Malus Sylvestris Research Articles

Responses of ‘Golden Delicious’ Apples to 1-MCP Applied in Air or Water

The efficacy of the ethylene action inhibitor 1-methylcyclopropene (1-MCP) applied in water to slow ripening of ‘Golden Delicious’ [Malus sylvestris var. domestica (Borkh.) Mansf.] apples was evaluated in comparison with 1-MCP applied in air. The material was applied by dipping fruit in 1-MCP water solutions (0.03, 0.3, or 3 mmol·m−3) for 4 min or by exposing fruit to 1-MCP gas (0.42, 4.2, or 42 μmol·m−3) in air for 12 h. Fruit were held in air at 20 °C for 25 days after treatment or stored at 0.5 °C in air for up to 6 months followed by 7 days in air at 20 °C. Application of 1-MCP in water or air delayed the increase in respiration and ethylene production associated with fruit ripening and reduced the amount of fruit softening, loss of acidity, and change in peel color. Treatments applied in water required a 700-fold higher amount of active ingredient compared with treatments applied in air to induce similar physiological responses. Fruit responses to 1-MCP varied with treatment concentration, and the maximum effects were obtained at concentrations of 4.2 or 42 μmol·m−3 in air and 3 mmol·m−3 in water. Peel color change was impacted less than retention of firmness and titratable acidity for 1-MCP treatments applied at concentrations of 4.2 or 42 μmol·m−3 in air and 0.3 or 3 mmol·m−3 in water. Treatment with 1-MCP in air or water was less effective for slowing peel degreening when treated fruit were stored at 0.5 °C compared with storage at 20 °C. Fruit treated with 1-MCP and stored in air at 0.5 °C developed a peel disorder typified by a gray·brown discoloration that is unlike other disorders previously reported for this cultivar. Symptoms were present when fruit were removed from cold storage and no change in symptom appearance was observed during a 7-d holding period at 20 °C.

External carbon dioxide injury and 1-methylcyclopropene (1-MCP) in the ‘Empire’ apple

The effects of several factors on the susceptibility of untreated and 1-MCP-treated ‘Empire’ apple ( Malus sylvestris (L.) Mill var. domestica (Borkh.) Mansf.) fruit to external CO 2 injury have been investigated. These factors include CO 2 partial pressure, the timing of elevated CO 2 exposure, delays between harvest and exposure to elevated CO 2, DPA concentration, and the timing of DPA treatment after exposure of fruit to 1-MCP. 1-MCP-treated fruit were more susceptible to external CO 2 injury than untreated fruit when stored in 5 kPa, but not 1 kPa, CO 2 (in 2 kPa O 2). 1-MCP did not increase the period of highest susceptibility to injury during controlled atmosphere storage. The greatest sensitivity to injury occurred 0–3 weeks after harvest. Sensitivity to injury decreased when untreated fruit were kept in air for up to 14 days before exposure to 5 kPa CO 2, but not for 1-MCP-treated fruit. DPA treatment prevented development of CO 2 injury even at a level as low as 250 μL L −1. DPA treatment could be delayed for 4 days after 1-MCP treatment while fruit were exposed to 5 kPa CO 2 in air without injury development. Overall the data indicate that 1-MCP enhances the risk of external CO 2 injury, but this injury can be prevented by treatment of fruit with DPA at harvest. In the absence of DPA treatment, handling procedures such as maintaining low CO 2 partial pressures in the storage atmosphere must be used.

Flavonoids from leaves of Pyrus communis, Malus sylvestris, and Malus domestica

Flavonoids from leaves of Pyrus communis, Malus sylvestris, and Malus domestica

Investigation of Changes in Phytohormone Levels Depending on Effects of Exogenous Indole-Butyric-Acid and Callus Formation in the Stem Cuttings of Some Apple Kinds (Malus sylvestris Miller)

Investigation of Changes in Phytohormone Levels Depending on Effects of Exogenous Indole-Butyric-Acid and Callus Formation in the Stem Cuttings of Some Apple Kinds (Malus sylvestris Miller)

Salt stress-mediated changes in free polyamine titers and expression of genes responsible for polyamine biosynthesis of apple in vitro shoots

