three-year height performance of pinus elliottii engelm. var ......three-year height performance of...

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Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites in Rhodesia By R. D. BARNES~) aad L. J. MULL IN^) (Received December 1977 1 August 1978) circumstances. P. elliottii is rnare drought resistant and also more tolerant of poior drainage than P. taeda. It is less severely affected by the group dying fungus, Rhizina in- flata (SCHAFF.) KARST., and less susceptible ta damage by fire. P. elliottii tends to have better stem and branch char- acteristics, allows more flexibility in management, its tim- ber is generally favourd by the trade and it is a high yielder of oleoresin. P. taeda on the other hand is more resistant to the pine woolly aphid, Pineus pini (L.), and to the honey fungus, Armillaria mellea (VAHL ex FR.) KUMMER. It captures the site more rapidly, is more tolerant of cold conditions and is much more resistant to windthrow and wind breakage at higher altitudes. P. taeda generally yields more timber than P. elliottii. Breeding Programmes far P. elliottii and P. taeda were started in Rhoidesia in 1960. Populations of plus trees (su- perior phenotypes) were built up and controlled crosses were made for progeny tests. Both the incentive and the conditions were therefore present for experimentation with hybrids between these two closely related species and Summary Pinus elliottii ENGELM. var. elliottii and P. taeda L. are irnportant exotics in Rhoidesia and each has desirable at- tributes. Reciprocal hybrid crosses were planned between eight clones from the breeding populations of each species. The croiss P. elliottii X P. taeda produced an average 25 viable seeds per cone whereas the reciprocal produced less than one. Progeny tests, each comprising 20 full-sib hybrid families between four P. elliottii female, and five P. taeda male parents, and nine open-pollinated half-sib non- hybrid families frolm the parents, were planted at three localities. An analysis of three-year height measurements showed that there was more within-family variation in the hybrid than there was in the no~n-hybrid families. General cambining ability variance remained high in the hybrids although there was a statistically significant amount of specific co~mbining ability associated with cer- tain clones. Over all families, heterosis was negative at the locality where envimnrnental conditions were most favour- able far growth and positive where conditions were mar- ginal for the two species. The prolspects for using hybrids between P. elliottii and P. taeda to assemble diverse pop- ulations for selection are golod and may have special value where the species are grown as exoitics when there may be increased chances of finding a "hybrid habitat". Table 1. - Analysis of variance, ranked means and Duncan Mul- tiple Range test of three-year height (m) for four half-sib P. ellio- ttii (E), five half-sib P. taeda (T) and 20 full-sib hybrid families at three localities. Key words: Pinus elliottii ENGELM., P. taeda L., hybrids, cross- ability, heterosis, hybrid habitat, general combining ability, specific combining ability. I Variance component 1 percent Source of variation Mean square Zusammenfassung Localities (L) Families (F) FxL Replications in L Error (pooled) Trees in plots Pinus elliottii ENGELM. var. elliottii und Pinus taeda I,. sind wirtschaftlich wichtige Baumarten in Rhdesien. Nach Kreuzung von 8 P. elliottii-Klonen und 8 P. taeda-Klonen in sag. Samenplantagen ko~mten bei den P. elliottii X P. taeda-Kombinatimen ungefähr 25 keimfähige Samen pro Zapfen erzielt werden, während in umgekehrter Richtung im Durchschnitt nur weniger als 1 Samen pro Zapfen zu erzielen war. Die dreijährigen Hybridnachkoimmenschaften aus dies~enKreuzungen wiesen im Höhenwachskum eine größere Variation auf als die beteiligten Kreuzungsfamilien innerhalb der jmeiligen Art. Heterosiseffekte zeigten sich nur dort, wo die Standortsbedingungen für beide Arten we- niger günstig waren. Repeatability of famiiy performance over locaiities = 78% Family Parents 8 Rank Mean height (m) and D.M.R.T. mixed T mixed T mixed T mixed T 8(T) 10(T) 10(T) 162(T) 13(T) 130) mixed T 8(T) 162(T) 10( T) 13(T) 10(T) 8(T) mixed E 162(T) mixed E mixed E 13(T) 8(T) mixed E 171(T) 162(T) 171(T) 171(T) 171(T) Introduction Many successful hybrids have been made amoag pines. Most species are hetermygous but heterosis in growth rate has been the exception ralther than the rule. Heterosis in fitness occurs moire frequently and a few interspecific cros- ses exhibiting this type of hybrid vigour are uised com- mercially. Although the expected potential of hybridiza- tim in the genus has noit been realised, most pine hybrids do tend to be viable and fertile and there are still pros- pects that further experimentation will lead to the pro- duction of valuable material. Pinus elliottii ENGELM. var. elliottii and P. taeda L. are both imprtant affo~restatian exotics in Rhodesia. Although their aptimum enviroinmental requirements are similar, each species has adaptive and morphological attributes which make it more suitabl'e than the other in certain I) Commonwealth Forestry Institute, Oxford, England. 2, Forest Research Centre, Salisbury, Rhodesia. Overall mean Standard error Silvae Genetica 27, Heft 6 (1978)

