Seed maturity indicators in Myrica esculenta, Buch-Ham.Ex. D.Don.: a multipurpose tree species of subtropical-temperate Himalayan region

Shruti Shah • Ashish Tewari • Bhawna Tewari • R. P. Singh

Received: 16 June 2008 / Accepted: 29 October 2009 / Published online: 18 November 2009� Springer Science+Business Media B.V. 2009

Abstract Myrica esculenta, Buch-Ham. Ex. D.Don is a common under-canopy associate

of chir pine and banj oak forests in the Kumaun and Garhwal Himalayas. The species

commonly known as Kaiphal is a small moderate sized evergreen dioecious tree. The

species is well known for its edible fruit and other by products and has emerged as a

potential income generating species in the Kumaun and Garhwal region. The regeneration

of Myrica esculenta is poor in natural habitats mainly due to intense anthropogenic

pressure on it. To synchronize artificial regeneration of such species exact knowledge of

maturity time is essential to avoid the collection of immature and non-viable seeds. Fruits

of Myrica esculenta were collected from nine sites varying from lower, mid and high range

of species distribution (900–2,100 masl) for assessing seed maturation indicators. Across

all the sites the colour change from green to dark red/blackish red appears as one of the

indicator of maturity. In addition to the colour change seed maturity and maximum ger-

mination was found in seeds when the weight of 100 fruits was between 41 and 60 g, fruit

moisture content between 62 and 69% and seed moisture content between 30 and 32%.

Keywords Seed maturation � Seed moisture content � Mean germination �Germination capacity

Introduction

The Himalayan ranges (27.38� N latitude and 72.98� E longitude) support a variety of

forests, which vary in species composition from east to west and from low to high ele-

vations. These forests have traditionally played a key role in safeguarding the environment

and have provided long term ecological security to the subcontinent. The central Himalaya

is divisible into three major regimes, the sal (Shorea robusta) towards the lower elevation

(300–1,000 m) with warmer climate, and the oaks (Quercus species) towards the high

elevation (1,800–2,600 m) with the chir pine zone (Pinus roxburghii) in between. Chir

S. Shah (&) � A. Tewari � B. Tewari � R. P. SinghDepartment of Forestry, Kumaun University, Nainital, Uttarakhand 263002, Indiae-mail: shrutisah@indiatimes.com

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New Forests (2010) 40:9–18DOI 10.1007/s11056-009-9179-y

pine, however, climbs up-to 2,100 m and mixes with the banj oak (Quercus leucotricho-phora) forests and the banj oak commonly descends down to 1,600 m to mix with chir pine

because of aspect related climatic variations.

In the Kumaun and Garhwal Himalaya Myrica esculenta, Buch-Ham. Ex. D.Don is a

common under-canopy associate of chir pine and banj oak forests (Bhatt et al. 2000a, b).

The species usually prefers moist sites and often forms distinctive clusters of its own

within the communities. The species commonly known as Kaiphal is a small moderate

sized evergreen dioecious tree. The species is widely distributed in sub-tropical-temperate

region of the Himalaya from Ravi eastwards towards Assam and in Khasi, Jaintia, Naga

and Lushai hills between altitudes of 900 and 2,100 masl (Anonymous 1962; Hooker

1876). The species is well known for its edible fruit and other by products and has emerged

as a potential income generating species in the Kumaun and Garhwal regions. (Dhyani and

Dhar 1994; Bhatt et al. 2000a, b). The popularity of the species can be judged from the fact

that the local people of the Kumaun region can earn over Rs. 14.00 lakh/season from

selling the fruits of the species (Bhatt and Dhar 2004). The species is also known for its

unique medicinal and industrial uses (Gupta et al. 1989). The ecological importance of

genus Myrica is of being non-leguminous angiosperm, nodulated by Frankia species. This

species is even used as fodder in some parts of Uttaranchal and Himanchal Pradesh in

times of scarcity (Singh 1982).

