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Cold tolerance in warm season turfgrasses 

 

 

 

 

 

A thesis presented in partial fulfilment of the requirements 

for degree of 

 

Doctor of Philosophy 

in Plant Science 

 

at Massey University 

Palmerston North, New Zealand 

 

 

 

 

 
 

 

 

 

Umer Habib 
 

2017 
  



 

 

 
 

 



 

i 

Abstract 
 

Warm season (C4) turfgrasses are a popular choice for sports and public venues in 

tropical, subtropical, arid and semiarid climates due to their spreading characteristics, 

multiple stress resistance, including water deficit and heat tolerance, and faster 

establishment. However intolerance of low temperature is the key limitation to their use 

in temperate regions. The New Zealand turf industry has a growing interest in warm 

season (C4) grasses due to their water use efficiency under heat stress and summer 

dormancy of cool season (C3) grasses, especially in the upper parts of the North Island. 

Twelve commercially available cultivars of four warm season grass genera (Cynodon, 

Zoysia, Paspalum and Pennisetum) were established in a glasshouse and ten cultivars in 

field at Palmerston North, New Zealand, using seeds and stolon cuttings. This phase of 

the project was carried out from November 2012 to January 2015, with three major 

aspects of turf function measured. Established plots were scored for quality attributes 

(colour, texture, uniformity, ground cover and overall quality) as prescribed by NTEP 

(National Turf Evaluation Program, USA). Field plots became dormant and began 

browning in late autumn. Browning progressed and became more visible by the end of 

winter. Glasshouse plots displayed better overall turf quality than field plots except for 

seeded Cynodon varieties which showed susceptibility to Anthracnose fungal attack. 

Vegetative Cynodon varieties (Agridark, Windsor green and Santa Ana) performed well 

along with Sea spray (Paspalum vaginatum). Regal Staygreen (Pennisetum clandestinum) 

proved more cold tolerant than other varieties but, being coarse textured, cannot attain 

high acceptance in the turf industry. A subsequent experiment was focused on detailed 

morphology and growth pattern of these varieties. It was observed that glasshouse plots 

developed fewer roots per node and a lower total root mass compared with those grown 

in field conditions. In field plots stolon structures were more compact with a high number 

of horizontal stolons. Rhizome appearance differed between the glasshouse and the field  

and during the first year of establishment only vegetatively established Cynodon varieties 

developed rhizomes under field conditions and only Agridark in the glasshouse. 

However, during the next growing season all varieties in the field, except Zenith, had 

formed rhizomes. Seeded couches failed to produce rhizomes in the glasshouse even after 

their 2nd growing season.  Detailed study of stolon morphology confirmed findings on 

turf mat quality from visual scoring, and identified a pattern of ecological interest in that 



 

ii 

varieties of the genera Cynodon and Zoysia formed compound or triplet nodes, with root, 

branch and internode formation allocated to different leaves. 

A second phase of the research investigated cold tolerance in warm season turf 

grasses and the response of four varieties from three different warm season turf species 

Agridark and Windsor Green (Cynodon dactylon), Sea Spray (Paspalum vaginatum), and 

Zenith (Zoysia japonica) when exposed to low but non freezing temperatures. This 

experiment aimed to identify low temperature tolerance thresholds at various exposure 

durations, to help turf mangers define temperature tolerance of available varieties. Plants 

were established in trays in a glasshouse and were exposed to a series of progressively 

decreasing temperatures (16/10°C, 12/8°C, 10/6oC, 8/4°C and 6/2°C, day/night) with 2 

weeks at each temperature step, or to sudden, short exposure to the same temperatures for 

2 weeks. Colour change during the various combinations of low temperature exposure, 

and recovery after damage were observed along with measurement of selected 

physiological indices including proline, malondialdehyde (MDA) and carbohydrate 

accumulation. It was found that longer exposure with gradually lowered temperature was 

more detrimental to plants than sudden, short exposure. Seashore paspalm (Sea spray) 

exhibited better colour retention during cold exposure than the other three varieties in this 

experiment. Levels of proline and MDA in leaf and stolon tissue, and carbohydrate status 

tended to return towards pre-stress levels when plants were placed in a glasshouse for 

recovery from these cold-stress challenges.  

The ecological significance of the triplet stolon structure is unclear but deserves 

further study. Understanding that cold damage is a cumulative process rather than a 

sudden event when a threshold is reached, will be helpful to development of 

recommendations for turf industry use of C4 grasses in temperate climates. 

 
 
 
  



 

iii 

Acknowledgements 
 

I am thankful to almighty Allah for all his blessings and bounties in my life. I duly 

regard Higher Education Commission of Pakistan for providing me a scholarship and 

Massey University, New Zealand for providing me excellent learning environment and 

support to pursue my goals. I am thankful to PMAS Arid Agriculture University, 

Rawalpindi, Pakistan for granting me study leave. I am also thankful to the Institute of 

Agriculture and Environment (IAE), Massey University for providing me the opportunity 

to attend the International Horticultural Congress 2014 in Brisbane, Australia and 

scholarship support for completion of studies and to the New Zealand Sports Turf Institute 

for their collaboration and funding support for my project. 

I would like to express my deepest gratitude to my principal supervisor Professor 

Cory Matthew for all his kindness, guidance, patience, hard work and support throughout 

my study tenure. He provided the best learning environment and created excellent skill 

development opportunities for me. He is a true mentor and I feel myself fortunate having 

the opportunity to work with him. Undoubtedly I would not be able to achieve my goals 

without his kind support and supervision. 

I am also grateful to my co-supervisor Dr Andrew Mitchell, Research manager 

and Agronomist at NZSTI, Palmerston North for his technical insight and help during the 

project. He not only helped from his industry knowledge but also provided support to 

establish the experiments and their maintenance on a long-term basis. I wish to express 

my thanks to my co-supervisor Dr Kerry Harrington, Senior Lecturer, IAE, Massey 

University for his counselling time management suggestions and availability at short 

notice when help was needed. It was an enjoyable experience working with him. Special 

thanks are due to Bill Walmsley, Keith Salisbury, Leigh Hunt and Nick Jones for their 

support in acquiring germplasm and helping me to familiarise myself with New Zealand 

sports turf industry. 

My thanks are due to the staff members of the Plant Growth Unit, Massey 

University, Steve Ray, Lesley Taylor, and Lindsay Sylva for their help during my 

experiments in the glasshouse and controlled growth chamber. I was impressed with their 

problem solving capacity. I wish to express my thanks to Mark Osborne and Simon 

Orsborn for their technical and manual support. I am also thankful to Chris Rawlingson 



 

iv 

for his equipment tutorials and technical support and Kay Sinclair for helping with lab 

management and material acquisition, as and when it was required. I would like to 

appreciate help from Sunmeet Bhatia and Cécile Duranton and Roberta Carnevalli to 

carry out analyses for me. Very special thanks to the kind face of the Institute, Denise 

Stewart for her advice in making this manuscript look like a thesis. 

 I have to say special thanks to my PhD fellow Lulu He, for teaching me analysis 

skills and her guidance during my laboratory work. My heart felt gratitude also goes to 

fellow PhD students Wei Zhang, Januarius Gobilik and Mauricio Maldonado, and other 

graduate student colleagues. Time spent in New Zealand and Massey University has 

become much more valuable with lots of memories due to the presence of my sincere 

friends Muhammad Naveed Anwar and Ahmed Raza. I am thankful to Dr Muhammad 

Ajmal for his presence and support who was always like a big brother to me. I would also 

like to commend other Pakistani scholars and community members for their presence 

which never let me feel homesick. I am greatly thankful to Dr. Nadeem Akhtar Abbasi, 

Dean FCFS for being, boss, brother, friend and mentor; your encouragement and follow 

up will be always remembered. 

I greatly appreciate the contributions of my family to pursue my ambitions 

especially to my parents for their endless efforts to help in my life and my career. I am 

thankful to my Uncle Abdul Wahid for his mentoring and extraordinary support and love 

throughout my life. Most due love and gratitude to my sister Sadia Habib and brother 

Qasim Habib always like friends than siblings to me. 

At last I will like to express my thoughts and sincere thanks for my loving wife 

Fatima Mazhar, being with me encouraging and supporting as and when it was needed, 

and to my gorgeous little man Muhammad Danyal for all the joys and love. You both are 

absolute treasure for me. 

