,

Southeastern Forest Experiment Station. 2 pp.

Bcyer, W.D.; Evans, S.R. 1987. Early flowering in longleaf pine

related to seed source. Journal of Forestry 85808.

Boyer, William D. 1978. Heat accumulation: an easy way to

anticipate the flowering of southern pines. Journal of Forestry

7820-23.

Chapman, H.H. 1922. A new hybrid pine (Pinus palustris x Pinus

taeda). Journal of Forestry 20:729-734.

Collins, Arthur B., Ill. 1984. Longleaf pine seed source planting in

Georgia-tenth year results. Res. Note SE-19. Asheville, NC:

U.S. Department of Agriculture, Forest Service, Southeastern

Forest Experiment Station. 3 pp.

Derr, Harold J. 1971. Brown-spot resistance among progenies of

longleaf plus trees. In: Proceedings of the 11 th conference on

southern forest tree improvement; 1971 June 15-18; Atlanta GA.

Macon, GA: Southern Forest Tree Improvement Committee.

45-51, In cooperation with: Southeastern Area, State and Private

Forestry, Region 8, U.S. Department of Agriculture, Forest

Service.

Harlow, William Morehouse; Harrar, Ellwood S. 1958. Textbook

of dendrology: covering the important forest trees of the United

States and Canada. 4th ed. New York: McGraw-Hill Book Co.

581 PP.

Harms, William R. 1987. A test of local longleaf pine seed source.

Charleston, S.C: U.S. Department of Agriculture, Forest Service,

Southeastern Forest Experiment Station; final report summary,

FS-SE-1 108,Study S-20. 1 p.

Henry, B.W.; Wells, Osborn 0. 1987. Variation in brown-spot

infection of longleaf pine from several geographic sources.

Res. Note SO-52. New Orleans, IA: U.S. Department of

Agriculture, Forest Service, Southern Forest Experiment Station.

4 PP.

Kraus, John F. 1985. Fusiform rust infection in longleaf pine from

various seed sources in two Georgia plantings. In: BarrowsBroaddus, Jane; Powers, Harry R., Jr., comps.,eds. Proceedings

of the rusts of hard pines working party conference S2.08-10,

IUFRO; 1984 October 1-3; Athens, GA. Athens, GA: Georgia

Center for Continuing Education, University of Georgia: 5358.

Kraus, John F. 1988. Brown-spot needle blight susceptibility of

longleaf pine seed sources in Georgia and Florida. In: Recent

research on conifer needle diseases: Conference proceedings;

1984 October 14-18; Gulfport, MS. Gen. Tech. Rep. WO-50.

Washington, DC: U.S. Department of Agriculture, Forest Service:

11-14.

Mohr, Charles. 1897. The timber pines of the southern United

States-together with a discussion of the structure of their wood

by Filbert Roth. Revised. Bull. 13. Washington, DC: U.S.

Department of Agriculture, Division of Forestry. 178 pp.

Perry, T.O.; Wang, Chl Wu. 1957. Cooperative forest genetics

research program. Res. Rep. 4. Gainesville, FL: University of

Florida, School of Forestry. 27 pp.

Rockwccd, D.L; Kok, H.R. 1977. Development and potential of a

longleaf pine seedling seed orchard. In: Proceedings of the

14th southern forest tree improvement conference; 1977 June

14-18; Gainesville, FL. Gainesville, FL: 7888.

Shoulders, Eugene. 1985. Seed origin affects longleaf pine in

Louisiana plantation. Res. Note SO-19. New Orleans, LA: U.S.

Department of Agriculture, Forest Service, Southern Forest

Experiment Station. 3 pp.

Shoulders, Eugene; Thomas, Terry A. 1988. Climate at the seed

source affects longleaf pine performance in Louisiana plantation.

Res. Note SO-78. New Orleans, LA: U.S. Department of

Agriculture, Forest Service, Southern Forest Experiment Station.

5 PP.

Snyder, E.B. 1981. Racial variation in root form of longleaf pine

seedlings. In: Proceedings of the 8th southern conference on

forest tree improvement; 1981 June 7-8; Gainesville, FL.

Gainesville FL: University of Florida, School of Forestry: 53-59.

Snyder, E.B.; Allen, R.M. 1988. Mountain longleaf pine excels

only in local plantings. Res. Note SO-83. New Orleans, LA: U.S.

Department of Agriculture, Forest Service, Southern Forest

Experiment Station. 4 pp.

Snyder, E. Bayne; [krr, Harold J. 1972. Breeding longleaf pines

for resistance to brown spot needle blight. Phytopathology

82:325-329.