In the present paper, changes in contents of free polyamines and expression of genes responsible for polyamine biosynthesis of apple [ Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] in vitro shoots exposed to 100 mM or 200 mM NaCl for 10 days were investigated. Salt stress led to severe growth retardation, based on electrolyte leakage (EL), net increase in fresh weight and shoot length. Na concentration in salt-treated shoots was significantly higher than that of the control, and K was unexpectedly enhanced, particularly by 100 mM NaCl. Three commonly occurring polyamines, putrescine (Put), spermidine (SPd) and spermine (Spm), were detected in the shoots, in which Spd and Spm were the most and the least abundant part, respectively. Both types of salt stress led to significant reduction of free Put, and Spm was significantly reduced by 100 mM NaCl, whereas Spd underwent negligible fluctuation. The genes encoding for polyamine biosynthetic enzymes showed diverse expression patterns in response to salt treatment. Salt stress led to induction of MdADC, MdSAMDC1, MdSPDS1 and MdSPMS, whereas it appreciably inhibited transcript level of MdACL5 in a concentration-dependent manner, indicating that the genes were differentially affected by salt stress. The data presented herein showed that salt stress led to conspicuous alteration of free polyamine contents and that mRNA levels of some genes involved in polyamine biosynthesis were strongly influenced. Mechanism underlying the decrease in free Put and gene expression patterns following long-term salt stress were discussed.

Sampling Damage to Tree Fine Roots: Comparing Air Excavation and Hydropneumatic Elutriation

Air excavation is commonly used to expose structural roots of trees, but its suitability for sampling fine roots for research has not been closely examined. We compared damage and root-diameter class distribution in roots sampled by air excavation with roots sampled by hydropneumatic elutriation of soil cores. We collected samples from six different tree species with a range of fine root diameters: Amur corktree (Phellodendron amurense Rupr.), apple [Malus sylvestris var. domestica (Borkh.) Mansf.], river birch (Betula nigra L.), striped maple (Acer pensylvanicum L.), swamp white oak (Quercus bicolor Willd.), and western redcedar (Thuja plicata D. Don). Root-diameter class distributions for each species were the same for samples collected by either air excavation or by elutriation. Median root diameter was greatest for Amur corktree and western redcedar (≈0.5 mm), intermediate in striped maple and oak, and least in river birch and apple (≈0.2 mm). Root damage was primarily due to loss of root tips. Although species varied in their susceptibility to root damage and whether air excavation caused more damage than elutriation, root diameter was not a good predictor of damage during sampling. Air excavation caused ≈26% greater damage to root samples of river birch and western redcedar than did elutriation. Both sampling methods caused equivalent root damage in all other species. Root anatomy influenced susceptibility to damage during sampling. Epifluorescence microscopy revealed a root hypodermis in all species except Amur corktree and western redcedar. Without the mechanical support of this suberized layer, the cortex of Amur corktree was easily stripped from the stele, leading to extensive damage by both sampling methods. Hydropneumatic root elutriation conferred some protection to roots relative to air excavation (in two of six species); final choice of root sampling method must depend upon the requirements of the individual study and characteristics of the site.

Cold Storage to Control Codling Moth Larvae in Fresh Apples

Codling moth [Cydia pomonella (Lepidoptera: Tortricidae)], found in exported apples (Malus sylvestris), can disrupt international markets. Cold storage at 1.1 °C was examined for possible control of three physiological larval states in ‘Fuji’ apples: diapausing (overwintering), diapause-destined, and nondiapausing. All nondiapausing larvae were dead within 12 weeks, diapaused-destined larvae were controlled by the seventh week, yet more than half of the original populations of diapausing larvae were still alive after 11 weeks. Because the diapaused-destined larvae were younger than the nondiapausing larvae, they may have been more susceptible to cold. Because larvae normally diapause outside the fruit, cold storage would not be applicable for controlling larvae in this state.