 

 
  



 

v 

Table of Contents 

 
Abstract i 

Acknowledgements ................................................................................................. iii 

Table of Contents ..................................................................................................... v 

List of Figures .......................................................................................................... x 

List of Tables......................................................................................................... xiii 

CHAPTER 1 ........................................................................................................... 1 

Introduction ............................................................................................................ 1 

1.1 Emergence of the modern turf industry ................................................ 1 

1.2 Thesis objectives ....................................................................................... 3 

1.2.1 Phase 1: Assessment of agronomic and mat forming properties of 

some commonly used warm season turfgrasses ................................ 3 

1.2.2 Phase 2: Response evaluation of selected varieties from Phase 1 

under controlled exposure to low temperature ................................ 4 

1.3 Thesis structure ........................................................................................ 4 

References ............................................................................................................... 6 

CHAPTER 2 ........................................................................................................... 9 

Review of Literature .............................................................................................. 9 

2.1 Grass to turfgrass ..................................................................................... 9 

2.2 Turfgrass classification .......................................................................... 11 

2.3 Warm season turfgrasses ...................................................................... 12 

2.3.1 Bermuda grass (Cynodon) ................................................................ 12 

2.3.2 Alternative species ............................................................................. 13 

2.4 Emergence of the global warm season turfgrass industry ................. 14 

2.4.1 Turfgrass evolution in the USA........................................................ 14 

2.4.2 The Australian turf industry ............................................................ 15 



 

vi 

2.4.3 The New Zealand turf industry ....................................................... 15 

2.5 Turfgrass quality attributes .................................................................. 17 

2.6 Turfgrass morphology ........................................................................... 18 

2.7 Temperature tolerance of cool and warm season grasses .................. 19 

2.8 Physiology of cold tolerance .................................................................. 22 

2.8.1 Carbohydrates ................................................................................... 22 

2.8.2 Proline ................................................................................................ 23 

2.8.3 Malondialdehyde (MDA) .................................................................. 23 

2.9 Experimental programme ..................................................................... 24 

References ............................................................................................................. 25 

CHAPTER 3 ......................................................................................................... 35 

Quality Attributes of Different Warm-Season Turfgrasses ............................. 35 

3.1 Introduction ............................................................................................ 35 

3.2 Materials and methods .......................................................................... 36 

3.2.1 Turf establishment ............................................................................ 36 

3.3 Results ..................................................................................................... 41 

3.3.1 Quality ................................................................................................ 42 

3.3.2 Ground cover ..................................................................................... 43 

3.3.3 Density ................................................................................................ 48 

3.3.4 Texture ............................................................................................... 48 

3.3.5 Colour ................................................................................................. 51 

3.4 Discussion ............................................................................................... 57 

3.4.1 Quality ................................................................................................ 57 

3.4.2 Groundcover ...................................................................................... 58 

3.4.3 Texture ............................................................................................... 58 

3.4.4 Colour ................................................................................................. 59 

3.5 Summary ................................................................................................. 60 



 

vii 

References ............................................................................................................. 62 

CHAPTER 4 ......................................................................................................... 65 

Morphological Attributes of Different Warm Season Turfgrasses ................. 65 

4.1 Introduction ............................................................................................ 65 

4.2 Materials and methods .......................................................................... 66 

4.2.1 Collection of turf cores ...................................................................... 67 

4.2.2 Data collection ................................................................................... 67 

4.2.3 Data analysis ...................................................................................... 67 

4.3 Results ..................................................................................................... 70 

4.3.1 Green mass, root mass and dead mass (thatch) .............................. 70 

4.3.2 Stolon length and rhizome length .................................................... 71 

4.3.3 Above ground attributes ................................................................... 77 

4.4 Discussion................................................................................................ 82 

4.4.1 Root mass, rhizome length and dead material (thatch) ................. 82 

4.4.2 Above -ground attributes.................................................................. 84 

4.5 Summary ................................................................................................. 86 

CHAPTER 5 ......................................................................................................... 91 

Stolon Morphology of Warm Season Turfgrasses ............................................ 91 

5.1 Introduction ............................................................................................ 91 

5.2 Materials and methods .......................................................................... 93 

5.2.1 Sample collection ............................................................................... 93 

5.2.2 Measurement ..................................................................................... 93 

5.2.3 Data analysis ...................................................................................... 94 

5.3 Results ..................................................................................................... 96 

5.3.1 Internode distance ............................................................................. 96 

5.3.2 Grass leaf attributes on a stolon ...................................................... 96 

5.3.3 Bud/branch and root occurrence ..................................................... 97 



 

viii 

5.3.4 Light microscopy ............................................................................... 97 

5.3.5 Principal component analysis ......................................................... 102 

5.4 Discussion ............................................................................................. 104 

5.4.1 Internode distance ........................................................................... 104 

5.4.2 Grass leaf attributes on a stolon .................................................... 104 

5.4.3 Bud/branch and root occurrence ................................................... 105 

5.4.4 Triplet morphology ......................................................................... 105 

5.5 Summary ............................................................................................... 106 

References ........................................................................................................... 108 

CHAPTER 6 ....................................................................................................... 111 

Evaluation of Cold Hardiness in Selected Warm Season Turfgrasses under 

Controlled Conditions........................................................................................ 111 

6.1 Introduction .......................................................................................... 111 

6.2 Materials and methods ........................................................................ 113 

6.2.1 Turf establishment .......................................................................... 113 

6.2.2 Experimental design........................................................................ 113 

6.2.3 Sample collection ............................................................................. 114 

6.2.4 Leaf proline contents....................................................................... 117 

6.2.5 MDA analysis ................................................................................... 117 

6.2.6 Carbohydrate content analysis ...................................................... 118 

6.2.7 Change in colour ............................................................................. 119 

6.2.8 Green up or recovery ...................................................................... 119 

6.2.9 Statistical analysis ........................................................................... 119 

6.3.1 Proline .............................................................................................. 121 

6.3.2 MDA ................................................................................................. 121 

6.3.3 Recalibration of the anthrone procedure for glucose and starch 124 

6.3.4 Low molecular weight sugars ......................................................... 124 



 

ix 

6.3.5 High molecular weight sugars ........................................................ 125 

6.3.5 Plant colour change during cold exposure and subsequent recovery

 129 

6.4 Discussion.............................................................................................. 131 

6.4.1 Proline .............................................................................................. 131 

6.4.2 MDA ................................................................................................. 131 

6.4.3 Carbohydrates ................................................................................. 132 

6.4.4 Discoloration .................................................................................... 133 

6.5 Conclusion ............................................................................................ 134 

References ........................................................................................................... 135 

CHAPTER 7 ....................................................................................................... 139 

General Discussion and Future Directions ...................................................... 139 

7.1 Introduction .......................................................................................... 139 

7.2  Turf establishment when using warm season grasses ...................... 140 

7.3 Qualitative traits of warm season turfgrasses ................................... 140 

7.4 Stolon and rhizome formation strategies ........................................... 141 

7.5 Phytomer specialisation in warm season turfgrasses ....................... 142 

7.6 Cold stress physiology.......................................................................... 143 

7.7 Key findings and their applications ................................................... 144 

7.8 Future research .................................................................................... 145 

References ............................................................................................................ 147 

INDEX TO APPENDICES .................................................................................. 149 

APPENDICES ..................................................................................................... 151 

 

 

  



 

x 

List of Figures 
 
 
Fig. 2.1 Structure of grasses with stolons and rhizomes (Christians, 

2011)  .   20 

Fig. 3.1 Establishment of field plots (left) and glasshouse plots (right) ......... 42 

Fig. 3.2 Established field plots (left) and glasshouse plots (right) in March 

2013. .................................................................................................. 42 

Fig. 3.3 Daily minimum/maximum temperatures in the field during summer, 

autumn, and winter in 2012-14. ........................................................ 46 

Fig. 3.4 Daily minimum/maximum temperatures in the glasshouse (GH) 

during 2012-13. ................................................................................. 47 

Fig: 3.5 Quality (overall visual appearance) of turf mat in glasshouse and open 

field conditions during Autumn/Winter 2013-2014. ......................... 48 

Fig: 3.6 Ground cover percentage of turf mat in glasshouse and open field 

conditions during Autumn/Winter 2013-2014. ................................. 49 

Fig: 3.7 Density of turf mat under glasshouse and open field during 

Autumn/Winter 2013-2014. .............................................................. 51 

Fig: 3.8 Texture of turf mat in glasshouse and open field conditions during 

Autumn/Winter 2013-2014. .............................................................. 52 

Fig: 3.9 Colour variations of turf mat in glasshouse and open field conditions 

during Autumn/Winter 2013-2014.. .................................................. 54 

Fig. 3.10 Progression in cold damage of experimental plots during 2013 under 

field conditions .................................................................................. 55 