The regeneration of most of the wild edible species is poor in their natural habitats

mainly due to intense biotic pressure on them (Sundriyal and Sundriyal 2001). In wild

edible plants availability of mature seeds is a problem as all accessible locations are

searched by the village dwellers for collection and selling them in the local markets. To

synchronize artificial regeneration of such species an exact knowledge of maturity time is

essential to avoid the collection of immature and non-viable seeds (Willan 1985) which can

cause nursery and plantation failure. Collection of forest fruits and seeds are greatly helped

by reliable guidelines of maturity that allow the earliest possible collection (Bonner 1988).

The general conclusion is that the more mature a seed the greater is its vigor and potential

to establish as a seedling. In many species seed maturation has been related to physical

attributes (Negi and Todaria 1995 and Pandit et al. 2002), but there is very scanty infor-

mation on the seed maturation of M. esculenta. In the present study M. esculenta fruits

were collected from nine different sites for 2 years to develop seed maturation indices for

better regeneration and multiplication of the species in the nursery for future plantations.

Materials and methods

Description of study site

The study sites, situated in the Kumaun Central Himalaya lie between 29�80–29�380 N

latitude and 79�210–79�450 E longitude. A total of 9 sites were selected; of these 3 sites

were located in the low range of the species (1,500–1,600 m), 3 in the mid range (1,700–

1,800 m) and 3 in the high range (2,000–2,100 m; Table 1). All 9 sites were on the south

and south eastern aspect.

The climate of the study area is subtropical monsoon type with higher temperature

towards the lower elevation and lower temperatures towards the high elevation. Rainfall is

governed by southwest monsoon and the average annual rainfall ranges from 2,000 to

2,200 mm.

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Tree characteristics

The trees selected had a clear bole, good crown, sufficient number of fruits and were

disease free. Five average sized, healthy trees were selected and marked at a distance of

about 100 m from each other at each site. The height and diameter at breast height were

measured for each marked tree in all the sites. The mean tree height ranged between

17.8 ± 0.90 and 19.7 ± 2.08 m. The mean tree diameter at breast height ranged between

79.6 ± 5.26 cm and 94.2 ± 6.47 cm (Table 2).

Seed collection and maturity parameters

Seeds collection of M. esculenta started from the last week of April up to the availability of

fruits towards the end of May from all the sites in two respective years, 2003 and 2004. For

maturity indices fruit collection was made at an interval of 1 week till the fruits were

available. The fruits were directly collected from the tree and de-pulped in the laboratory.

The fruit parameters taken were fruit colour, fruit surface area (mm2; length 9 width)

and weight of 100 fruits taking three replicates of 25 fruits (g). All these parameters were

taken for seeds also. Weight parameters were recorded with digital electronic balance and

the size was recorded with a hand held electronic digital vernier caliper (measuring range

between 1 and 150 mm).

The moisture content of fruits and seeds was expressed on fresh weight basis and

calculated for each collection date using three replicates of 25 fruits and seeds each, dried

at 103 ± 2�C for 16 ± 1 h (International Seed Testing Association (ISTA) 1993) and then

reweighed.

Table 1 Site characteristics of Myrica esculenta

S. no. Sites Altitude Aspect Forest type

Low altitude sites (1,500–1,600 m)

1 Patwadanger I (S1) 1,520 Southern Pine forest

2 Patwadanger II (S2) 1,580 South eastern Pine forest

3 Patwadanger III (S3) 1,520 Southern Pine forest

Mid altitude sites (1,700–1,800 m)

4 Bhowali I (S4) 1,740 South eastern Oak forest

5 Bhowali II (S5) 1,760 South eastern Oak forest

6 Bhowali III (S6) 1,720 Southern Oak forest

High altitude sites (2,000–2,200 m)

7 Maheshkhan I (S7) 2,150 Southern Oak-pine forest

8 Maheshkhan II (S8) 2,100 South eastern Oak-pine forest

9 Maheshkhan III (S9) 2,120 Southern Oak-pine forest

Table 2 Tree characteristics for different sites (±SE)

Sites Mean tree height (m) Mean tree diameter (cm)

Low altitudinal sites 18.6 ± 0.70 90.6 ± 4.72

Mid altitudinal sites 19.7 ± 2.08 94.2 ± 6.47

High altitudinal sites 17.8 ± 0.90 79.6 ± 5.26

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Moisture content was calculated as

MC% ¼ Fresh weight� dry weight

Fresh weight� 100

The seeds were surface sterilized with 0.1% HgCl2. For germination four replicates of

100 seeds each were used. Germination was carried out at 25 ± 1�C on top of the paper in

a seed germinator for each collection date. Germination was counted when the visible

radicle was about 1 mm. Water was added as required during the experiment.