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Page 1: Three-year height performance of Pinus elliottii Engelm. var ......Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites

Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites in Rhodesia

By R. D. BARNES~) aad L. J. MULL IN^)

(Received December 1977 1 August 1978)

circumstances. P. elliottii is rnare drought resistant and also more tolerant of poior drainage than P. taeda. It is less severely affected by the group dying fungus, Rhizina in- flata (SCHAFF.) KARST., and less susceptible ta damage by fire. P. elliottii tends to have better stem and branch char- acteristics, allows more flexibility in management, its tim- ber is generally favourd by the trade and it is a high yielder of oleoresin. P. taeda on the other hand is more resistant to the pine woolly aphid, Pineus pini (L.), and to the honey fungus, Armillaria mellea (VAHL ex FR.) KUMMER. It captures the site more rapidly, is more tolerant of cold conditions and is much more resistant to windthrow and wind breakage at higher altitudes. P. taeda generally yields more timber than P. elliottii.

Breeding Programmes far P. elliottii and P. taeda were started in Rhoidesia in 1960. Populations of plus trees (su- perior phenotypes) were built up and controlled crosses were made for progeny tests. Both the incentive and the conditions were therefore present for experimentation with hybrids between these two closely related species and

Summary Pinus elliottii ENGELM. var. elliottii and P. taeda L. are

irnportant exotics in Rhoidesia and each has desirable at- tributes. Reciprocal hybrid crosses were planned between eight clones from the breeding populations of each species. The croiss P. elliottii X P. taeda produced an average 25 viable seeds per cone whereas the reciprocal produced less than one. Progeny tests, each comprising 20 full-sib hybrid families between four P. elliottii female, and five P. taeda male parents, and nine open-pollinated half-sib non- hybrid families frolm the parents, were planted at three localities. An analysis of three-year height measurements showed that there was more within-family variation in the hybrid than there was in the no~n-hybrid families. General cambining ability variance remained high in the hybrids although there was a statistically significant amount of specific co~mbining ability associated with cer- tain clones. Over all families, heterosis was negative at the locality where envimnrnental conditions were most favour- able far growth and positive where conditions were mar- ginal for the two species. The prolspects for using hybrids between P. elliottii and P. taeda to assemble diverse pop- ulations for selection are golod and may have special value where the species are grown as exoitics when there may be increased chances of finding a "hybrid habitat".

Table 1. - Analysis of variance, ranked means and Duncan Mul- tiple Range test of three-year height (m) for four half-sib P. ellio- ttii (E), five half-sib P. taeda (T) and 20 full-sib hybrid families at

three localities. Key words: Pinus elliottii ENGELM., P. taeda L., hybrids, cross-

ability, heterosis, hybrid habitat, general combining ability, specific combining ability.