Fig: 3.11 Differences in appearance of experimental plots from early autumn to 

mid-winter (2013) under glasshouse conditions. .............................. 56 

Fig: 3.12 Differences in appearance of experimental plots from early autumn to 

mid-winter (2013) under Field conditions. ....................................... 57 

Fig: 3.13 Visual status of field plots in third summer, showing recovery of 

colour after winter (October 2014) and retention of colour through 

summer 2014–2015. .......................................................................... 58 

Fig. 4.1 Extraction of turfgrass cores (A) and cling-film wrapped turfgrass 

cores (B). ........................................................................................... 70 

Fig. 4.2 A typical grass core dissection, in this case Regal Staygreen 



 

xi 

 (P. clandestinum) .............................................................................. 71 

Fig. 4.3 Dry weight comparison of green mass, dead mass and root mass in 

glasshouse and field plots - June, 2013. ............................................ 74 

Fig. 4.4 Dry weight comparison of green mass, dead mass and root mass in 

glasshouse and field plots - February, 2014. ..................................... 75 

Fig. 4.5 Comparison of stolon length and rhizome length in glasshouse and 

field plots June, 2013. ....................................................................... 76 

Fig. 4.6 Comparison of stolon length and rhizome length in glasshouse and 

field plots - February, 2014. .............................................................. 77 

Fig. 4.7 Comparative turf height at Field and Glasshouse (before core 

extraction).......................................................................................... 78 

Fig:4.8 Dry weight comparison of structural components (leaf, vertical 

branches, stems/stolons, roots, and rhizomes) in June 2013. ............ 81 

Fig:4.9 Dry weight comparison of structural components (leaf, vertical 

branches, stem/stolon, roots, and rhizome) in Feb 2014. .................. 82 

Fig 5.1 Dissected 12-phytomer stolon segments of A) Pennisetum (Regal 

Staygreen), and B) Cynodon (Windsor green) .................................. 96 

Fig.5.2 Stolon growth pattern of different turfgrass species on the basis of 

internode distance in twelve successive nodes (ID1-ID12) .............. 99 

Fig. 5.3 Comparison of leaf blade length, sheath length, and blade width in 

glasshouse and open field (average of twelve successive leaves on a 

stolon) .............................................................................................. 100 

Fig. 5.4 Total number of bud/branch and roots on a stolon for 12 successive 

nodes................................................................................................ 101 

Fig. 5.5 Triplet structure of a Cynodon stolon (Left), No grouping of internodes 

in Pennisetum (Right)...................................................................... 101 

Fig. 5.6 Triplet formation of nodes in (A) Cynodon, (B) Zoysia japonica .. 102 

Fig. 5.7 Plot of principal component scores for principal components 1 and 2 

for nine warm season turfgrass varieties in glasshouse and field 

plots. ................................................................................................ 104 

Fig. 5.8 Mean principal component scores for nine warm season turfgrass 

varieties from a principal component analysis designed to quantify 

morphological variation. ................................................................. 104 

Fig. 6.1 Induction and removal mechanism of turf trays in CTR ................. 116 



 

xii 

Fig 6.2 Turfgrass trays during establishment in the glasshouse .................. 116 

Fig 6.3 Established turf in the glasshouse ................................................... 116 

Fig 6.4 Turf trays in the controlled temperature room (CTR) ..................... 116 

Fig 6.5 Colour development for the anthrone reagent in the cross-calibration 

with glucose, starch and 

inulin.

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

 ......................................................................................................... 127 

 

 
 
 
  



 

xiii 

List of Tables 
 
Table 2.1 Important warm season turfgrass species with their uses and 

characteristics ............................................................................... 14 

Table 3.1 Warm-season turfgrass varieties, sources establishment method, 

and seed rate ................................................................................. 40 

Table 3.2 Randomized layout plan .............................................................. 41 

Table 5.1 Eigenvalues, proportion of variation explained, and coefficients for 

principal components 1 to 3 of a principal component analysis 

designed to detect differences in morphological pattern between 

varieties. ..................................................................................... 103 

Table 6.1 Table of treatments as followed during the experiment ............ 115 

Table 6.2 Proline accumulations (mg g–1 of dry matter) in turfgrasses after 

sudden and prolonged exposure to low temperature. ................ 122 

Table 6.3 MDA accumulations (nmol g–1 of dry matter) in turfgrasses after 

sudden and prolonged exposure to low temperature. ................ 123 

Table 6.4 Variations in Low molecular sugar levels (mg g–1 of dry matter) in 

turfgrasses after sudden and prolonged exposure to low 

temperature. ............................................................................... 127 

Table 6.5 Variations in high molecular sugar levels (mg g–1 of dry matter) in 

turfgrasses after sudden and prolonged exposure to low 

temperature. ............................................................................... 128 

Table 6.6 Degree of colour change during and after exposure to low 

temperature based on % age and colour characteristics ............ 130 

  



 

xiv 

 

 
 

 
 
 



 

1 

CHAPTER 1 

Introduction 
 

1.1 Emergence of the modern turf industry 

Turfgrass is an essential component of landscaped ecosystems and symbolizes 

beauty and utility in integration with plants and other features (Roberts et al., 1992). Turf 

is grown at various venues (such as lawns at residential units, public and community 

parks, sports fields, city green belts, golf courses and many commercial properties) for 

functional, recreational and aesthetic benefits (Duble, 1996). Turfgrass enhances aesthetic 

appeal of a landscape, stimulates mental health with positive societal impact, public unity, 

enhanced productivity and enhanced living standards (Kaplan and Kaplan, 1989; Ulrich, 

1986). Sports activities are believed to contribute significantly to societal health in the 

modern world (Stiles et al., 2009). Turfgrass provides an economical surface for 

recreation and outdoor sports. The unique cushioning effect provided by turfgrass 

minimizes the risk of injuries to players in games like soccer and rugby (Orchard, 2002; 

Beard & Green, 1994). Therefore a large proportion of the area under turf comprises 

sports and athletic fields (Morris, 2003). 

Turfgrasses are separated into cool season (C3) grasses and warm season (C4) 

grasses based on their photosynthetic pathways (Christians, 2011), and the two groups 

also have distinct ecological distributions and temperature adaptations. Cool season 

grasses include among others the genera Agrostis, Festuca and Lolium, historically 

widely used in New Zealand (NZ). This group is often considered to be resource hungry 

and to have comparatively high maintenance and input requirements. Susceptibility to 

heat stress is a basic limitation of cool season or C3 grasses in warmer climates. C3 grasses 

exhibit low water use efficiency due to increased evapotranspiration rates under high 

temperatures resulting in a water imbalance that in more extreme conditions can lead to 

dehydration (Jiang and Huang, 2001). 

Warm season turfgrasses primarily belong to the genera Cynodon, Eremochloa, 

Paspalum, Pennisetum, and Zoysia (Busey, 1989) and members of these genera typically 



 

2 

have horizontal creeping stems, resulting in a spreading growth habit. Warm season 

turfgrasses are widely grown in tropical regions of the world and even sometimes in 

temperate regions, where hot summers are experienced (Geren et al., 2009). They exhibit 

higher levels of heat and drought tolerance due to their morphology, growth pattern, 

internal cell metabolism and photosynthetic efficiency (Zhou and Abaraha, 2007). 

Intolerance of low temperature is a limiting factor for most of the warm season grass 

species (Thomas et al., 2009). Cold temperature damage in warm season turfgrasses 

lowers the aesthetic appeal. Their growth usually stops below 16ºC, while foliage starts 

turning brown below 10ºC (Anderson et al., 2002). 

  Cynodon (often referred to by the common names Bermuda grass, or couch grass) 

is the most widely cultivated genus among the warm season group of grasses (Taliaferro, 

2003) and is extensively used for golf courses and athletic surfaces (Munshaw et al., 

2006) in southern USA and many other countries worldwide where summer temperatures 

exceed 25°C. The most commonly encountered species is C. dactylon (L.) Pers., but C. 

transvaalensis Burtt Davy with a number of sterile hybrid crosses of these two species, 

are also used in the industry. Other genera, especially Zoysia and Paspalum are now 

attracting industry attention as some members of these genera display particular 

adaptations such as shade and salinity tolerance (Duncan, 1996). 

The turf industry typically produces and manages lawns and other land areas for 

landscape, public and recreational purposes (Nutter, 1965). The industry started to 

develop in its present structure after World War II and grew dramatically in the next three 

decades (Ralph and Busey, 1987). The turf industry as an entity evolved radically during 

the latter half of 20th century with USA becoming the hub for research and development. 