Wakeley, Philip C. 1959. Five-year results of the southwide pine

seed source study. In: Proceedings of the 5th southern conference on forest tree improvement; 1959 June 11; Raleigh, NC.

Raleigh, NC: North Carolina State University, School of Forestry:

511.

Wakeley, Philip C. 1981. Results of the southwide pine seed source

study through 1960-61. In: Proceedings of the 8th southern

conference on forest tree improvement; 1981 June 7-8;

Gainesville, FL. Gainesville, FL: The University of Florida, School

of Forestry: 10-24.

17
Wakeley, Phlllp C. 1966. How far can seed be moved? In:

Proceedings of the 7th southern conference on forest tree

improvement; 1966 June 26-27; Gulfport, MS. New Orleans,

IA: U.S. Department of Agriculture, Forest Service, Southern

Forest Experiment Station: 3643.

Wells, 0.0. 1969. Results of the Southwide Pine Seed Source

Study through 196669. In: Proceedings of the 10th southern

conference on forest tree improvement; 1969 June 17-19;

Houston, TX. College Station, TX: Texas Forest Service, Texas

A 8 M University: 117-129.

Wells, 0.0.; Snyder, E.B. 1976. Longleaf pine half-sib progeny

test. Forest Science 22M4-406.

Wells, Oaborn 0. 1975. Geographic variation in southern tree

species. In: Thielges, B.A., ed. Forest tree improvement-the

third decade. Baton Rouge, LA: Louisiana State University,

Division of Continuing Education: 19-66.

Wells, Oaborn 0.; Wakelay, Philip C. 1970. Variation in longleaf

pine from several geographic sources. Forest Science 16:26-42.

18
Table l--Numbers of longleaf pines Table 2--Seedlots with xanthescens' seedlings

having solid-colored vs. striped seed wings at Morgan Nursery

Seed source, by

State and county

Trees with seedwings

Solid

color Striped

Alabama Baldwin 18

Florida Glades 15

Jackson 18

Marion 15

Taylor 20

Georgia Bryan

Colquitt

Jefferson

Jenkins

Meriwether

Paulding

Pulaski

Richmond

Taylor

Telfair

Terre11

Ware

Washington

20

20

18

18

19

17

17

20

18

18

16

23

18

17

Number Number

Bulked lots

2

5

2

0

0

0

0

2

2

2

z

0

2

G

3

2

?

2754 Bryan, GA

2755 Colquitt, GA

2756 Jefferson, GA

2762 Taylor, GA

2765 Ware, GA

2766 Washington, GA

2767 Wayne, GA

3065 Marion, FL

Individual-tree lots

2912 Richmond, GA

2927 Colquitt. GA

2999 Washington, GA

3084 Taylor, FL

3961 Jackson, FL

1

2

1

2

:

1

1

1

t

3

1

1

Cotyledons are normal green;

primary foliage is yellow.

Seed source, by

Seedlot number county and State Seedlings

Table 3--Individual-tree seedlots fm Georgia

with Sonderegger seedlings

Total 345 37

Seedlot number Seed source

(county) Seedlings

2814 Jefferson 1 1 of 22

2821 Jefferson 10 1 2 0f 27

2824 Jefferson 14 2 of 26

2862 Meriwether 15 1 of 27

2886 Paulding 20 2 0f 25

2901 Pulaski 16 1 of 39

2918 Richmond 13 4 of 35

3010 Washington 16 1 of 33

Number

19
Table &-Needle lengths of longleaf pine seedlings after 1 year of

growth in a Florida and a Georgia nursery

Needle length

Andrews Morgan Morgan

Seedlot Origin Physiographic Nursery, Nursery,

number County State province FL GA

Nursfry,

GA

2759 Paulding GA

2758 Meriwether GA

2762

2766

2761

Taylor GA

Washington GA

Richmond GA

2764 Terre11 GA

2760 Pulaski GA

2756 Jefferson GA

2755 Colquitt GA

2763 Telfair GA

2757 Jenkins GA

2765 Ware GA

2767 Wayne GA

2754 Bryan GA

2753 Baldwin AL

2769 Jackson FL

3066 Taylor FL

3065 Marion FL

2768 Glades FL

3085 Pasco FL

Piedmont

Sandhills

Upper Coastal

Plain

Lower Coastal

Plain

Flatwoods 42.9

Alabama &

Florida

35.2 43.1 37.7

33.1 42.5 35.7

37.3 43.6 39.6

39.6 44.5 40.3

39.9 43.0 38.9

39.8 45.2 41.0

39.1 45.2 41.0

40.3 42.1 39.2

41.1 41.9 40.2

42.4 42.8 39.2

37.3 41.5 38.3

41.6 46.5 41.7

44.1 48.4 44.7

41.4 46.5 41.7

39.5 44.4 40.1

43.0 45.2 40.3

40.5 47.0 39.7

42.2 46.6 42.4

45.8 49.9 42.0

45.0 47.8 _-

44.3 46.2 37.4

40.4 34.3 _-

+* ns. . ** ns. .