Germination percent was calculated as the total number of germinated seeds out of 100

tested seeds within the test period. At the end of the germination test un-germinated seeds

were classified as firm seed and rotten seeds by floating test. The result was expressed as

germination capacity (GC), calculated following Paul (1972):

GC% ¼ Total germinated seedsþ total un-germinated sound seed

Total seeds tested� 100

The data was statistically analyzed for multiple analysis of variance (ANOVA; Snedecor

and Cochran 1967) to show the significant difference between sites, years and dates.

Results

Fruit colour

The colour of the fruits was green at first collection during the last week of April, which

gradually changed to dark red/blackish red at final collection, in the last week of May in

both the years (Table 3).

Fruit characteristics

The fruit surface area in Yr2 was marginally larger than Yr1 both at the time of initial

collection and final collection across all the three altitudinal sites. Across all the sites the

mean difference in size between initial collection and final collection was 26.6 mm2 in Yr1

and 19.75 mm2 in Yr2. During both initial and final collection fruit surface area was

smaller at low altitudinal sites and larger at high altitudinal sites.

Like fruit surface area, weight of 100 fruits was also higher in Yr2 both at initial and

final collection. Across all the sites the mean difference in weight of 100 fruit between

initial collection and final collection was 29.18 g in Yr1 and 27.75 g in Yr2 (Table 3).

The moisture content of fruit was higher across all collection dates in Yr1 in comparison

to Yr2. Across all sites and with each collection date the moisture content gradually

declined. The decline in moisture was 7.99% in Yr1 and 6.48% in Yr2 between first and

final collection (Fig. 1).

ANOVA showed that fruit surface area, weight of 100 fruits and the moisture content of

fruit varied significantly across sites, years and dates of collection (P \ 0.01). The inter-

action between year 9 date was significant for weight of 100 fruits (P \ 0.01). The

interaction between site 9 year 9 date was significant for weight of 100 fruits (P \ 0.01),

moisture content of fruit (P \ 0.05; Table 4).

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Table 3 Variations in physical parameters of fruit and seeds of M. esculenta over the collection periodfrom April end to May end in Yr-1 and Yr-2

Collection Fruit characteristics Seed characteristics

Fruit Colour Fruit surfacearea (mm2)

Weight of 100fruits (g)

Seed surfacearea (mm2)

Weight of 100seeds (g)

Low altitude sites (Yr 1)

27 April Green 60.94 ± 6.54 29.32 ± 1.62 39.55 ± 2.08 7.65 ± 0.40

4 May Reddish green 65.43 ± 3.47 35.69 ± 1.09 45.02 ± 2.10 9.67 ± 0.70

11 May Red 76.34 ± 5.28 40.51 ± 2.20 48.22 ± 2.60 10.73 ± 0.35

18 May Dark red 79.55 ± 2.38 48.63 ± 1.86 50.95 ± 2.05 12.05 ± 0.40

25 May Dark red 87.61 ± 5.24 58.36 ± 2.17 55.44 ± 3.13 16.54 ± 0.44

Mid altitude sites (Yr 1)

28 April Green 63.3 ± 4.11 29.58 ± 1.41 39.64 ± 3.28 7.36 ± 0.16

5 May Green 72.88 ± 2.04 33.76 ± 1.43 50.12 ± 2.95 10.12 ± 0.41

12 May Reddish green 77.06 ± 4.27 39.93 ± 1.53 51.27 ± 2.91 11.54 ± 0.38

19 May Red 79.65 ± 6.38 50.69 ± 2.73 52.6 ± 2.12 12.2 ± 0.52

26 May Dark red 88.06 ± 4.54 56.88 ± 3.52 56.49 ± 3.61 14.01 ± 0.24

High altitude sites (Yr 1)