The United States Department of Agriculture (USDA) and the United States Golf 

Association (USGA) were two of the main organizations that influenced this 

transformation of the pre-existing traditional turf culture (Jenkins, 1994). Research into 

warm season turf species is now well documented in scientific literature from USA, China 

and Australia, to name three countries that have strong links with New Zealand, but also 

in many other countries in Europe, Asia, Africa and South America. Turf breeders seek 

to develop turfgrass cultivars that can offer acceptable growth and surface quality under 

a wide range of atmospheric and soil conditions. The ultimate goal for any turfgrass 

breeding programme is the development of cultivars with smaller shoot size than forage 



 

3 

grasses, and with the capacity to form a finely structured turf. A turf variety should also 

have reduced maintenance requirements, but ability to tolerate biotic (disease, pest, 

traffic) and abiotic (drought, cold, heat, salinity) stress factors (Duncan and Carrow, 1999; 

Lee et al., 2004). 

The turf industry in New Zealand was founded on cool season grasses from the 

United Kingdom (UK), particularly species of Agrostis, Festuca, Lolium, and Poa. The 

presence of warm season turfgrass species was noted in the 19th and early 20th centuries 

(C. dactylon – 1871, Paspalum dilatatum Poir. – 1896, Pennisetum clandestinum Hochst. 

Ex. Chiov.  – 1940) (Field and Forde, 1990), and P. clandestinum in particular has become 

spontaneously dominant in household lawns in many coastal and northern areas of New 

Zealand. However, there was generally little interest in using C4 grasses in sown turf 

initially. With increasing urbanisation and indications of a warming climate, the period 

since 2000 has seen an increased interest in warm season turfgrasses (C4) by the NZ turf 

industry, especially for the more northern areas of the North Island, due to their drought 

tolerance and high water use efficiency. However, warm season turfgrasses are often 

intolerant of low temperature and susceptible to winter injury. 

1.2 Thesis objectives 

This project was designed with a primary interest to evaluate warm season 

turfgrass varieties available in NZ for their cold tolerance characteristics. Data collection 

was targeted towards describing aspects of the growth morphology and physiology of 

selected warm season turfgrasses. Envisaged outcomes were development of 

management practices for the New Zealand turf industry to reduce the occurrence of 

problems such as winter browning in warm season turfgrasses. The project was a joint 

venture between the Institute of Agriculture and Environment, Massey University and the 

New Zealand Sports Turf Institute (NZSTI), to evaluate a range of warm season turf 

species for their turf characteristics and cold tolerance under glasshouse and field 

conditions. The project comprised two phases: 

1.2.1 Phase 1: Assessment of agronomic and mat forming properties of 

some commonly used warm season turfgrasses  

The core objectives of the study were:  



 

4 

• To assess the mat quality for a range of warm season turfgrasses protected from 

and exposed to frost and evaluation of each variety in terms of visual attributes. 

• To compare different cultivars for their agronomic/morphological characteristics 

under frost free (glasshouse) and frost exposed (field) growing conditions. 

1.2.2 Phase 2: Response evaluation of selected varieties from Phase 1 under 

controlled exposure to low temperature 

A follow-up experiment was designed to explore detailed physiological responses to 

different levels of cold exposure to selected varieties from the first phase. Key aims were 

to: 

• Explore the level of cold tolerance among selected warm season turfgrasses under 

controlled conditions to precisely define their damage threshold levels for industry 

extension purposes. 

• To develop an understanding about morphological changes occurring during cold 

exposure and comparison of discoloration and re-greening rates. 

• To evaluate different physiological changes (e.g., levels of sugars) taking place in 

plants during cold exposure for an improved understanding of the physiology of 

cold tolerance. 

• To determine the rate of recovery from cold stress (“green up”) under optimal 

growing conditions (spring). 

1.3 Thesis structure 

The thesis comprises seven chapters. Following this brief introduction, Chapter 2 

provides a review of literature with insights into turf culture and history focused towards 

warm season turfgrasses and their potential use in the New Zealand turf industry. 

Turfgrass qualitative and quantitative attributes and cold stress physiology are also 

explored in Chapter 2. 

Chapter 3 is the first data chapter and provides qualitative assessment of turf 

quality traits of the investigated warm season turfgrass species (10 cultivars from 5 



 

5 

species) selected for initial evaluation. Chapter 4 is complementary to Chapter 3 and 

presents data describing above and below ground growth and morphology of selected 

turfgrass varieties under the two different growth environments investigated (glasshouse 

and field).  

In view of the C4 grass creeping growth habit being so different from the erect 

tufted growth habit of the C3 grasses traditionally used in New Zealand, Chapter 4 and 

Chapter 5 provide a detailed investigation of C4 turfgrass morphology. Chapter 4 includes 

a combination of quantitatively measured and visually observed characteristics. In the 

course of measurements in Chapter 4 it was noted that there appeared to be different plant 

anatomy (Niklas and Kutschera, 2009) in certain genera for the organisation of phytomers 

in the construction of stolons. As this fundamental morphology difference has been only 

occasionally and briefly reported in the literature, morphology data to explore this point 

were collected and are presented in Chapter 5. 

Chapter 6 describes an experiment designed to explore temperature thresholds at 

which chilling damage occurs. The chilling damage experiment included selected 

cultivars from those evaluated in the turf quality and plant morphology studies reported 

in Chapters 3, 4, and 5. Plant growth dynamics, leaf colour change, and some 

physiological status indicators were evaluated at various points in a regime of 

progressively decreasing temperatures.  

The thesis concludes with a brief summary of the main findings and a general 

discussion of future research requirements in Chapter 7. 

  



 

6 

References 

Anderson, J. A., Taliaferro, C. M., Martin, D. L. (2002) Freeze tolerance of Bermuda 

grasses: Vegetatively propagated cultivars intended for fairway and putting green 

use, and seed-propagated cultivars. Crop Science 42: 975–977. 

Beard, J. B., Green, R. L. (1994) The role of turfgrasses in environmental protection and 

their benefits to humans. Journal of Environmental Quality 23: 452–460. 

Busey, P. (1989) Progress and benefits to humanity from breeding warm-season grasses 

for turf. Contributions from Breeding Forage and Turfgrasses 14: 49–70. 

Christians, N. (2011) Fundamentals of Turfgrass Management (4th ed.). John Willey and 

sons, Inc. Hoboken, New Jersey, USA. 

Duble, R. L. 1996. Turfgrasses, Their Management and Use in the Southern Zone. Texas 

A&M University Press, College Station, TX. 

Duncan, R. R., Carrow, R. N. (1999) Turfgrass molecular genetic improvement for 

biotic/edaphic stress resistance. Advances in Agronomy 67: 233–305.  

Duncan, R. R. (1996) The environmentally sound turfgrass of the future. USGA Green 

Section Record 34: 9–11. 

Field, T. R. O., & Forde, M. B. (1990). Effects of climate warming on the distribution of 

C4 grasses in New Zealand. Proceedings of the New Zealand Grassland 

Association 55: 47–50. 

Geren, H., Avcioglu, R., Curaoglu, M. (2009) Performances of some warm-season 

turfgrasses under Mediterranean conditions. African Journal of Biotechnology 8: 

4469–4474. 

Jenkins, V. (1994). The lawn: a history of an American obsession. Smithsonian Institution 

press, Washington DC. 

Jiang, Y., & Huang, B. (2001). Drought and heat stress injury to two cool-season 

turfgrasses in relation to antioxidant metabolism and lipid peroxidation. Crop 

Science 41: 436–442. 



 

7 

Kaplan, R., Kaplan, S. (1989) The Experience of Nature: A Psychological Perspective. 

Cambridge University Press, New York. 

Lee, G. J., Carrow, R. N., Duncan, R. R. (2004) Salinity tolerance of selected seashore 

paspalums and bermudagrasses: root and verdure responses and criteria. 

Horticulture Science 39: 1136–1142. 

Morris, K. N. (2003) The National Turfgrass Research Initiative. National Turfgrass 

Federation, Inc. National Turfgrass Evaluation Program, Beltsville Agricultural 

Research Center, Beltsville, Maryland, 20705 USA.  

Munshaw, G. C., Ervin, E. H., Shang, C., Askew, S. D., Zhang, X., Lemus, R. W. (2006) 

Influence of late season iron, nitrogen and seaweed extract on fall color retention 

and cold tolerance of bermudagrass cultivars. Crop Science 46: 273–283. 

Niklas, K.J., Kutschera, U. (2009) The evolutionary development of plant body plans. 