48.2

46.4

$9.

42.6

44.0

42.2

41.5

** Within the indicated column, means differ significantly at the l-percent level.

* Within the indicated column, means differ significantly at the 5-percent level.

n.s. Within the indicated column, means do not differ at the 5-percent level.

1

Means of 10 individual-tree progenies per source.

20
Table 5--Summary of lst-year survival by seed source origin

Seedlot

number

Mean lst-year Deviation

Physiographic survival from

province Seedlot Province grand mean

------~---------Per~ent-~~~-----~~~~~~~~

2759

2758

2762

2766

2761

2764

2760

2756

- 2755

2763

2757

2765

2767

2754

2753

2769

3066

;z:

2768

Piedmont

Sandhills

Upper Coastal

Plain

Lower Coastal

Plain

Flatwoods

n.s.

68.3

71.8

~~~~

63:5

64.4

63.6

68.2

55.4

61.6

53.8

:z

56:3

it+

70.0

64.0

10.4

4.4

65.4 5.8

56.9 -2.7

55.0 -4.6

Alabama &

Florida

Grand mean

50.1

;Ei

50:4

55.0

53.2

54.1 -5.5

59.6

** Within the indicated column, means differ significantly at the l-percent

level.

n.s. Within the indicated column, means do not differ significantly at the

5-percent level.

21
Table 6--~uamary of overall second-year survival of longleaf pine seed sources

Plantation location, by county and State Deviation

Origin Physiographic Crawford Houstpn Laurens Baker Means from grand

Seedlot County State province GA GA GA FL Seedlot Province mean

______---------pe rcent______ l *

l * 63.2 11.0

63.3

63.2

58.9 6.7

51.2

65.1

60.5

60.0 7.8

2759 Paulding GA

2758 Meriwether GA

Piedmont

----__

n.s.

82.0

74.5

Sandhills

l * l * **

56.3 42.4 72.4

68.7 47.9 61.6

2762 Taylor GA

2766 Washington GA

2761 Richmond GA

38.8

68.4

75.6

Upper Coastal

Plain

55.2

66.1

69.0

Lower Coastal

Plain

34.2

66.1

44.8

32.2 56.6 77.2

55.9 71.1 65.1

35.9 63.4 67.1

2764 Terre11 GA

2760 Pulaski GA

2756 Jefferson GA

43.4 65.6 68.6 58.2

29.5 74.7 67.7 59.5

50.0 62.3 67.9 62.3

51.2 -1.0

2755 Colquitt GA

2763 Telfair GA

2757 Jenkins GA

20.6 50.2 67.4 43.1

49.5 63.4 66.4 61.4

21.6 57.5 72.5 49.1

2765 Ware GA

2767 Wayne GA

2754 Bryan GA

Flatwoods

27.0 24.4 57.6 69.4 44.6

43.7 33.1 27.6 62.1 41.6

53.7 32.7 48.8 67.9 50.8

45.7 -6.5

Alabama & 43.9 -8.3

Florida

2753 Baldwin AL 31.6 41.7 35.4

2769 Jackson FL 49.3 32.9 47.6

3066 Taylor FL 35.7 33.6 48.3

3065 Marion FL 32.2 29.7 37.7

66.0

74.8

63.8

56.3

__

43.7

51.2

45.4

39.0

3085 Pasco FL -- -_ 51.8 - -

Mean** 50.2 36.0 53.9 68.5 52.2

l * Within the indicated row or column, means differ significantly at the l-percent level.

n.6. Within the indicated column, means do not differ significantly at the s-percent level.
Table 7--Summary of overall percentage of trees out of the “grass stage” 2 years after planting for longleaf pine

seed sources

Plantation location, by county and State Deviation

Origin Physiographic Crawford Houston Laurens Baker Means from grand

Seedlot County State province GA GA GA FL Seedlot Province mean

_______________percent-_-_---__________ l

* l

12.6 2.6

11.9 23.2

12.4 21.3

28.4 15.3

29.8 4.5

n.s. l

3.3 4.0

6.7 9.9

Piedmont

Sandhills

l * 9.2

5.6

12.9

12.6

12.4

15.3

10.1

11.9

0.5

3.9

2759 Paulding GA

2758 Meriwether GA