I V a r i a n c e component 1 p e r c e n t S o u r c e o f v a r i a t i o n Mean s q u a r e

Zusammenfassung L o c a l i t i e s ( L ) F a m i l i e s ( F ) F x L R e p l i c a t i o n s i n L E r r o r ( p o o l e d ) T r e e s i n p l o t s

Pinus elliottii ENGELM. var. elliottii und Pinus taeda I,. sind wirtschaftlich wichtige Baumarten in Rhdesien. Nach Kreuzung von 8 P. elliottii-Klonen und 8 P. taeda-Klonen in sag. Samenplantagen ko~mten bei den P. elliottii X P. taeda-Kombinatimen ungefähr 25 keimfähige Samen pro Zapfen erzielt werden, während in umgekehrter Richtung im Durchschnitt nur weniger als 1 Samen pro Zapfen zu erzielen war. Die dreijährigen Hybridnachkoimmenschaften aus dies~en Kreuzungen wiesen im Höhenwachskum eine größere Variation auf als die beteiligten Kreuzungsfamilien innerhalb der jmeiligen Art. Heterosiseffekte zeigten sich nur dort, wo die Standortsbedingungen für beide Arten we- niger günstig waren.

R e p e a t a b i l i t y o f f a m i i y p e r f o r m a n c e o v e r l o c a i i t i e s = 78%

F a m i l y P a r e n t s

8

Rank Mean h e i g h t (m) a n d D.M.R.T.

mixed T mixed T mixed T mixed T

8 ( T ) 1 0 ( T ) 1 0 ( T )

1 6 2 ( T ) 1 3 ( T ) 1 3 0 )

mixed T 8 ( T )

162(T) 1 0 ( T ) 1 3 ( T ) 1 0 ( T )

8 ( T ) mixed E 162(T)

mixed E mixed E

1 3 ( T ) 8 ( T )

mixed E 1 7 1 ( T ) 1 6 2 ( T ) 171(T) 171(T) 171(T)

Introduction Many successful hybrids have been made amoag pines.

Most species are hetermygous but heterosis in growth rate has been the exception ralther than the rule. Heterosis in fitness occurs moire frequently and a few interspecific cros- ses exhibiting this type of hybrid vigour are uised com- mercially. Although the expected potential of hybridiza- t im in the genus has noit been realised, most pine hybrids do tend to be viable and fertile and there are still pros- pects that further experimentation will lead to the pro- duction of valuable material.

Pinus elliottii ENGELM. var. elliottii and P. taeda L. are both imprtant affo~restatian exotics in Rhodesia. Although their aptimum enviroinmental requirements are similar, each species has adaptive and morphological attributes which make it more suitabl'e than the other in certain

I) Commonwealth Forestry Institute, Oxford, England. 2, Forest Research Centre, Salisbury, Rhodesia.

O v e r a l l mean S t a n d a r d e r r o r

Silvae Genetica 27, Heft 6 (1978)

Page 2: Three-year height performance of Pinus elliottii Engelm. var ......Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites
Page 3: Three-year height performance of Pinus elliottii Engelm. var ......Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites
Page 4: Three-year height performance of Pinus elliottii Engelm. var ......Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites
Page 5: Three-year height performance of Pinus elliottii Engelm. var ......Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites
Page 6: Three-year height performance of Pinus elliottii Engelm. var ......Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites
Page 7: Three-year height performance of Pinus elliottii Engelm. var ......Three-year height performance of Pinus elliottii Engelm. var. elliottii X P. taeda L. hybrid families on three sites

and ANDERSON, E.: Spruce and pine racial hybrid variations in sented at the second meeting of the Working Group on Quantita- Northern Europe. In Papers presented a t the second meeting of tive Genetics. Raleigh, N.C., 18-19 August, 100-112 (1970). - STUBER, the Working Group on Quantitative Genetics. Raleigh, N.C. 18-19 C. W., and COCKERHAM, C. C.: Gene effects and variantes in hybrid August, 118-128 (1970). - SCHMITT, D. M.: Interspecific hybridization populations. Genetics 54, 1279-1286 (1966). - WOESSNER, R. A.: Inter- in forest trees: potential not realized. Proceedings of the 14th provenance crosses of loblolly pine. Proceedings of the 14th meet- meeting of the Canadian Tree Improvement Association, Part 2. ing of the Canadian Tree Improvement Association, Part 2. Fredericton, N.B., August 28-30, 17-23 (1973). - STUBER, C. W.: Fredericton, N.B., August 28-30, 17-23 (1973). Theory and use of hybrid population statistics. In Papers pre-

Computer organised orchard layouts (COOL) based on the permutated neighbourhood design concept

By G. D. BELL and A. M. FLETCHER

(Received May / August 1978)

Forestry Commission Research Division, Northern Research Station, Roslin, Midlothian

Summary

COOL is a computer program which produces seed or- chard layouts based on the permutated neighbourhood design colncept. The program is flexible, coping with irreg- ular areas, up to 100 clones, varying numbers of ramets per clone, and hybrid orchards. It is easy to use, being coa- trolled by a set of English command words, and can be operated without specialised statistical or computing knowledge. The use of COOL should eliminate much of the hard work involved in designing seed orchards. Key words: seed production, seed orchard design, tree breeding.

