Trop. Agr. Develop. 64（3）： 125 － 134，2020

Studies on Growth Characteristics and Starch Productivity of the Sago Palm (Metroxylon sagu Rottb.) Folk Varieties in Seram and Ambon Islands, Maluku,

Yoshinori YAMAMOTO 1, *, Tetsushi YOSHIDA 1, Isamu YANAGIDATE 1, Febby Jeanry POLONAYA 2, Willem Anthon SIAHAYA 2, Foh Shoon JONG 3, Yulius Barra PASOLON 4, Akira MIYAZAKI 1, Tomoko HAMANISHI 5, and Kazuko HIRAO 6

1 Faculty of Agriculture, Kochi University, Nankoku, Kochi 783-8502, Japan 2 Faculty of Agriculture, Pattimura University, Ambon, Maluku 97233, Indonesia 3 PT. National Timber and Forest Products, Selatpanjang, Riau 28753, Indonesia 4 Faculty of Agriculture, Haluoleo University, , Southeast 93232, Indonesia 5 Faculty of Home Economics, Kyoritsu Women’s University, Chiyoda-ku, Tokyo 101-8437, Japan 6 Aikoku Gakuen Junior College, Edogawa-ku, Tokyo 133-8585, Japan

Abstract The growth characteristics and starch productivities of four major sago palm folk varieties (spiny types: Ihur, Tuni, Makanaru; non-spiny type: Molat) were compared on Seram and Ambon Islands, Maluku, Indonesia. These folk varieties were recognized there by spiny or non-spiny; length and density of the spine; the angle of leaf crown; the attaching features of leaflets to the rachis (horizontal or V-shaped); drooping of the leaflet tip; starch productivity; starch color, etc. There were no significant differences in growth characteristics, leaf and leaflet characteristics, starch content, and characteristics related to starch content among the four major folk varieties on Seram Island. The trunk growth characteristics and starch content of the Ihur and Tuni on Ambon Island were comparable to those of Seram Island. In addition, the starch productivities of the four major folk varieties, 500–600 kg plant-1, were superior to those of sago palms in Malaysia and in western areas of Indonesia. The starch contents were as high as those of the late-flowering folk varieties of the Indonesian territory on New Guinea. Key words: Folk variety, Growth characteristics, Leaf characteristics, Macronutrients, , Starch productivity

ha, respectively. Rasyad and Wasito (1986) reported Introduction the sago palm forest area on the Maluku Islands was Sago palm is estimated to have originated in New 30,000 ha and that on Seram Island was considered to Guinea (Barrau, 1959; Flach and Schuiling, 1989; occupy 11,000 ha. Regarding the folk varieties of sago Takamura, 1990; Flach, 1997), and in particular, in palms grown in the Maluku Islands, Sastrapradja and Irian Jaya, Indonesia. The morphological and growth Mogea (1977) and Flach (1980) reported five sago palm characteristics and the starch productivity there were folk varieties (Sagu Mulat, Sagu Makanaro, Sagu Ihor, highly diverse (Widjono et al., 2000; Matanubun and Sagu Tuni, Sagu Ikau) on Ambon Island (Although Maturbongs, 2006; Matanubun, 2015; Yamamoto et al., the spelling of name for the same folk variety differs 2020). It was estimated that sago palms were transmit- depending on the reporters, such as “Mulat” and ted from the origin to western areas such as Indonesia, “Molat”, “Ihor” and “Ihur”, “Makanaro” and “Makanaru”, Malaysia, and Thailand and to northern area such as the the authors followed the description by Louhenapessy Philippines (Ehara, 2015). (1992) and the latter spelling of each variety was used On the other hand, Beccari (1918) set the center of in this report). Yamamoto (1999) confirmed these diversity for sago palm on the Maluku Islands, and the five folk varieties on Ambon Island and also identified sago starch has been utilized as a staple food there for four of the five on Seram Island, with the exception of long time (Sastrapradja and Mogea, 1977). Flach (1977) Ikau. Among these folk varieties, Tuni, Molat, Ihur and estimated the wild and semi-/cultivated sago palm Makanaru were presumed to be major folk varieties on areas of the Maluku Islands to be 50,000 and 10,000 Ambon and Seram Islands (Sastrapradja and Mogea, 1977; Yamamoto, 1999). Molat is a non-spiny type,

Communicated by H. Ehara while the other varieties are spiny types. In addition, Received Jan. 10, 2020 Louhenapessy (1992) recognized nine sago palm folk Accepted Apr. 16, 2020 varieties (Tuni, Molat, Ihur, Makanaru, Duri Rotan, * Corresponding author [email protected] Molat Berduri, Molat Merah, Molat Berduri Merah, Sagu 126 Trop. Agr. Develop. 64（3）2020