2762 Taylor GA

2766 Washington GA

2761 Richmond GA

6.4 9.5

12.0 5.4

4.0 2.0

Upper Coastal

Plain

3.2

2764 Terre11 GA 24.8 21.0 6.6 4.6 14.2

2760 Pulaski GA 10.6 11.9 6.4 5.6 8.6

2756 Jefferson GA 19.8 23.5 5.7 3.1 13.0

Lower Coastal 8.0 -0.7

Plain

2755 Colquitt GA 10.7 .3 4.3 5.3 5.2

2763 Telfair GA 17.3 24.4 16.0 .2 14.5

2757 Jenkins GA 4.5 7.4 4.1 .9 4.2

Flatwoods 6.8 -1.9

2765 Ware GA 7.1 7.6 1.9 4.0 5.2

2767 Wayne GA 15.5 9.1 1.1 4.8 7.6

2754 Bryan GA 4.2 13.8 5.4 6.5 7.5

Alabama &

Florida

5.7 -3.0

2753 Baldwin AL

2769 Jackson FL

3066 Taylor FL

3065 Marion FL

2768 Glades FL

3085 Pasco FL

1.7 13.3 2.0 1.3 4.6

4.8 10.2 .5 9.6 6.3

16.0 10.9 5.5 8.6 10.2

2.6 1.6 2.8 6.7 3.4

8.4 2.2 2.2 2.3 3.8

-- _- 28.9 -- -_

Mean +* 12.8 11.8 6.3 5.0 8.7

** Within the indicated column or row, means differ significantly at the l-percent level.

l Within the indicated column, means differ significantly at the 5-percent level.

n.s. Within the indicated column, means do not differ significantly at the 5-point level.
$2 Table 8--Summary of overall fifth-year survival of longleaf pine seed sources

Plantation location, by county and State Deviation

Seedlot Origin Physiographic Polk Crawford Houston Laurens Ware Baker Lake Means from grand

number County State province GA GA GA GA GA FL FL Seedlot Province mean

2759 Paulding GA 47.4 53.3 48.6 73.3 89.5 78.0 54.7 63.5

2758 Meriwether GA 58.7 63.8 52.8 68.4 91.3 72.1 61.9 67.0

Sandhills

2762 Taylor GA 29.5 39.3 40.5 71.9 94.7 74.2 52.8 57.6

2766 Washington GA 35.9 63.8 60.7 82.3 93.5 58.8 50.2 63.6

2761 Richmond GA 42.5 71.4 38.9 66.4 77.2 63.6 43.2 57.6

Upper Coastal

Plain

2764 Terre11 GA 55.4 51.8 46.7 71.8 87.3 64.1 51.4 61.2

2760 Pulaski GA 41.7 62.6 35.6 77.7 93.0 67.3 47.6 60.8

2756 Jefferson GA 61.9 63.8 56.8 73.5 95.0 68.2 51.4 67.2

Lower Coastal

Plain

2755 Colquitt GA 20.0 34.2 28.7 59.0 73.0 66.1 52.8 47.7

2763 Telfair GA 47.6 52.8 54.4 68.2 80.5 62.3 37.6 57.6

2757 Jenkins GA 41.3 45.3 25.8 65.1 81.1 71.1 22.5 50.3

Flatwoods 48.4 -5.6

2765 Ware GA 41.3 25.9 32.1 57.5 78.5 62.3 36.2 47.7

2767 Wayne GA 18.9 41.7 39.3 43.0 73.5 50.7 48.6 45.1

2754 Bryan GA 30.5 48.8 38.4 56.4 88.0 63.6 40.5 52.3

Alabama &

Florida

2753 Baldwin AL 24.2

2769 Jackson FL 37.4

3066 Taylor FL 17.5

3065 Marion FL la.5

2768 Galdes FL 13.9

3085 Pasco FL _-

27.8 44.6

45.8 38.2

33.6 33.4

31.0 30.5

35.7 28.1

90.6 62.4

82.7 74.8

al.7 60.4

71.1 51.9

58.0 46.2

- - - -

47.6

53.7

56.1

47.6

27.5

56.8 - - - -

43.9 48.7

48.6 54.5

54.2 48.1

52.1 43.2

22.3 33.1

42.2 49.5

Piedmont

n.s. n.s. n.s.

65.3 11.3

59.6 5.6

63.1 9.1

51.9 -2.1

45.7 -8.3

Mean l * 36.0 47.0 40.7 61.2 83.2 64.1 45.7 54.0

l * Within the indicated row or column, means differ significantly at the l-percent level.

l Within the indicated column, means differ significantly at the s-percent level.

n.s. Within the indicated column, means do not differ significantly at the s-percent level.
'fable +-Summary of average number of years out of the "grass stage" 5 years after planting for longleaf pine *ource*

Plantation location, by county and State Deviation

Seedlot Origin Physiographic Polk Crawford Houston Laurens Ware Baker Lake Means from grand