Linked vs. unlinked markers: multilocus microsatellite haplotype‐sharing as a tool to estimate gene flow and introgression

We have explored the use of multilocus microsatellite haplotypes to study introgression from cultivated (Malus domestica) into wild apple (Malus sylvestris), and to study gene flow among remnant populations of M. sylvestris. A haplotype consisted of alleles at microsatellite loci along one chromosome. As destruction of haplotypes through recombination occurs much faster than loss of alleles due to genetic drift, the lifespan of a multilocus haplotype is much shorter than that of the underlying alleles. When different populations share the same haplotype, this may indicate recent gene flow between populations. Similarly, haplotypes shared between two species would be a strong signal for introgression. As the expected lifespan of a haplotype depends on the strength of the linkage, the length [in centiMorgans (cM)] of the haplotype shared contains information on the number of generations passed. This application of shared haplotypes is distinct from using haplotype-sharing to detect association between markers and a certain trait. We inferred haplotypes for four to eight microsatellite loci on Linkage Group 10 of apple from genotype data using the program phase, and then identified those haplotypes shared between populations and species. Compared with a Bayesian analysis of unlinked microsatellite loci using the program structure, haplotype-sharing detected a partially different set of putative hybrids. Cultivated haplotypes present in M. sylvestris were short (< 1.5 cM), indicating that introgression had taken place many generations ago, except for two Belgian plants that contained a haplotype of 47.1 cM, indicating recent introgression. In the estimation of gene flow, F(ST) based on unlinked loci indicated small (0.032-0.058) but statistically significant differentiation between some populations only. However, various M. sylvestris haplotypes were shared in nearly all pairwise comparisons of populations, and their length indicated recent gene flow. Hence, all Dutch populations should be considered as one conservation unit. The added value of using sharing of multilocus microsatellite haplotypes as a source of population genetic information is discussed.

Gymnosporangium spp. (non‐European)

<i>Gymnosporangium</i> spp. (non‐European)

RADIO FREQUENCY-HOT WATER DIPS FOR POSTHARVEST CODLING MOTH CONTROL IN APPLES

A combination radio frequency-hot water dip method was examined as a potential quarantine treatment against fifth instars of the codling moth, Cydia pomonella L. (Lepidoptera: Tortricidae), in apples, Malus sylvestris (L.) var. domestica (Borkh.) Mansf., which were intended for export to Japan. The apples were initially exposed to 27.12-MHz radio frequency energy at 12 kW for 2.75 min and were then submerged in a range of hot water dips (48‐50C) for different durations. Efficacious tests were at 48C for >2 h, at 49C for >50 min and at 50C for >40 min. Fruit quality tests indicated that the best hot water parameters were at 50C for 40 min. Fruit quality after 2 weeks was cultivar dependent where “Fuji” apples tolerated heat treatment better than “Delicious” and “Gala” apples. None of the treated fruits were acceptable after 60 days. Regardless of cultivar, heat treatment resulted in loss of both peel and fresh colors, coupled with reduced firmness and increased external and internal damage.

RADIO FREQUENCY PULSE APPLICATION FOR HEATING UNIFORMITY IN POSTHARVEST CODLING MOTH (LEPIDOPTERA: TORTRICIDAE) CONTROL OF FRESH APPLES (MALUS DOMESTICA BORKH.)

ABSTRACT A radio frequency (RF) treatment, with a pulse mode to increase heating uniformity, was examined as a potential quarantine treatment against fifth instars of the codling moth, Cydia pomonella L. (Lepidoptera: Tortricidae), in apples, Malus sylvestris L. var. domestica (Borkh.) Mansf., intended for export to Japan. Apples were exposed in water baths to 27.12‐MHz RF energy at 12 kW with a pulse mode of 30 s‐on/30 s‐off for different durations. The temperature uniformity was determined by measurements at 12 sites evenly spaced at two depths in the fruit. The most promising exposure time was 29 min. The efficacy of the treatment with the pulse mode was examined against the codling moth larvae by using exposures between 27 and 30 min and up to 30 min of holding time in bath after treatment. The treatment that killed all larvae with the least amount of energy was the 29‐min exposure with 50‐min holding time. This treatment was appraised for fruit quality in “Delicious” apples. Some injury was observed when treated fruits were removed from 1C storage after 30 days. When these fruits were held at 25C for 1 week, all were too damaged to evaluate for fruit quality. Although pulse mode increased heating uniformity in the fruit, the thermal requirements to control codling moth larvae may exceed the injury threshold of the fruits.