Functional Plant Biology 36: 682–695. 

Nutter, G. C., Watson, J. R. (1969). The turfgrass industry. Turfgrass Science, In Hansan, 

A. A., Juska F.V. (Eds.). Turfgrass science. (9–26) American Society of 

Agronomy, Wisconsin, USA. 

Orchard, J. (2002) Is there a relationship between ground and climatic conditions and 

injuries in football?. Sports Medicine 32: 419–432. 

White, R. W., Busey, P. (1987) History of turfgrass production in Florida. 

In Proceedings of Florida State Horticulture Society 100: 167–174. 

Stiles, V. H., James, I. T., Dixon, S. J., Guisasola, I. N. (2009) Natural Turf Surfaces. 

Sports Medicine 39: 65–84. 

Taliaferro, C. M. (2003) Bermudagrasses. In Casler, M. D., & Duncan, R R. (Eds.). 

Turfgrass biology, genetics, and breeding. (235–256) Hoboken, N.J.: John Wiley. 

Thomas, R. S., Rodgers, C. A., VanDyke, R., Williams, D. W., and Phillips, T. D. (2009) 

The Inheritance of Cold Tolerance and Turf Traits in a Seeded Bermuda grass 

Population. Crop Science 49: 1489–1495. 

http://link.springer.com/journal/40279


 

8 

Ulrich, 1986 Ulrich, R. S. (1986) Human responses to vegetation and landscapes. 

Landscape and Urban Planning 13: 29–44. http://dx.doi.org/10.1016/0169-

2046(86)90005 

Zhou, S., Abaraha, A. (2007) Response to heat stress in warm season and cool season 

turfgrass cultivars. Scientific Research and Essays 2: 95–100. 

 

  

http://dx.doi.org/10.1016/0169-2046(86)90005
http://dx.doi.org/10.1016/0169-2046(86)90005


 

9 

CHAPTER 2 

Review of Literature 
 

2.1 Grass to turfgrass 

Grasses belong to a major group within the flowering plants, or Angiosperms, that 

are known as monocotyledons because there is only a single embryonic first leaf or 

cotyledon inside the seed.  More than 10,000 individual species of grasses exist in the 

plant kingdom, and they exhibit a great diversity in form, growth and habitat (Christians, 

2011), including among others, maize, wheat, rice, and the bamboos. Turfgrasses are 

recognized for a compact horizontal growth with mat forming ability, and are capable of 

maintaining a high density when subject to regular mowing and significant treading, 

traffic, or other forms of wear (Roberts et al., 1992). Almost 50 grass species worldwide 

fit these criteria (Christians, 2011).   

The historic use of grasses is referred to in many parts of the Bible (Roberts et al., 

1992). Some artists’ depictions of the “Garden of Eden”, though tree-dominated, also 

show a green surface covered with grass like vegetation. The modern concept of turf can 

be traced back to early domestication of plants and animals when grazed lands were used 

as natural playgrounds. Historical records hint at development of vast and luxurious 

pleasure gardens around imperial palaces in China about 157–87 B.C. (Huffine and Grau, 

1969). Similar development of gardens has been reported after this era in Persia, India 

and Europe (Spain, Italy, and France). However the emergence of ball games (about 5000 

years ago) has had more influence on the evolution of turfgrasses to their current form. 

The awareness of turfgrass husbandry practices grew rapidly after 15th century with 

development of golf (Casler and Duncan, 2003; Roberts et al., 1992).  

Modern turfgrass varieties emerged through three major forces of selection, 

described by Darwin as natural, unconscious, and methodical selection (Casler and 

Duncan, 2003). Biotic and abiotic stress factors help grasses to develop and colonize over 

a large fraction of Earth’s terrestrial surface. Species and local ecotypes with natural 

adaptation to conditions in the geographic region where they occur, including resistance 



 

10 

to cold, salt or drought stress have emerged independently of any human activity (Duncan 

and Carrow, 1999). By developing such adaptations, grasses have colonised a range of 

habitats including coastal strips, alpine slopes, river plains, deserts and wetlands and in 

each case have developed specific adaptations to prevailing conditions. A good example, 

is North African germplasm of tall fescue (Schedonorus arundinaceus (Schreb.) Dumort 

and cocksfoot Dactylis glomerata L. which tends to have a summer dormancy that is 

recognised as a water conservation adaptation to help overcome summer moisture deficit 

(Volaire and Norton, 2006, and references therein).  

The unconscious selection process involves the early domestication of livestock 

and grasses. Perennial grasses present in grazed lands display a number of defence 

strategies including dwarfism, axillary bud utilisation to generate a branching growth 

habit, and the presence of rhizomes and stolons which both facilitate spreading and 

protect plant tissue from consumption by browsing animals (Casler and Duncan, 2003). 

Continued defoliation of grazing lands is assumed to have resulted in a shift towards a 

short stature and high tiller density (Roberts et al., 1992). Besides defoliation by grazing 

animals,  other factors such as trampling and fire would have contributed selection 

pressure, so that ecotypes subject to these selection pressures ultimately evolved the 

ability to grow as a mat of intertwined stems on the soil surface (Busey, 1977).  

Methodical selection processes were initially based upon selection and propagation of 

wild cultivars with required characters. Selections from old turf grounds were used as 

planting material with a broad adaptation to certain stress factors. These selections were 

then passed through recurrent selection processes to choose the best individual cultivars 

(Casler and Duncan, 2003; Moore and Moser, 1995).  

Western civilizations in the early eighteenth century domesticated wild grass 

species for turf purposes (Bormann, 1993). Cool season grass genera like Agrostis, 

Festuca, Lolium and Poa are widely reported as earlier selections from grazed lands in 

different regions of Europe and North America (Beard, 1998; Busey 1989). European 

botanists collected and successfully disseminated materials from colonized areas 

resulting in naturalization of non-native, long living, turf species (Duncan and Carrow, 

1999). In New Zealand anecdotal information available in turf industry circles suggests 

that as recently as the 1980s, many sports ground managers were using the same seed 

lines of browntop (Agrostis capillaris L.), as were being sold to hill country sheep farmers 



 

11 

for pasture use, for new sowings of cricket wickets and outfields and local body parks 

and reserves grounds used for weekend sports.  

In the second half of the 20th century, breeding of turfgrasses was initiated by 

agronomists working on pasture grasses (Seagle and Iverson, 2002). Turf breeders are 

trying to develop turfgrass cultivars that can offer acceptable growth and surface quality 

for a wide scenario of atmospheric and soil conditions. Dwarf cultivars with ability to 

tolerate biotic (disease, pest, traffic) and abiotic (drought, cold, heat, salinity) stress 

factors with less maintenance requirements are the ultimate goals for any breeding 

program (Lee et al., 2004). In New Zealand, ‘Grasslands Egmont’ browntop and 

‘Grasslands Cook’ Chewing fescue are examples of such selection of pasture grasses 

aimed at producing varieties specifically for turf use (Rumball and Robinson, 1982; 

Rumball, 1982). This point is further discussed below. 

2.2 Turfgrass classification 

From a turf industry perspective, adaptability to cold climate is a major abiotic 

stress tolerance factor helpful in classification of turfgrasses and defining their 

distribution in different regions (Casler, 2006). On this basis turfgrasses are often grouped 

within turf industry circles into cool season grasses and warm season grasses, also known 

respectively as C3 and C4 grasses. The cool season grasses are mostly of European 

geographic origin and have the C3 photosynthetic pathway, while the warm season 

grasses are typically of tropical or subtropical geographic origin and possess the C4 

photosynthetic pathway (Christians, 2011). Hence, the two groups also have distinct 

ecological distributions and temperature adaptations. The C3 and C4 terminology is 

derived from the identity of the first photosynthetic products which are 3-carbon and 4-

carbon molecules respectively (Ehleringer and Cerling, 2002). These two photosynthetic 

pathways respond differently to available atmospheric CO2. C4 plants achieve a higher 

photosynthetic rate and resource utilization than C3 plants (Shay & Kubien, 2013) by 

increased CO2 concentration around the enzyme Rubisco. The C4 pathway is an energy 

expensive pathway but eliminates photorespiration, and also results in reduced stomatal 

conductance, giving a strong advantage for drought tolerance, (Sage, 2004; Sage and 

McKown, 2006). The CO2 enrichment process of C4 plants enables them to synthesize 

2.4 g sugar compared with 1.9 g in C3 plants for each MJ of light energy intercepted by 

the plant’s leaf surface area (Christians, 2011), while the reduced stomatal conductance 



 

12 

and lower water use allow C4 plants to tolerate high temperatures (>20°C) better than C3 

plants, and maintain growth longer in water deficit conditions before stored soil moisture 

is depleted. Alternatively, under irrigation, C4 grass turf has lower water requirement than 

a C3 grass turf, which can be an attractive consideration when planning maintenance of a 

playing surface or golf course in a modern city, since a saving of 100 mm irrigation water 

applied in a growing season translates to 1ML ha–1.  