number County State province GA GA GA , GA GA FL FL Seedlot Province mean

___-----______---__________-----yeBr---____________________________

l l * l n.s. n.s. n.s. n.s.

1.3 1.2

1.6 1.6

1.5 1.7

1.5 1.7

1.3 1.5

1.4 1.8

1.4 1.6

1.4 1.7

1.3 1.4

1.3 1.3

1.0 1.4

1.6 1.0

1.1 1.4

1.5 1.6

l * n.6.

0.0

.2

Piedmont 1.9

1.8

1.9

2.1

2.1

2.2

2.2

2.1

2759 Paulding GA 1.8 1.8 2.8 1.9 2.0

2758 Meriwether GA 2.0 1.9 3.0 1.5 1.8

Sandhills

2762 Taylor GA 1.9 1.9 3.0 2.2 2.2

2766 Washington GA 2.3 2.3 3.1 2.1 2.5

2761 Richmond GA 2.4 2.4 2.8 2.4 2.4

Upper Coastal

Plain

2764 Terre11 GA 2.4 2.2 3.2 2.1 2.3

2760 Pulaski GA 1.7 1.8 2.7 2.0 2.0

2756 Jefferson GA 2.6 2.2 3.0 1.7 2.5

Lower Coastal

Plain

2755 Colquitt GA 2.4 2.2 3.0 1.6 1.8

2763 Telfair GA 2.0 2.1 3.4 2.5 2.5

2757 Jenkins GA 1.9 2.1 2.9 2.2 2.2

Flatwoods

2765 Ware GA 1.9 1.9 2.3 1.6 1.8

2767 Wayne GA 1.9 2.0 2.9 1.4 1.7

2754 Bryan GA 1.8 1.7 3.2 2.1 2.4

.2

2.2

1.9

2.2

2.0 .I

2.0

2.2

2.0

1.8

1.7

1.8

2.0

-.I

Alabama & 1.8 -.l

Florida

2753 Baldwin

2769 Jackson

3066 Taylor

3065 Marion

2768 Glades

3085 Pasco

AL 1.5 1.7 3.2 1.3 1.9 1.3 1.9 1.8

FL 1.9 2.1 3.0 1.9 2.1 1.7 1.9 2.1

FL 2.2 2.0 3.0 1.5 2.0 1.5 1.7 2.0

FL 1.5 1.6 2.7 1.5 1.8 1.4 1.2 1.7

FL 1.5 1.6 2.1 .8 1.1 .7 .8 1.2

FL _- -- __ 2.5 __ __ 2.6 2.6

Mean ** 2.0 2.0 2.9 1.8 2.0 1.4 1.6 1.9

** Within the indicated row or column, means differ significantly at the l-percent level.

l Within the indicated column, means differ significantly at the s-percent level.

n.s. Within the indicated column, means do not differ significantly at the s-percent level.

.I
% Table lo--Summary of overall fifth-year average height of longleaf pine seed sources

Plantation location, by county and State Deviation

Seedlot Origin Physiographic Polk Crawford Houston Lauren6 Ware Baker Lake Means from grand

number County State province GA GA GA , GA GA FL FL Seedlot Province mean

2759 Paulding GA 2.77 2.59 5.34 2.88 3.67 1.67 1.59

2758 Meriwether GA 3.07 3.12 6.76 2.35 3.34 2.27 2.59

Sandhills

2762 Taylor GA 3.24 3.09 6.98 2.69

2766 Washington GA 3.99 4.20 6.91 3.54

2761 Richmond GA 4.09 4.17 5.71 2.98

4.41

5.36

5.08

2.23 2.63

2.35 2.47

1.71 2.53

Upper Coastal

Plain

2764 Terre11 GA 4.20 3.89 7.39 2.98 4.82 2.29 3.27

2760 Pulaski GA 2.49 3.00 6.00 3.07 4.26 2.02 2.68

2756 Jefferson GA 4.64 3.58 7.42 2.79 5.41 2.11 3.04

Lower Coastal

Plain

2755 Colquitt GA 3.99 4.51 6.34 2.78 3.82 1.97 2.46

2763 Telfair GA 3.29 3.42 8.17 3.57 6.31 1.89 2.23

2757 Jenkins GA 3.20 3.45 6.49 2.69 4.71 1.44 2.09

Flatwoods

2765 Ware GA 2.94 2.88 5.22 2.42 3.64 2.38 1.81

2767 Wayne GA 3.29 3.41 6.55 1.92 3.68 1.56 2.41

2754 Bryan GA 2.93 2.46 6.93 3.28 5.23 2.09 2.58

Alabama &

Florida

2753 Baldwin AL

2769 Jackson FL

3066 Taylor FL

3065 Marion FL

2768 Glades FL

3085 Pasco FL

2.29 2.57 7.73 2.17 4.10 1.83 3.18 3.41

3.16 3.67 6.60 2.69 4.49 2.83 3.40 3.83

3,72 3.52 7.16 2.44 4.20 2.38 3.20 3.80

2.04 2.40 6.09 2.27 3.79 2.20 2.01 2.97

2.06 2.05 4.40 1.19 1.75 1.14 1.31 1.99

_- -- -_ 4.63 -_ __ 6.09 5.36

3.23 3.26 6.54 2.77 4.32 2.02 2.68 3.53

Piedmont

Mean l *

n.s. n.s. n.s. n.s. l * n.s.