28 April Green 64.4 ± 3.88 28.02 ± 1.39 37.7 ± 6.15 8.12 ± 0.18

5 May Green 70.57 ± 5.25 31.73 ± 1.07 43.05 ± 2.33 10.25 ± 0.17

12 May Reddish green 76.86 ± 2.17 38.75 ± 0.84 50.46 ± 1.95 10.91 ± 0.29

19 May Red 80.39 ± 6.51 51.46 ± 1.70 53.63 ± 2.49 11.59 ± 0.32

26 May Dark red 89.92 ± 4.45 59.24 ± 1.55 57.43 ± 2.96 12.99 ± 2.33

Low altitude sites (Yr 2)

27 April Green 80.51 ± 2.91 35.59 ± 1.20 53.41 ± 3.17 10.75 ± 0.20

4 May Reddish green 88.28 ± 2.02 43.80 ± 1.75 52.9 ± 2 14.11 ± 0.33

11 May Red 89.71 ± 4.39 52.03 ± 2.35 56.91 ± 1.25 14.44 ± 0.19

18 May Dark red 91.68 ± 2.63 58.03 ± 3.27 57.98 ± 0.93 15.24 ± 0.50

25 May Dark red 94.47 ± 4.89 62.82 ± 2.56 61.20 ± 2.62 16.98 ± 0.41

Mid altitude sites (Yr 2)

28 April Green 75.16 ± 3.34 34.40 ± 1.36 49.50 ± 1.56 12.34 ± 0.10

5 May Green 83.83 ± 3.20 42.71 ± 1.36 52.20 ± 2.23 12.83 ± 0.11

12 May Reddish green 85.89 ± 0.82 45.25 ± 1.54 54.35 ± 1.65 13.44 ± 0.27

19 May Red 88.56 ± 3.46 53.53 ± 2.67 58.02 ± 4.20 13.93 ± 0.30

26 May Dark red 93.37 ± 5.55 60.93 ± 3.29 61.79 ± 2.34 17.00 ± 0.76

High altitude sites (Yr 2)

28 April Green 74.21 ± 2.77 35.28 ± 1.12 47.88 ± 1.04 11.52 ± 0.31

5 May Green 81.16 ± 2.89 42.53 ± 1.56 51.2 ± 1.58 12.51 ± 0.69

12 May Reddish green 87.91 ± 2.29 51.39 ± 1.64 55.05 ± 1.44 13.21 ± 0.52

19 May Red 89.06 ± 5.94 59.89 ± 1.21 59.97 ± 3.06 14.79 ± 0.38

26 May Dark red 101.31 ± 5 64.77 ± 0.77 60.71 ± 2.67 16.4 ± 0.37

Source of variation Fruit surfacearea (mm2)

Weight of 100fruits (g)

Seed surfacearea (mm2)

Weight of 100seeds (g)

Site F test ** ** ** **

CD at 5% 3.99 1.82 2.60 0.83

** Significant at 1% (P \ 0.01)

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Seed characteristics

Seed surface area was also marginally larger in Yr2 than Yr1 across all the sites. During

the first collection the mean difference in seed surface area in Yr1 and Yr2 was 11.3 mm2.

Across all the sites the mean difference in seed surface area between initial collection and

final collection was 17.49 mm2 in Yr1 and 10.97 mm2 in Yr2. The difference in weight of

100 seeds, across all the sites between initial and final collection was 6.80 g in Yr1 and

5.25 g in Yr2 (Table 3). In both years the decline in moisture content between initial and

final collection was significant (P \ 0.01). The decline in moisture content was 12.29% in

Yr1 and 14.85% in Yr2 between initial and final collection (Fig. 2).