Taxonomically, all grasses are members of the botanical family Poaceae, and the 

commonly used cool season turfgrasses are members of the Poeae tribe within that family, 

while the major warm season turfgrasses are members of the Paniceae.     

2.3 Warm season turfgrasses 

The warm season turfgrasses are widely distributed throughout the regions with 

warmer temperatures including, humid, sub-humid, arid and semi-arid climates. (Beard, 

1973). Optimum air temperature suitable for warm season turfgrasses ranges between 17–

35ºC (Fry and Huang, 2004). They tend to become dormant when temperature drops 

below 10ºC or during heavy frosting (Beard, 1982). Despite this limited tolerance of cool 

temperatures, they are often found in regions with a continental climate, where they thrive 

in the warm conditions of summer but cease leaf production and lose colour in winter. 

Important warm season turfgrasses include, but are not limited to species of the genera 

Cynodon, Zoysia, Buchloe, Paspalum, Pennisetum, Eremochloa, and Stenotaphrum (Fry 

and Huang, 2004; Juska et al., 1969). Most of the C4 grasses with high functional quality 

grown for sports turf surfaces are propagated vegetatively (ie using sprigs or cuttings 

obtained from an established turf) rather than by seed (Geren et al., 2009). For species 

where both vegetatively propagated and seeded varieties are available, the vegetatively 

propagated varieties are generally considered within industry circles to be superior in 

performance, compared to seeded varieties. 

2.3.1 Bermuda grass (Cynodon) 

Bermuda grass is the most widely grown and adopted warm season turf species. 

It is commonly known as couch grass, green couch grass and Indian doab (Teliaferro, 

2003). Many commercial selections and seeded varieties are available for a range of 

climates and specific uses with good variation in form and habitat. Bermuda grass 



 

13 

originated and evolved in Eastern Africa. Biotic and abiotic stresses including 

environment and grazing pressure from larger herds of African wild mammals resulted in 

the grass to evolving a deep root system with plenty of lateral growth (stolons) and 

tramping resistance  (Beard, 1998). The genus Cynodon has nine different species with 

greater genetic variation. C. dactylon (L.) Pers. and C. transvaalensis Burtt Davy are the 

important turf types and most of the commercial varieties are crosses between these two 

species, which are generally sterile and need to be vegetatively propagated (McCarty and 

Miller, 2002).  

Table 2.1 Important warm season turfgrass species with their uses and 

characteristics (Extracted from Casler, 2006). 

Common name Latin name Region Traits Uses 

 

https://doi.org/10.1017/S0021859606006137 

 

2.3.2 Alternative species 

Zoysia grass is another very important member of the warm season turfgrasses. 

This genus is known to be native from western Pacific Rim along the western coasts of 

Indian Ocean. This genus comprises 11 known species with a distribution map spreading 

from New Zealand to Japan and French Polynesia through to Mauritius. Species of Zoysia 

grass (Zoysia japonica Steud., Zoysia matrella (L.) Merr., Zoysia pacifica (Goudsw.) M. 

Hotta & Kuroki) are used in the turf industry, especially in the Southeast Asian tropics, 

in countries such as Malaysia and Indonesia. Among the Zoysia species, Z. japonica is 

considered most tolerant to low temperature stress and to have a superior shade tolerance 

compared to Cynodon (Zhang et al., 2009; Engelke and Anderson, 2003). Zoysia has a 



 

14 

good reputation in the turf industry as it has good turf quality characters and low 

maintenance requirements compared to some other warm season grasses (White et al., 

2001). 

Paspalum vaginatum Sw. or seashore paspalum is a more recent inclusion in lists 

of turfgrass species, predicted to become much more widely used in the turf industry in 

the 21st century (Duncan and Carrow, 2000). The grass is attracting attention among turf 

scientists and industry stakeholders as it displays resistance to multiple abiotic stresses, 

including soil salinity, drought, cold and shade (comparable to Bermuda grass) tolerance 

(Duncan, 2003). 

2.4 Emergence of the global warm season turfgrass industry  

As defined by Watson et al., 1992: the “Turfgrass industry in its broader aspect is 

a group of specialized individuals and organizations sharing their common interest in 

production, development and maintenance of green space”. The turf industries in various 

countries are dynamic in nature and their definition encompasses various factors 

including geographic location, scale of operation and demand for products, manpower, 

investment and governmental policies (Nutter, 1965). The origins of the turf industry are 

closely aligned with the history of sports turf usage particularly with the ball games; 

initially golf (Watson et. al., 1992), and more recently sports such as football. Evolution 

of the turf industry was slow in the first half of the 20th century but very steady (Beard, 

1982). However, it has experienced tremendous growth during the latter half of 20th 

century, encompassing the farming and maintenance of targeted species for 

environmental, aesthetic and recreational utilities (Shearman, 2006). The international 

turf industry is expected to grow and change rapidly during the 21st century, with an 

increased emphasis on environmental protection and best management practices (Seagle 

and Iverson, 2002).  

2.4.1 Turfgrass evolution in the USA 

Turfgrasses gained their current status as products of an established industry with 

an extensive range of private, private, commercial, and government users only during the 

past five decades in USA (Roberts et al., 1992). At present turfgrass is the fourth largest 

‘crop’ in USA covering more than 50 million acres (Milesi et al., 2005) with an estimated 



 

15 

annual economic turnover of up to 60 billion USD. It is estimated that 2% of total land 

area in USA is under turf cover (Milesi et al., 2005) and it has been estimated that more 

than 90% of American citizens use or come into contact with turfgrass, in some way, in 

their daily life. 

2.4.2 The Australian turf industry 

The key developmental period for the Australian turf industry is considered to 

have been between World Wars I and II. Rapid urbanization and the increased need for 

sophisticated and well developed sports centres are considered the main drivers of this 

expansion (Loch et al., 2016). According to the Turf Producers Association (TPA) around 

4,400 hectares of turf sod is produced for sale, with a total annual value of $300 million 

and the Turf maintenance sector has over $500 million annual turnover. Queensland’s 

share of the total turf production is 38%, New South Wales has a 33% share, and Victoria 

and Western Australia account for 15 and 11% of production, respectively (TPA 2017). 

A survey conducted during 2006-2007 showed that about 85% of the total area under turf 

production in Australia comprised warm season turfgrass species, of which couches 

(Cynodon sp.), buffalo grass (Buchloe dactyloides (Nutt.) Engelm) and Kikuyu 

(Pennisetum clandestinum Hochst. ex Chiov.) make up 91% (Haydu et al., 2008). 

2.4.3 The New Zealand turf industry 

The New Zealand turf industry is also growing in volume very rapidly. Many 

public institutes and private organizations are working in research, training, production 

and maintenance sectors of the industry. According to a study conducted in New Zealand 

during 2006, an estimated area of 122,328 hectares was planted in turf, of which schools 

comprise about 48% (58,217 ha). Council administered areas were estimated at 30% 

(36,326 ha) followed by 20% (25,361 ha) for golf courses. Approximately 48% (58,139 

ha, previously reported 60,000 ha in 2000) of the total area under turf was managed for 

sports and amenity use only (Haydu et al., 2008; Haydu et al., 2006; Way, 2001).  

The New Zealand sports turf industry has been assessed as having $33 billion 

worth of resources including land, buildings and equipment. Schools were estimated as 

the largest stakeholder comprising 70% of total assets followed by city councils at 20% 

and golf courses at 6% respectively. Almost 94% of turf industry business is dependent 

on domestic customers. In 2006, the New Zealand turf industry was reported to have a 



 

16 

financial turnover of $NZ 356.6 million for operational expenditure. The major expense 

was the salaries of 24,000 people employed by the industry under different categories 

(NZSTITO, 2011). According to a recent report by SPARC (Sports and Recreation New 

Zealand), 92% of young people and 83% of adults surveyed indicated active participation 

in sports (Dalziel, 2011). 