3.14 -0.39

2.93

3.36

3.83 .3o

3.61

4.12

3.75

3.87 .32

4.12

3.36

4.14

3.75 .22

3.70

4.13

3.44

3.31 -.22

3.04

3.26

3.64

3.32 -.21

l * Within the indicated row or column, means differ significantly at the l-percent level

l Within the indicated column, means differ significantly at the s-percent level.

n.6. Within the indicated column, means do not differ significantly at the s-percent level
Table 11--Su~=ary of overall annual height growth rate to age 5 years of longleaf pine seed sources

Plantation location, by county and State Deviation

Seedlot Origin Physiographic Polk Crawford Houston ,Laurens Ware Baker Lake Means from grand

number county State province GA GA GA GA GA FL FL Seedlot Province mean

____________________________________________Feet_________________----______-_______________________

n.s. n.s. n.s. n.s. n.s. n.s. n.s. l * n.s.

1.59 -0.11

1.51

1.67

1.71 .Ol

1.69

1.75

1.69

1.75 .o5

Piedmont

2759 Paulding GA

2758 Meriwether GA

1.55 1.41 1.86 1 49 1.77 1.17 1.29

1.54 1.60 2.22 1 57 1.85 1.33 1.56

Sandhills

44 1.99 1.40 1.51

67 2.09 1.43 1.41

43 2.09 1.33 1.49

2762 Taylor

2766 Washington

2761 Richmond

GA

GA

GA

GA

GA

GA

Upper Coastal

Plain

1.66 1.56 2.27

1.69 1.77 2.19

1.72 1.70 2.04

53 2.05 1.40 1.73 1.78

59 2.10 1.37 1.59 1.69

58 2.15 1.43 1.65 1.79

2764 Terre11

2760 Pulaski

2756 Jefferson

1.72 1.70 2.30

1.43 1.56 2.16

1.77 1.55 2.43

Lower Coastal 1.73 .03

Plain

1

1

1

Flatwoods

1

1

1

Alabama &

Florida

1

1

1

1

1

2755 Colquitt GA

2763 Telfair GA

2757 Jenkins GA

63

70

66

90 2.11

54 2.39

58 2.23

66 2.06 1 . 41 1.71

53 2.38 1 . 32 1.68

36 2.10 1. 29 1.05

1.78

1.79

1.61

1

1.64

1.68

1.69

1

67 -.03

69 -.Ol

56

66

55

46 2.21

69 2.18

40 2.17

48 1.88 1. 38 1.54

40 2.01 1. 31 1.52

57 2.23 1. 36 1.55

2765 Ware GA

2767 Wayne GA

2754 Bryan GA

64

61

58

45

36

__

45

72

68

45

25

--

2.36

2.15

2.35

2.16

2.03

--

58

35

55

56

40

85

2.05 1 . 25 1.62 1.71

2.06 1 . 54 1.78 1.74

2.07 1 . 41 1.79 1.78

2.08 1. 41 1.63 1.68

1.40 1. 37 1.45 1.47

__

2753 Baldwin AL

2769 Jackson FL

3066 Taylor FL

3065 Marion FL

2768 Glades FL

3085 Pasco FL

Mean ** 1.60 1.58 2.20 1.53 2.02 1.36 1.58 1.70

** Within the indicated row or column, means differ significantly at the l-percent level.

Y

n.s. Within the indicated column, means do not differ significantly at the s-percent level.
Table 12--Summary of lOth-year data from longleaf pine seed sources. all plantations

Average annual Fusiform

Seedlot Origin Physiographic growth Volume rust

number County State province Survival Height 5-10 x 100 infection Forking

2759 Paulding G A

2758 Meriwether GA

2762 Taylor GA

2766 Washington GA

2761 Richmond GA

2764 Terre11 GA

2760 Pulaski GA

2756 Jefferson GA

2755 Colqaitt GA

2763 Telfair GA

2757 Jenkins GA

2765 Ware GA

2767 Wayne GA

2754 Bryan GA

2753 Baldwin AL

2769 Jackson FL

3066 Taylor FL

3065 Marion FL

2768 Glades FL

3085 Pasco FL

Piedmont

Sandhills

Upper Coastal

Plain

Lower Coastal

Plain

Flatwoods

Alabama &

Florida

Percent

l * l

‘58.6

56.3

60.9

55.6

52.3

59.0

55.4

58.5

57.2

55.8

62.4

47.3

42.7

52.8

46.3

46.2

45.5

43.1

50.0