ANOVA showed that seed surface area, weight of 100 seeds and the moisture content of

seed varied significantly across sites, years and dates of collection (P \ 0.01). The inter-

action between year 9 date was significant for seed surface area (P \ 0.05) and weight of

100 seeds (P \ 0.01). The interaction between site 9 year 9 date was significant for

weight of 100 seeds (P \ 0.01; Table 4).

Germination

In Yr1 seeds collected initially in the last week of April from low altitudinal sites and first

and second collection seeds of mid and high altitudinal sites failed to germinate. Germi-

nation increased thereafter with each collection and was maximum during the third col-

lection in low altitudinal sites (60 ± 4.86%) and in the mid and high altitude sites mean

germination was maximum during the fifth collection and was 54.44 ± 3.33 and

62.22 ± 3.33%, respectively (Fig. 3).

In Yr2 at low altitudinal sites germination started from first collection in the last week of

April whereas in the mid and high altitudinal sites it started from the second collection. The

germination increased with each collection and was maximum during the second collection

in low altitudinal sites (63.33 ± 7.08%). In the mid and high altitudinal sites mean ger-

mination was maximum during the fourth collection and was 64.42 ± 4.04% and

59.9 ± 9.55, respectively (Fig. 3).

ANOVA showed that the germination varied significantly across sites, years and dates

of collection (P \ 0.01). The interaction between year 9 date was significant for germi-

nation (P \ 0.01). The interaction between site 9 year 9 date was also significant for

germination (P \ 0.01; Table 4).

50

60

70

80

Low Altitude sites Mid altitude sites High altitude sites

Dates of collection

Yr1

Yr2

Fig. 1 Relation between dates of collections and moisture content of fruits in 2 years at low, mid and highaltitude sites

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Table 4 Analysis of variance (ANOVA) for different fruit and seed parameters across different collectiondates, sites and years

Character Source of variation DF Mean square F-value

Fruit surface area Site 8 541.33 8.88**

Year 1 8,781.53 144.19**

Date 4 3,920.63 64.38**

Site 9 year 8 103.42 1.69 NS

Site 9 date 32 93.00 1.52*

Year 9 date 4 105.81 1.73 NS

Site 9 year 9 date 32 43.08 0.71 NS

Weight of 100 fruits Site 8 635.99 50.16**

Year 1 3,688.83 290.93**

Date 4 7,065.86 557.28**

Site 9 year 8 61.80 4.87**

Site 9 date 32 41.53 3.27**

Year 9 date 4 62.43 4.92**

Site 9 year 9 date 32 47.63 3.75**

Moisture content (Fruits) Site 8 41.06 6.97**

Year 1 1,681.70 285.83**

Date 4 417.14 70.90**

Site 9 year 8 24.66 4.19**

Site 9 date 32 13.48 2.29**

Year 9 date 4 4.78 0.81 NS

Site 9 year 9 date 32 11.20 1.90*

Seed surface area Site 8 297.09 11.50**

Year 1 2,720.82 105.38**

Date 4 1,565.96 60.65**

Site 9 year 8 91.11 3.52**

Site 9 date 32 48.82 1.89*

Year 9 date 4 100.24 3.88*

Site 9 year 9 date 32 29.23 1.13 NS

Weight of 100 seeds Site 8 38.77 67.83**

Year 1 575.09 1,005.99**

Date 4 266.18 465.62**

Site 9 year 8 6.79 11.87**

Site 9 date 32 2.35 4.12**

Year 9 date 4 4.74 8.30**

Site 9 year 9 date 32 4.78 8.37**

Moisture content (seeds) Site 8 257.57 28.49**

Year 1 1,639.34 81.37**

Date 4 1,437.83 159.08**

Site 9 year 8 43.68 4.83**

Site 9 date 32 13.22 1.46 NS

Year 9 date 4 20.46 2.26 NS

Site 9 year 9 date 32 12.62 1.39 NS

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Table 4 continued

Character Source of variation DF Mean square F-value

Germination Site 8 481.03 4.99**

Year 1 3,134.81 32.51**

Date 4 17,054.11 176.91**

Site 9 year 8 306.84 3.18**

Site 9 date 32 960.31 9.96**

Year 9 date 4 5,067.81 52.57**

Site 9 year 9 date 32 587.09 6.09**

NS non significant

** Significant at 1% (P \ 0.01)

* Significant at 5% (P \ 0.05)