Analysis of Genetic Diversity of Processing Apple Varieties

Genetic diversity of 18 processing apple varieties and two fresh varieties were evaluated using 12 simple sequence repeats (SSR) primer pairs previously identified in Malus domestica Borkh. A total of 87 alleles in 10 loci were detected using 10 polymorphic SSR markers selected within the range of 5–14 alleles per locus. All the 20 varieties could be distinguished using two primer pairs and they were divided into four groups using cluster analysis. The genetic similarity (GS) of groups analyzed using cluster analysis varied from 0.14 to 0.83. High acid variety Avrolles separated from other varieties with GS less than 0.42. The second group contained Longfeng and Dolgo from Northeast of China, the inherited genes of Chinese crab apple. The five cider varieties with high tannin contents, namely, Dabinette, Frequin rouge, Kermerrien, M.Menard, and D.Coetligne were clustered into the third group. The fourth group was mainly composed of 12 juice and fresh varieties. Principal coordinate analysis (PCO) also divided all the varieties into four groups. Juice and fresh apple varieties, Longfeng and Dolgo were clustered together, respectively, using both the analyses. Both the analyses showed there was much difference between cider and juice varieties, cider and fresh varieties, as well as Chinese crab apple and western European crab apple, whereas juice varieties and fresh varieties had a similar genetic background. The genetic diversity and differentiation could be sufficiently reflected by combining the two analytical methods.

Nitric Oxide and Nitrite Treatments Reduce Ethylene Evolution from Apple Fruit Disks

`Golden Delicious' apple [Malus sylvestris var. domestica (Borkh.)] cortex disks suspended in solutions containing a nitric oxide (•NO) donor [S-nitrosoglutathione (GSNO) or sodium nitroprusside (SNP)], •NO gas, or nitrite (KNO2) were used to identify impacts of •NO on ethylene production and NO2– on •NO and ethylene production. Treatment with GSNO or SNP reduced ethylene biosynthesis compared with control treatments containing equimolar concentrations of oxidized glutathione (GSSG) or Na4(CN)6 respectively. Apple disk exposure to •NO gas did not impact ethylene production. Treatment with NO2– resulted in increased •NO production and decreased ethylene biosynthesis. Generation of •NO increased linearly whereas ethylene generation decreased exponentially with increasing NO2– treatment concentration. •NO was enhanced in autoclaved tissue disks treated with NO2–, suggesting that its production is produced at least in part by nonenzymatic means. Although this evidence shows •NO is readily generated in apple fruit disks by NO2– treatment, and ethylene synthesis is reduced by •NO/NO2– generated in solution, the exact nature of •NO generation from NO2– and ethylene synthesis modulation in apple fruit disks remains to be elucidated.

Polyamine biosynthesis of apple callus under salt stress: importance of the arginine decarboxylase pathway in stress response

To clarify the involvement of the arginine decarboxylase (ADC) pathway in the salt stress response, the polyamine titre, putrescine biosynthetic gene expression, and enzyme activities were investigated in apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] in vitro callus under salt stress, during recovery after stress, and when ADC was inhibited by D-arginine, an inhibitor of ADC. Salt stress (200 mM NaCl) caused an increase in thiobarbituric acid-reactive substances (TBARS) and electrolyte leakage (EL) of the callus, which was accompanied by an increase in free putrescine content, during 7 d of treatment. Conjugated putrescine was also increased, but this increase was limited to the early stage of salt stress. Accumulation of putrescine was in accordance with induction of ADC activity and expression of the apple ADC gene (MdADC). When callus that had been treated with 200 mM NaCl was transferred to fresh medium with (successive stress) or without (recovery) NaCl, TBARS and EL were significantly reduced in the recovery treatment, indicating promotion of formation of new callus cells, compared with the successive stress treatment. Meanwhile, MdADC expression and ADC activity were also decreased in the callus undergoing recovery, whereas those of the callus under successive stress were increased. Ornithine decarboxylase (ODC) activity showed a pattern opposite to that of ADC in these conditions. D-Arginine treatment led to more serious growth impairment than no treatment under salt stress. In addition, accumulation of putrescine, induction of MdADC, and activation of ADC in D-arginine-treated callus were not comparable with those of the untreated callus. Exogenous addition of putrescine could alleviate salt stress in terms of fresh weight increase and EL. All of these findings indicated that the ADC pathway was tightly involved in the salt stress response. Accumulation of putrescine under salt stress, the possible physiological role of putrescine in alleviating stress damage, and involvement of MdADC and ADC in response to salt stress are discussed.