New Zealand turf culture is dominated by cool season grasses. Settlers from 

United Kingdom introduced amenity grasses early in 20th century (e.g. species of 

Agrostis, Festuca and Lolium). Initially browntop (Agrostis capillaris L.) and fine fescues 

(Festuca rubra L.) were extensively used to establish amenity and sports areas. Festuca 

rubra was introduced in the 19th century, and a growth form lacking creeping rhizomes 

and with high shoot density, now known as F. rubra subsp. commutata, was identified on 

a farm in the South Island, owned by a Mr Chewings, who sold seed and promoted it as 

a superior type. The variety was widely sown around New Zealand, and significant 

volumes of seed were exported to Europe and USA, where it remains well known in turf 

industry circles (Rumball, 1983). The name Chewings fescue is now widely recognised 

in Europe and the USA, and even listed in Merriam-Webster online dictionary 

(https://www.merriam-webster.com/dictionary/chewings%20fescue). The first New 

Zealand breeding program for cool season turfgrasses was started at the Department of 

Scientific and Industrial Research, Grasslands Division (DSIR Grasslands, now 

AgResearch) in 1973 and they released five varieties of cool season grasses, developed 

by selection of pasture types for shoot density and short stature. Chewings fescue was one 

of these. Descriptive notes on ‘Grasslands Cook’ Chewings fescue were published by 

Rumball (1982), and the wider turfgrass development programme at that time is described 

by Rumball (1983). However, the DSIR Grasslands turf varieties were not widely adopted 

by the turf industry, and during the 1980s many groundkeepers in New Zealand began 

importing planting material from USA and Europe. Additionally, from the 1990s 

establishment of a plant variety rights scheme in New Zealand facilitated the development 

of commercial breeding of turf varieties. PGG Wrightson is a Christchurch-based 

company that has been active in turfgrass breeding and evaluation, with interest centred 

on but not confined to the Australasian market.  

In the last two decades, the New Zealand turf industry has developed an interest 

in warm season grasses due to summer dormancy and poor heat and moisture deficit 



 

17 

tolerance of cool season grasses, especially in the upper parts of the North Island. Kikuyu 

(P. clandestinum) was introduced to New Zealand in the 20th century to improve summer 

pasture production in northern North Island regions. In these areas, Kikuyu has been 

found to spontaneously displace C3 grass species and is now widespread as a major 

species in many urban lawns and roadside berms, but because of its coarse growth habit 

it has not generally been used in turf applications. One of the warm season turf species 

commonly employed in New Zealand is Indian doab (C. dactylon), also known as “couch” 

in the turf industry. Interest in this species is especially strong for golf courses and sports 

fields.  Seeded (Princess-77, Yukon, Southern Star, La Paloma) and non-seeded couches 

(Agridark, Windsor Green, Legend, Santa Ana) are available for the turf market in New 

Zealand (Hunt, 2011). However seeded varieties of Z. japonica (Zenith and Compadre), 

P. vaginatum (Sea Spray) and P. clandestinum (Regal Staygreen) are also available in the 

market. Warm season turf species, being efficient in water and nutrient use, in addition 

to their high temperature tolerance, can potentially have a large future role in the New 

Zealand sports turf industry. 

2.5 Turfgrass quality attributes 

The aesthetic appearance of turf venues is a high priority for turf managers, and 

is often demanded by users as well, even in those situations where there is an intense use 

or unfavourable weather conditions. Very objective criteria are required for determining 

the grass type to be employed under any particular condition, in order to match varieties 

established with environmental and soil conditions and available resources (Carrow et al., 

2010). Quality assessment or visual field assessment in turfgrasses is a type of ranking 

system that varies with species, season, growing conditions and the evaluator (Krans and 

Morris, 2007). 

Turfgrass quality is the combination of qualitative traits and visual appearance. 

Turfgrass quality is measured as the combined visual effect of density, uniformity, 

texture, smoothness, growth habit and colour (Beard and Beard, 2005). A scale widely 

used for visual field assessment in turfgrasses is scoring at 1–9 where 1 is the poorest 

denomination and 9 as the best (Krans and Morris, 2007). Turfgrass evaluators judge the 

turfgrass quality on the basis of visual observations. Turfgrass quality assessment ratings 

differ from individual to individual. It is recommended that only one evaluator should 

perform the procedure throughout one study (Bell et al., 2009; Morris, 2003). Qualitative 



 

18 

ratings of turfgrasses vary to a large extent depending upon variety/species, seasonal 

changes, cultural and management practices (Morris and Shearman, 2014). 

2.6 Turfgrass morphology 

All grasses share a segmental morphology whereby growth units called phytomers 

are continuously formed at an apical meristem (Sharman, 1947), and progress through a 

series of development phases producing first a leaf, then a bud within the leaf axil which 

may initiate to form a branch. There may or may not be elongation of the stem between 

successive leaves. Leaves normally undergo programmed senescence after a defined life 

span, at which point root primordia may form and generate adventitious roots, which take 

several leaf appearance intervals to fully develop. Unlike familiar plants such as shrubs 

or trees where the seedling stem persists for the life of the plant and expands by secondary 

thickening; in grasses the constancy of shoot form over time comes from coordinated 

cycling of the component phytomers through their development stages. The ‘stem’ may 

either be a ‘pseudostem’ which is actually a whorl of leaves with the youngest emerging 

from the centre and the oldest shed from the exterior when the leaf dies, or where 

internode elongation has occurred the stem may be a stolon on or just above the soil 

surface or an underground rhizome. There are many studies of grass morphology, often 

in the context of optimising herbage intake by animals. Among these, for ryegrass, 

Silsbury (1970) has described the coordination of phytomers within the grass shoot or 

tiller, while Fulkerson and Donaghy (2001) described leaf turnover, noting that there are 

typically three live leaves at any one time in a ryegrass shoot. A comprehensive study of 

the fate of individual phytomers in Poa pratensis plants over a series of years was 

published by Etter (1951). The author is not aware of similar studies of grass morphology 

at the phytomer level in C4 grasses. 

Warm season turfgrasses primarily belong to the genera Cynodon, Eremochloa, 

Paspalum, Pennisetum, and Zoysia (Busey, 1989) and members of these genera typically 

display a spreading nature, since the internode regions of their phytomers are normally 

elongated to form stolons or rhizomes. In modern turf breeding many commercial C4 

grass cultivars do not have the capacity to produce seeds, often as a result of development 

through interspecies hybridisation or artificially induced polyploidy. Turf managers 

frequently report that seeded varieties exhibit poor mat quality, an open growth habit and 



 

19 

less traffic resistance, compared to vegetatively propagating varieties of the same species 

(Geren et al., 2009). 

 

Fig. 2.1 Structure of grasses with stolons and rhizomes (from Christians, 2011). 

 

2.7 Temperature tolerance of cool and warm season grasses 

Cool season grasses have an optimal growth temperature ranging from 16°C to 

24°C, temperatures, and above this range results in decreased metabolism (Du et al., 

2009). With extremely high temperatures and drought conditions they undergo dormancy, 

cease to grow and turn brown in colour (Beard, 1994).  

The C4 pathway naturally occurs mainly close to the equator under warm and 

sunny conditions. It involves complex metabolic and anatomical differences, compared 

to C3 plants, which allow efficient water use and retain productive capacity under high 

light intensity and temperatures (Beard, 1998). Warm season or C4 turfgrasses are widely 

grown in tropical regions of the world and even sometimes in temperate regions, where 

hot summers are experienced (Geren et al., 2009). The C4 grasses are able to tolerate high 

temperatures (i.e. warmer maximum summer temperatures ranging from 25 – 38°C) and 

water deficiency very well. Tolerance to low temperature is one major limiting factor for 

most of the warm season turf species (Thomas et al., 2009). Cold temperature damage in 

warm season turfgrasses lowers their aesthetic appeal. Their growth usually stops below 

16°C, while foliage starts turning brown below 10°C (Anderson et al., 2002). 



 

20 

Since playing fields and parks are key amenities in urban infrastructure, the need 

for a sustainable turf surface always keeps managers under pressure. The tension between 

the desire for scheduled events to go ahead without cancellation, and the need to avoid 

damage and maintain visual appearance, is a central consideration nowadays when 

developing maintenance practices (Cisar, 2004).  Damaged or dormant grasses lose their 

aesthetic and environmental values and are considered undesirable in sports and athletic 

fields. The costs of redevelopment of injured or dead turf can be a significant burden on 

municipal maintenance budgets (Anderson et al., 2007). There is, therefore, an increasing 

interest in superior varieties capable of survival and growth under both extremes of 

temperature. Turf breeders have made substantial progress with increasing stress 

tolerance in warm season grasses (Anderson et al., 2007). Research work with warm 

season turfgrasses has been carried out in many countries but mainly concentrated in 

USA, Australia and China.  