0

10

20

30

40

50

Low Altitude sites Mid altitude sites High altitudesites

Dates of collection

Yr1Yr2

Fig. 2 Relation between dates of collections and moisture content of seeds in 2 years at low, mid and highaltitude sites

0

10

20

30

40

50

60

70

Low Altitude sites Mid altitude sites High altitude sites

Dates of collection

Yr1Yr2

Fig. 3 Relation between dates of collection and germination in two different years at low, mid and highaltitude sites

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Across all the sites germination percent increased with the decline of moisture content

of fruit and seed. The maximum germination was observed when the moisture content of

fruit was between 62.01 ± 2.34% (high altitudinal sites) and 69.61 ± 0.56% (low altitu-

dinal sites) and range of moisture content of seed was 30.38 ± 0.70% (low altitudinal

sites) to 32.22 ± 0.66% (low altitudinal sites) across both the years.

Discussion

Seed collectors have long been aware that most mature and immature fruits and seeds can

be distinguished in a number of ways e.g. by colour difference, increased firmness or

brittleness, decreased moisture content and specific gravity or by change in physical

dimensions (Edwards 1980). Willan (1985) has considered colour change in fruit and cone

to be reliable criteria for judging seed maturity. In M. esculenta colour change from green

to dark red was good maturity indicator. Colour change has also been recommended as a

ripeness indicator in the Himalayan wild cherry (Prunus cerasoides D.Don, Rox.; Tewari

2005) and in Bauhinia retusa Ham. (Upadhayay et al. 2006). In Ailanthus excelsa Roxb.

the physiological maturity was indicated by a direct colour change of the fruit from

yellowish brown to brown (Ramakrishnan et al.1990). Negi and Todaria (1995) also find

colour to be the best indicator of maturity for five species of the Garhwal Himalayas.

Moisture content has also been used as a reliable maturity indicator by numerous

researchers. Decline in moisture content percent from maturing seeds is closely related to

seed maturity (Adams and Rinni 1981). In M. esculenta decline in moisture content was a

good indicator of maturity. The seeds matured when the moisture content of seeds was

between 30.46 and 31.72%. According to Maideen et al. (1990) in Casuarina equisetifoliaseed maturity is attained when the cone moisture content is below 50%. This is also

observed in different pine species. In pine maturity is generally reached when moisture

content of cone is below 50% (Edwards 1980). In Pyracantha crenulata the moisture

content of fruit between 30 and 36% and the seed moisture content between 68 and 71%

have been adjudged indicators of seed maturation (Shah et al. 2006). Pandit et al. (2002)

also observed that in Populus ciliata, a broad leafed species the drop in moisture content of

capsule from 80 to 60% during maturation coincide with the maximum germination in seed.

In low altitudinal sites seeds matured in the first and second week of May, whereas in mid

and high altitude sites they matured in the third and fourth weeks of May. Maturity for

M. esculenta was earlier at low altitudinal sites in comparison to mid and high altitude sites.

Earlier maturation in lower sites was possibly due to higher temperature than the high sites.

Pandit (2002) also observed similar results in Pinus roxburghii and Cupressus torulosa.

Edwards (1980) reported a significant correlation between maturity and physical

parameters but there are also many unsuccessful attempts. From this study it becomes

evident that colour change from green to dark red, fruit moisture content ranging between

62 and 69% and seed moisture content varying from 30 to 32% are best indicators of

maturity in M. esculenta.