41.4

47.4

51.2

44.9

36.8

21.7

m

l *

m

**

3

Ill _-_--percent______

-

l * l * l

4.5 0.65 1.03 11.0 5.0

5.0 .72 1.40 14.5 5.0

5.0 .72 1.35 12.9 4.0

5.4 .73 1.51 13.7 5.0

5.1 .69 1.43 13.1 3.5

5.5 .75 1.56 20.2 6.4

5.4 .a0 1.62 20.9 5.3

5.3 .70 1.64 15.3 6.9

5.6 .79 1.64 la.4 3.1

5.6 .73 1.71 17.5 5.5

5.7 .a5 1.63 15.9 3.3

5.0 .78 1.27 15.7 6.7

5.4 .73 1.66 a.4 4.3

5.3 .73 I. 58 11.7 3.2

5.3 .79

5.6 .78

5.4 .78

5.1 .77

4.5 .62

1.43

1.57

1.65

1.49

1.24

1.96

la.0

21.6

14.3

a.0

5.8

.4

6.3

4.4

2.6

3.9

1.2

46.3 6.2 .91 2.9

Mean 49.4 5.3 0.75 1.52 13.9 4.4

** Within the indicated column, means are significant at the l-percent level.

l Within the indicated column, means are significant at the 5-percent level.

28
Table 13--Summary of lsth-year data from longleaf pine seed sources. all plantations

Average annual Fusiform

Seedlot Origin Physiographic growth Volume rust

number county State province Survival Height 10-15 x 100 infection Forking

2759 PaUlding GA

2758 Meriwether GA

2762 Taylor GA

2766 Washington GA

2761 Richmond GA

2764 Terre11 GA

2760 Pulaski GA

2756 Jefferson GA

2755 Colquitt GA

2763 Telfair GA

2757 Jenkins GA

2765 Ware GA

2767 Wayne GA

2754 Bryan GA

2753

2769

3066

3065

2768

3085

Baldwin

Jackson

Taylor

Marion

Glades

Pasco

AL

FL

FL

FL

FL

FL

Overall Mean

Piedmont

Sandhills

Upper Coastal

Plain

Lower Coastal

Plain

Flatwoods

Alabama &

Florida

Percent m

l * l l *

55.3

53.1 7.9

57.5 8.3

53.5

50.3 8.4

56.8 9.2

53.3 8.7

55.8

54.8 9.2

53.2 9.1

59.5 8.9

44.3

39.4 9.4

49.7 9.4

43.8 9.0

43.4

42.0 9.0

40.7 9.3

47.4 9.0

38.8

45.3 8.9

48.2 9.4

40.7 9.2

33.8 9.0

19.1 7.8

3

m m ______percent______

- -

l * l * l * n.s.

0.72 3.88 12.1 5.2

.72 4.41 14.0 4.8

.75 4.56 15.6 8.2

.79 5.02 15.3 6.3

.76 5.06 17.2 5.7

.79 5.25 19.6 8.1

.77 5.21 21.3 6.9

.76 5.38 15.6 8.1

.80 5.99 19.3 3.7

.79 5.54 17.9 5.8

.78 5.05 15.1 4.9

.85 5.05 15.7 3.1

.82 5.63 13.2 3.6

.76 5.67 15.6 5.6

.77 4.64 18.4 5.5

.80 5.63 20.8 4.9

.81 5.69 17.0 4.1

.82 5.32 12.8 6.5

.72 4.92 9.2 2.9

45.7 10.2 .79 6.38 10.2 2.6

46.7 9.0 .78 5.21 15.8 5.3

l * Within the indicated column, means are significant at the l-percent level.

* Within the indicated column, means are significant at the 5-percent level

n.s. Within the indicated column, means are not significant at the 5-percent level.

29
Table l&--Summary of 15th-year combined analysis. showing variance components and their

significance

Traits

Source of

variation

Average annual

Fusiform height growth

rust lo-15 years Volume

d.f. Survival infection Height DBH x 10 x 10 Forks

_____________________ _--Variance comoonents________________________

Plantations 6 76.82""

Replications in

plantations 35

5

13

30

79

175

495

9.98**

Province 21.54'

Seedlots in

provinces 18.34*' 6.02~~ .18** .25** .38* .20** .58

Plantations x

province .88 0 .07* .16* .43 .19* 3.79*

Plantations x

seedlots in

province 16.45" 4.52 .06** .21** .53* .11 2.72

Replications in

plantations x

provinces

Error

0

71.17

413.17** 3.05"' 7.87**

7.66,' .31** .53**

0 .o3 0

0 0 0 0 0 0

81.81 1.00 1.87 10.46 2.70 90.42

23.63~1 11.30** 37.55""