Lack of tolerance to chilling temperatures is the most often investigated 

performance trait for selection and usage of warm season species (Bermuda grass, Zoysia 

and seashore paspalum) under cooler climates (Stefaniak et al., 2009; Kopec et al., 2007; 

Casler, 2006; Anderson et al., 2003; Beard, 1982).  Warm season grasses are usually 

considered to be intolerant to both sudden and prolonged exposure to low temperatures.  

Low temperature exposure caused by rapid temperature drop (frosting) has been referred 

to as ‘direct’ cold exposure, and exposure to low temperatures for a prolonged duration 

is termed ‘indirect’ cold exposure (Fry, 1990). Cold temperature exposure results in 

chilling injury leading to dormancy, chlorosis and necrosis, and such damage is followed 

by death in more sensitive species. Frost kill occurs because of disruption of cell 

membranes and other components due to expansion on freezing of solutes. Overall 

reduction in chlorophyll accumulation, photosynthesis, metabolite translocation, enzyme 

activity and protein synthesis can be observed (Sanghera et al., 2011; Samala et al., 1998; 

Salisbury and Ross, 1992). 

Winter field performance is a typical assessment method for cold hardiness in 

grasses. One measure of cold hardiness is the LT50 (LT denotes lethal temperature), which 

refers to the low temperature exposure that will kill 50% of plants in a grass population 

(Zhang et al., 2009). Uncertainty over what kinds of fluctuations in temperature may be 

most damaging to plants limits the interpretive power of field experiments. In vitro 



 

21 

experiments using controlled environment cabinets and freeze chambers provide better 

information and understanding of cold stress mechanisms (Anderson et al., 2005).  

Cynodon (Bermuda grass, couch grass) is the most common and widespread genus 

among the warm season group of grasses (Taliaferro, 2003) and is extensively used for 

golf courses and athletic surfaces. However Cynodon varieties often lack winter hardiness 

and so are vulnerable to frost damage or winter kill (Munshaw et al., 2006). The cold 

tolerance level of different Bermuda grass cultivars was evaluated by Anderson et al. 

(2002) using a laboratory freeze method and found a great variation in tolerance levels 

based on cultivar type and usage. Varieties particularly recommended for fairways were 

more tolerant compared with those used for greens.  

Anderson et al. (2007) observed that vegetatively propagated Bermuda grass had 

a lower LT50 (–6.2°C to –11.3°C) than seeded varieties (–5.3°C to –8.7°C). Similarly 

Zhang et al., 2006 compared two seeded cultivars of Cynodon (Riviera and Princess-77) 

for their level of cold tolerance. LT50 was lower (–8.3°C) for Riviera compared to 

Princess–77 (–6.3°C). Tolerance to cold, while partly mediated by physiological factors, 

is also influenced by grass morphology. A positive correlation can be found with stolon 

length, stolon size and rhizome length (Ahring et al., 1975). Bermuda grass cultivars with 

increased cold tolerance are mostly selections of clonal (vegetatively propagating) 

material from cooler regions which were further hybridized with superior mat forming 

cultivars (Samudio and Brede, 2002). 

As noted above, Zoysia is one of the genera of C4 grsses used in the turf industry. 

Species, with the three species most commonly used by the industry being (Z. japonica, 

Z. matrella, and Z. pacifica. Z. japonica is considered most tolerant to low temperature 

stress and to have a superior shade tolerance compared to Cynodon (Zhang et al., 2009; 

Engelke and Anderson, 2003). Exposure to air temperatures below 10°C is problematic 

for common Z. japonica and can cause leaf discoloration and increase in carbohydrate 

and proline contents (Wei et al., 2008). Patton and Reicher, 2007 evaluated in vitro freeze 

tolerance of different Zoysia varieties, and in that work a cultivar Diamond displayed 

inferior performance (LT50 = –8.4°C) compared with Meyer and Zenith (LT50 = –11.5°C). 



 

22 

2.8 Physiology of cold tolerance 

Intolerance to low temperature is the key factor limiting the use of warm season 

(C4) grasses (Bush, 2000). Warm season grasses start to turn dormant when exposed to 

temperatures below 15°C followed by browning of the leaves at 10°C (Beard, 1973). This 

intolerance can cause chilling injury (discoloration, chlorosis, and necrosis) known as 

winter browning. Low temperature thresholds vary greatly among species and varieties 

depending upon genetic and environment interactions (Stier, 2007).  

The circumstances where damage is triggered rapidly below a certain threshold 

temperature or occurs slowly and cumulatively in response to temperatures that would 

not be low enough to cause immediate symptoms, and where a protective effect from 

‘conditioning’ may occur, are not well clarified in the existing research literature. It is 

generally assumed that gradual but prolonged exposure would not prove as lethal as a 

sudden freezing exposure. However, there are reports that continuous exposure to low 

temperature above the sudden damage threshold can kill a significant proportion of an 

existing population. Low temperature exposure, whether sudden or gradual, results in  

several physiological and morphological changes in the plants, leading to reduced 

transpiration rates, changes in concentration or induction of particular enzymes, 

production of proteins related to stress tolerance and carbohydrate translocation (Bocian 

et al., 2011; Hisano et al., 2008). Prior exposure to non-lethal low temperatures (cold 

acclimation) triggers the expression of many physiological and metabolic responses and 

enables the plant to cope with cold stress (Cyril, 2002; Dionne, 2001). 

2.8.1 Carbohydrates 

An increase in non-structural carbohydrate concentration is often correlated with 

low temperature tolerance in warm season turfgrasses, and this effect involves substances 

responsible for membrane stabilization and limiting crystallization. Concentrations of 

soluble carbohydrates in buffalo grass vary significantly in samples taken at different 

times of the year (Bell et al., 2002). There is a strong association between higher 

concentrations of non-structural carbohydrates and cold tolerance (Huang et al., 2014; 

Bush et al., 2000). DiPaola and Beard (1992) have also reported higher levels of sugars 

in Zoysia compared with Cynodon, and linked this to improved survival. Different 



 

23 

varieties of Cynodon also showed variation in total non-structural carbohydrate contents 

during and after cold acclimation (Zhang et al., 2006).  

2.8.2 Proline 

C4 grasses are usually susceptible to chilling injury (1°C ̶ 12°C), which may cause 

membrane damage and loss to plant proteomic activity, decreased photosynthesis and 

respiration (Stier, 2007). Proline is an amino acid widely studied as an indicator of 

response to abiotic stress, mainly drought and cold stress. Many scientists have reported 

an increase in proline concentration during exposure to low temperature in their 

experiments performed under various conditions on warm season turfgrasses (Munshaw 

et al., 2006; Zhang et al., 2006 & 2011). 

 Increased proline content in turfgrasses is often correlated with increased cold 

tolerance. Patton et al. (2007) studied 13 different genotypes of Zoysia and find out their 

difference in cold tolerance in relation to proline and carbohydrate accumulation. They 

concluded that proline levels increase in Zoysia during cold acclimation but it was found 

that proline is less beneficial to plants as temperature drops. Kauffman (2010) found 

Diamond as the most hardy variety when compared with TifEagle, Champion, and 

SeaDwarf and reported an increase in proline contents. Patton et al. (2007) further 

confirmed that a rise in carbohydrate and proline contents help plants to tolerate cold 

temperatures. 

2.8.3 Malondialdehyde (MDA) 

The cell membrane is a primary site of cold injury and any alteration in its 

composition can influence cell metabolism and fluid exchange, and especially a 

transformation to a rigid state can reduce its permeability (Huang et al., 2014). Increase 

in fatty acid desaturation (FAD) is related to improved cold tolerance in plants (Wang et 

al., 2003) and cold susceptible plants increase lipid desaturation during cold acclimation 

for better membrane permeability (Taiz and Zeiger, 1991). Malondialdehyde (MDA) 

formation as a response to lipid peroxidation is considered a precursor to environmental 

stress (Wang et al., 2010). With prolonged exposure to chilling stress, MDA levels tend 

to increase in leaves (Wang et al., 2009).  



 

24 

2.9 Experimental programme 

Considering the above discussion, a research programme was planned that would 

first provide data relevant to greenkeepers and grounds supervisors in New Zealand on 

the performance in the field at Palmerston North of a selection of warm season turfgrass 

species and varieties available in New Zealand. An intended outcome of the planned 

research was to identify, from the results of that study, research questions specific to 

performance of warm season turfgrasses in New Zealand that could be the focus of some 

detailed follow up research.  

  



 

25 

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