References

Adams CA, Rinnie RW (1981) Seed maturation in Soyabean (Glycine max (L.)Merr.) is independent of seedmass and of the parent plant. Yet is necessary for production of viable seed. J Exp Bot 32:615–620

Anonymous (1962) A dictionary of Indian raw material and industrial products, vol VI L-M. CSIR Pub-lication, New Delhi, pp 471–472

New Forests (2010) 40:9–18 17

123

Bhatt ID, Dhar U (2004) Factors controlling micropropagation of Myrica esculenta buch.-Ham. Ex D.Don: ahigh value wild edible of Kumaun Himalaya. Afr J Biotechnol 3(10):534–540

Bhatt ID, Rawal RS, Dhar U (2000a) The availability, fruit yield and harvest of Myrica esculenta Buch-Hamex D.Don in Kumaun (West Himalaya). Indian Mt Res Dev 20(2):146–153

Bhatt ID, Rawal RS, Dhar U (2000b) Improvement in seed germination of Myrica esculenta Buch-Ham exD.Don—a high value tree species of Kumaun Himalaya. India Seed Sci Technol 28:597–605

Bonner FT (1988) Using leachate conductivity of bulked samples to estimate seed quality. In: Proceedingsof the 1988 southern forest nursery association, 1988, July 25, 218-Charleston, SC. Southern ForestNursery Association, Columbia Sc, pp 164–172

Dhyani PP, Dhar U (1994) Myrica esculenta box myrtle (Kaiphal). Himavikas Occasional Publ.3.GBPIHED, Almora

Edwards DGW (1980) Maturity and quality of tree seeds- a state of the art review. Seed Sci Technol 8:625–657Gupta PN, Chandra R, Govind S (1989) Bio-chemical studies on some Myrica genotype. Prog Hort 21(1–2):

168–170Hooker JD (1876) Flora of British India, vol V. Bishan Singh Mahendra Pal Singh, DehradunInternational Seed Testing Association (ISTA) (1993) International rules for seed testing (Supplement).

Zurich, SwitzerlandMaideen SK, Selvaraj JA, Vinayarai RS (1990) Cone attributes as indices of seed maturity and effect of

cone and seed grades on seed germination and vigour in Casuarina equisetifolia. J.R. & G. For SeedSci Technol 18:483–489

Negi AK, Todaria NP (1995) Effect of seed maturity and development on germination of five species fromGarhwal Himalaya, India. J Trop For Sci 8(2):255–258

Pandit A (2002) Development of technology for assessment of seed viability during storage of Pinusroxburghii and Cupressus torulosa in Kumaun Himalaya. Ph.D. Thesis, Kumaun University, Nainital

Pandit A, Pant K, Ram J (2002) Effect of collection date on capsule moisture content and germination ofPopulus ciliata wall. Ex. Royle from central Himalaya. New For 23:121–130

Paul DK (1972) A hand book of nursery practice for Pinus caribaea var. hondurensis and other conifers inwest Malaysia Working paper no. 19, FO:SF/MAL 12, UNDP/FAO, Kualalumpur

Ramakrishnan HB, Jacqueline AS, Vinaya Rai RS (1990) Studies on ripeness index and pre-sowing seedtreatment in Ailanthus excelsa Roxb. Seed Sci Tecnol 18:491–498

Shah S, Tewari B, Bisht S, Tewari A (2006) Seed maturation indicators in Pyracantha crenulata Rox. inKumaun Central Himalaya. New For 32:1–7

Singh RB (1982) Fodder trees of India. Oxford & IBH Publication, New Delhi, p 560Snedecor GW, Cochran WG (1967) Statistical methods. Oxford and IBH, New Delhi, p 593Sundriyal M, Sundriyal RC (2001) Seed germination and response of stem cuttings to hormonal treatment in

six priority wild edible fruit species of Sikkim Himalaya. Ind For 127(6):695–705Tewari B (2005) Regeneration and nursery techniques of Prunus cerasoides. Ph.D. thesis, Department of

Forestry, Kumaun University, NainitalUpadhayay L, Singh RP, Tewari A, Bisht S, Shah S (2006) Seed maturation indicators in Bauhinia retusa

Ham. in Kumaun Central Himalayas. Indian J For 29(4):367–371Willan RL (1985) A guide to forest seed handling with special reference to the tropics. FAO forestry paper

20/2 FAO, Rome

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