(203) Responses of `Golden Delicious' Apples to 1-MCP Applied in Air and Water

The efficacy of the ethylene action inhibitor 1-methylcyclopropene (1-MCP) applied in water to slow ripening of `Golden Delicious' [Malus sylvestris var. domestica (Borkh.) Mansf.] apples was evaluated in comparison with 1-MCP applied as a gas in air. The material was applied by dipping fruit in 1-MCP water solutions (0, 0.03, 0.3 or 3 μM) for 4 min, or by exposing fruit to 1-MCP gas (0, 0.01, 0.1 or 1 μL·L-1) in air for 12 h. Fruit were held in air at 20 °C for 25 days after treatment or stored at 0.5 °C in air for up to 6 months followed by 7 days in air at 20 °C. Application of 1-MCP in water or air delayed the increase in respiration and ethylene production associated with fruit ripening, and reduced the amount of fruit softening, loss of acidity and change in peel color. Treatments applied in water required a concentration 700-fold higher compared to those applied in air to induce similar physiological responses. Fruit responses to 1-MCP varied with treatment concentration, and the maximum effects were obtained at concentrations of 0.1 or 1 μL·L-1 in air and 3 μM in water. Peel color change was impacted less than retention of firmness and titratable acidity for some 1-MCP treatments. Treatment with 1-MCP was less effective for slowing peel degreening when treated fruit were stored at 0.5 °C compared to storage at 20 °C. In 1 of the 3 years of this study, fruit treated with 1-MCP and stored in air at 0.5 °C developed a peel disorder typified by a gray-brown discoloration that is unlike other disorders previously reported for this cultivar.

AROMA VOLATILE BIOSYNTHESIS IN APPLES AT HARVEST OR AFTER HARVEST AFFECTED BY JASMONATES

Effects of jasmonates on production of aroma volatile compounds by 'Tsugaru' apples [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] at harvest, and by 'Delicious' apples after harvest were examined. Among volatiles classified as alcohols, esters, ketones, aldehydes, acetic acid, and α-farnesene, esters were the most prevalent compounds, followed by alcohols. Jasmonate treatment at pre-climacteric stage increased the production of esters (butyl propanoate, butyl butyrate, and propyl butyrate) and of anthocyanin. In addition, jasmonate treatment stimulated 1-aminocyclopropane-1-carboxylate oxidase 1 (ACO 1) mRNA transcript and thylene production at 3 days after treatment. This result suggests that jasmonates may influence ACC oxidase activity. The impact of jasmonate application after harvest on volatile production differed with cultivar. The combination of ethephon with jasmonates reduced volatile production by 'Delicious' compared with ethephon only. The effect of jasmonates on volatile production was related to the effect of jasmonates on internal ethylene concentration. The results show that the effect of jasmonates on aroma volatiles in apples may be mediated by ethylene. Furthermore, the effect of jasmonates on aroma volatiles may depend on the fruit development stage when treated with jasmonates.

Seasonal changes of total carbohydrate contents in three varieties of apple (Malus sylvestris Miller) stem cuttings

Abstract Stems of apple ( Malus sylvestris Miller) varieties ‘Golden delicious’, ‘Starkrimson delicious’ and ‘Misket delicious’ were taken at different times of the year and investigated to find out seasonal changes of their total carbohydrate concentrations. All investigated varieties exhibited seasonal differences in carbohydrate contents. Total carbohydrate contents of these varieties reached at the highest value in February but the lowest in May. For all the three varieties, soluble sugars and starch contents in stem tissues were found to be similar and also soluble sugars contents of stems to be higher than starch through all the sampling period (August–May).

Chloroplast diversity in the genus Malus: new insights into the relationship between the European wild apple (Malus sylvestris (L.) Mill.) and the domesticated apple (Malus domestica Borkh.)