2.82** .64**

.35 .08

3.02*

1.06

l Significant at the 5-percent level.

l * Significant at the l-percent level.

30
Table 15--Components of variation as a percentage of the total variation at 15 years

from combined data of all longleaf pine sources and plantations

Source of

variation Survival

Average annual Fusiform

height growth rust

Height lo-15 years Volume infection

Physiographic

provinces 10.01" 0.63 0.93 0.54 0.00

Seedlots 17.41"" 4.48"" 1.80"" 1.79"" .96

Seedlots in

provinces 8.52"" 3.94"" 1.00" 1.32"" 1.18""

Plantations x

provinces .41 1.44" 1.13 1.28" .oo

Plantations x

seedlots 8.11** 2.52"" 2.40"" 1.84"" .87

Plantations x

seedtots in

provinces 7.65"" 1.28"" 1.40" .73 .89

* Significant at the 5-percent level.

** Significant at the l-percent level.

31

; 1

Klaus, John F.; Sluder, Earl R. 1990. Genecology of longleaf pine in 1

Georgia and Florida. Res. Pap. SE-278. Asheville, NC: U.S. Department 1

Kraus, John F.; Sluder, Earl R. 1990. Genecology of longleaf pine in

Georgia and Florida. Res. Pap. SE-278. Asheville, NC: US. Department

1

2 I of Agriculture, Forest Service, Southeastern Forest Experiment Station. of Agriculture, Forest Service, Southeastern Forest Experiment Station.

Fourteen seed sources of Pinus palustris from Georgia, 5 from Florida,

and 1 from Alabama were grown at five locations in Georgia and at

two in Florida. Results through the 15th year show: (1) survival and

early height growth were best for the northern sources; (2) individual

stand or seed source contributed strongly to the components of variation;

(3) the seed source x plantation interaction was significant for growth

traits; and (4) the species has a generally broad zone of adaptability in

Georgia.

KEYWORDS: Pinus palusfris, geographic variation, survival and growth

traits, adaptability.

Fourteen seed sources of Pinus palusrris from Georgia, 5 from Florida,

and 1 from Alabama were grown at five locations in Georgia and at I

two in Florida. Results through the 15th year show: (I) survival and

early height growth were best for the northern sources; (2) individual 1

stand or seed source contributed strongly to the components of variation; 1

(3) the seed source x plantation interaction was significant for growth 1

traits; and (4) the species has a generally broad zone of adaptability in 1

Georgia.

I

KEYWORDS: Pinus palusrris, geographic variation, survival and growth 1

traits, adaptability. I

I

The Forest Service, U.S. Department of

Agriculture, is dedicated to the principle of

multiple use management of the Nation’s forest resources

for sustained yields of wood, water, forage, wildlife, and

recreation. Through forestry research, cooperation with the

States and private forest owners, and management of the

National Forests and National Grasslands, it strives-as

directed by Congress-to provide increasingly greater

service to a growing Nation.

USDA policy prohibits discrimination because of race,

color, national ongin, sex, age, religion, or handicapping

condition. Any person who believes he or she has been

discriminated against in any USDA-related activity should

immediately contact the Secretary of Agriculture,

Washington, DC 20250.