Zusammenfassung

Für die Planung und Anhge von Samenplantagen wird ein Computerprogra~mrn (COOL) vorgestellt, mit dem viele Schwierigkeiten bei der Erstellung von Samenplantagen bleiholben werden können. COOL ist einfa~ch zu handhaben

. und vielseitig anwendbar. Anlagepläne können auch für unregelmäßig geformte Flächen und für unterschiedlich große Klongruppen konstruiert werden.

Introduction

The first clonal seed orchard in Britain was planted in 1952. Several orchards of variolus species have been planted since then using a range of designs (FAULKNER 1965). In this eiarly period clonal components of the orchards had not been properly tested and the area of individual orchards was generally less than one hectare. Brmding Programmes for the main commercial species have now developed to the stage where most selected trees are in half-sib progeny tests an a nurnber of sites. Selection ob clones is based on a full consideration of data from these tests including a detailed analysis of any faimily X site interaction ( J O H N S ~ E m d SAMUEL 1978).

New clonal seetd orchards of the 1.5 generation type (WEIR

and ZOBEL 1975) are being established and before embarking on this Programme of orchards a set of basic rules was established.

1. On the basis of past experience, and, to keep the breed- ing population at an acceptable size in order to minirnise problems of relatednoss, the number of clones in any or- chard should be between 30 and 50. This number can be reduced by thinning as more information becolmes available general and specific cmbining ability, pm,portims of male and female flowers, dates of fliolwering and self-fertility levels.

2. The minimum area of an orchard should be four hec- tares (KOSKI 1975) to ensure satisfactory cross-pollination within the orchard.

3. Spacing should be related to the expected shape and development of the crowns for the species and the expected year of the first commercial coae crops.

4. Risk of self-pollinaticm should be minimised. 5. Theoretically all po~ssible crosses should occur with

similar frequency to obtain panmixis. 6. For ease of layout, management and the use of ma-

chinery for ground Cover coatrol, planting should be based on a Square or rectangular grid system.

Several types of design have k e n suggested in the past such as, shifting-clone design (MALAC 1962) or a modifica- t im of it as used in the southern states of Arnerica; cyclic balanced incomplete block designs (FREEMAN 1967; DYSON and FREEMAN 1968); most orchards however use simple ran- d~mised designs (GIERTYCH 1975).

GIERTYCH (1975) concidered a range olf orchard designs and coincluded that most tree breeders require minimal self- pollination and good panmixis from their designs. Several designs fulfill these two objectives to a greater or lesser degree but when other factors are considered a permutated neighbourhood design is the molst suitable. In commercial orchards fadors such as, compariscm of clones olr the divi- sim of an orchard into replicates for experimentation, are unimportant. The permutated neighbourhood design has not been widely used because it is difficult to coastruct without the aid of a computer, particularly when orchards are extensive andlor have mixtures of clones which are not evenly represented.

LA BASTIDE (1967) developed a computer program for permutated neighbourhood designs which allows all Cross- combinations to occur with approximately equal frequency and minimal celfing. However, the program is t m rigid and requires the orchard to be rectangular and have equal num- bers of ramets per clone.

Experience in Britain shows that prime seed orchard sites are difficult to obtain and are often irregular in shape. In additim the number of ramets available for each clone may not be constant due to varied grafting success, though in some cases varying numbers of ramets may be desirable due to imbalance in flower production between clones. An orchard design was required to take these factors into ac- count and LA BASTIDE having demonstrated the possibility of producing permutated neighbourhood designs by com- puter, it was decided to devebp a similar but more flexible program.

Silvae Genetica 27, Heft 6 (1978)