To unravel the relationship between the European wild apple, Malus sylvestris (L.) Mill., and its domesticated relative M. domestica Borkh., we studied chloroplast DNA variation in 634 wild and 422 domesticated accessions originating from different regions. Hybridization between M. sylvestris and M. domestica was checked using 10 nuclear microsatellites and a Bayesian assignment approach. This allowed us to identify hybrids and feral plants escaped from cultivation. Sixty-eight genotypes belonging to 12 other wild Malus species, including 20 M. sieversii (Ledeb.) Roem. accessions were also included in the analysis of chloroplast diversity. Marker techniques were developed to type a formerly described duplication and a newly detected transversion in the matK gene. Chloroplast DNA variation was further investigated using PCR-RFLP (Polymerase Chain Reaction-Random Fragment Length Polymorphism), and haplotypes were constructed based on all mutational combinations. A closer relationship than presently accepted between M. sylvestris and M. domestica was established at the cytoplasmic level, with the detection of eight chloroplast haplotypes shared by both species. Hybridization between M. sylvestris and M. domestica was also apparent at the local level with sharing of rare haplotypes among local cultivars and sympatric wild trees. Indications of the use of wild Malus genotypes in the (local) cultivation process of M. domestica and cytoplasmic introgression of chloroplast haplotypes into M. sylvestris from the domesticated apple were found. Only one of the M. sieversii trees studied displayed one of the three main chloroplast haplotypes shared by M. sylvestris and M. domestica. This is surprising as M. sieversii has formerly been described as the main maternal progenitor of the domesticated apple. This study hereby reopens the exciting discussion on the origin of M. domestica.

Salicylic Acid, Ethephon, and Methyl Jasmonate Enhance Ester Regeneration in 1-MCP-Treated Apple Fruit after Long-Term Cold Storage

Volatile esters, primarily synthesized in peel tissues, are major aromatic components of apple fruits [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.]. The use of cold storage combined with 1-methylcyclopropene (1-MCP) treatment prolongs the life of apples but represses the regeneration of esters during poststorage ripening. In this study, the regeneration of total esters was significantly increased in apple fruits treated with salicylic acid (SA) and Ethephon (ETH) that had been treated once or twice with 1-MCP. However, methyl jasmonate (MeJA) treatment resulted in regeneration of total esters after a single 1-MCP treatment. To determine the mechanism by which SA, ETH, and MeJA regulate ester regeneration, the apple alcohol acyltransferase gene (MdAAT2) was investigated at the mRNA, protein, and enzyme activity levels. Genes associated with ethylene perception were also investigated by RT-PCR. The results suggest that MdAAT2 controls ester regeneration and that MdETR1 plays a key role in ethylene perception and regulation of downstream MdAAT2 gene expression during poststorage. Ester compounds and concentrations differed in peels treated with different signal molecules, indicating that regulation of the pathway upstream of straight-chain ester biosynthesis depended on the regulation of lipoxygenase (LOX) and alcohol dehydrogenase (ADH) activity by SA, ETH, and MeJA during poststorage ripening.

Penicillium digitatum . [Distribution map

Abstract A new distribution map is provided for Penicillium digitatum (Pers.) Sacc. Ascomycota: Eurotiales. Hosts: Citrus spp. and crab apple ( Malus sylvestris ). Information is given on the geographical distribution in Europe (Cyprus, France, Greece, Italy, Netherlands, Spain, Sweden, UK), Asia (Bangladesh, Bhutan, Brunei Darussalam, China (Fujian, Guangdong, Guangxi, Guizhou, Hong Kong, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Zhejiang), India (Assam, Gujarat, Haryana, Jammu and Kashmir, Karnataka, Maharashtra, Punjab, West Bengal), Indonesia, Iraq, Israel, Japan, Jordan, Korea Democratic People's Republic, Malaysia (Peninsular Malaysia), Pakistan, Saudi Arabia, Singapore, Taiwan, Turkey, Vietnam, Yemen), Africa (Egypt, Kenya, Libya, Malawi, Morocco, Nigeria, South Africa, Tanzania, Tunisia, Zimbabwe), North America (Mexico, USA (California, Connecticut, Florida, Hawaii, Indiana, Oklahoma, Washington)), Central America and Caribbean (Barbados, Cuba, Dominica, El Salvador, Panama, Puerto Rico, United States Virgin Islands), South America (Argentina, Brazil, Chile, Venezuela), and Oceania (American Samoa, Australia (New South Wales, Queensland, South Australia, Western Australia), Cook Islands, Fiji, New Caledonia, New Zealand, Niue, Papua New Guinea).