Swanggi) in the Maluku Islands. In addition to the four after varieties) in Seram and Ambon Islands, Maluku, major folk varieties, among the fi ve folk varieties listed Indonesia and to compare them with those of the sago by him, three folk varieties are considered to be variants palms we have already investigated in Malaysia and of Molat: Molat Berduri (it has spines when young, but Indonesia. In addition, we interviewed the sago growers they disappear with growth), Molat Merah (the leaf axis on Seram and Ambon Islands about the characteristics is red when young), and Molat Berduri Merah (with the of the four major varieties. combined characteristics of Molat Berduri and Molat Materials and Methods Merah). Of the other two folk varieties, Duri Rotan is a spiny type with short spines, high density, and low Sampling investigation starch productivity, and Sagu Swanggi (Swanggi means We selected Kairatu Village on Seram Island in “ghost”) accumulates starch immediately after trunk 2007 and Rutong Village on Ambon Island in 2008 as formation. Moreover, Schuiling (2009) reported six sago research sites based on the interview surveys of Prof. J. palm folk varieties as well as their characteristics, Ihur, E. Louhenapessy and Prof. E. T. H. Maris of Pattimura Tuni, Molat Licin, Molat Duri Putih, Molat Duri Merah, University in Ambon (Fig. 1). Makanaru Duri Hitam, and Makanaru Duri Putih, on Seram Island based on the interview with the most Kairatu Village on Seram Island in 2007 experienced sago processor. In his report, Molat Licin The research was conducted in September 2007. and Makanaru Duri Hitam or Makanaru Duri Putih Three plants each of four major sago palm varieties, Ihur, may correspond to Molat and Makanaru in the other Tuni, and Makanaru (spiny types), and Molat (non-spiny reports (Sastrapradja and Mogea, 1977; Flach, 1980; type) at harvest stage (fl ower bud formation to fl owering Louhenapessy, 1992; Yamamoto, 1999). stage), which are considered to be suitable for harvest- As described above, several reports on the folk ing, were sampled in a small-scale sago palm garden in varieties and their characteristics of the sago palms on Kairatu Village. However, two plants were sampled for Maluku Islands are available, but some characteristics Molat due to diffi culty in fi nding out the plant at harvest for each variety were not necessarily consistent among stage. In addition, one sampled plant of Tuni was in the the reports. Moreover, the growth characteristics and young fruit stage, and it was slightly over the appropri- starch productivities of these sago palm folk varieties ate harvest time, so the plant was excluded from the grown in the Maluku Islands have not been adequately calculation of mean values of all characteristics. The soil clarifi ed. types of the surveyed gardens were loam to silt loam. Clarifi cation of the differences in the growth char- The leaf and growth characteristics were examined for acteristics and starch productivities of sago palm folk sampled sago palms. varieties is important for the selection of the best variety Regarding the leaf characteristics, three leaves in line with the production target, and it is a problem were samples from the basal, middle and top positions that must be solved before promoting the development of a plant, and the leaf length (the length from the base of sago palm plantations that have been developing in of the leaf sheath to the tip of the uppermost leafl et) and Indonesia and Malaysia in recent years. number of leafl ets were measured. The longest leafl ets Based on this background, the purpose of this from the left and right sides of the sampled leaves were study is to clarify the growth characteristics and starch collected, and the length, maximum width, and SPAD productivities of four major sago palm varieties (here- value (SPAD-502, Minolta Co.) were measured. The

Seram 1. Kairatu Masohi

Liang Ambon 1. Rutong 100 km Ambon

Fig. 1. Map of Ambon and Seram Islands, Maluku, Indonesia Yamamoto et al.: Starch productivity of sago palm in Seram and Ambon Islands 127

SPAD values were measured at the central parts of the Methods of analyzing inorganic and organic com- longest leaflets. The leaf area was determined accord- ponents ing to the method of Omori (2001). The leaf area was Analysis of macronutrients calculated as follows: the length of the longest leaflet × The dried materials of the pith samples in Kairatu its maximum width × the number of leaflets per leaf × Village on Seram Island were ground through 100 mesh 0.54. The mean leaflet area was calculated by dividing or less and 0.5 g of the ground material was decomposed the leaf area by the number of leaflets per leaf, and the with sulfuric acid-hydrogen peroxide solution; nitrogen leaf area per plant was calculated by the leaf area × the (N) and phosphorous (P) were analyzed using the semi- number of leaves per plant. In this research, since the micro Kjeldahl method and Murphy and Riley’s method individual plants were from flower bud formation to the (1962), respectively. Potassium (K), calcium (Ca) flowering stage, the leaf length and leaflet characteris- and magnesium (Mg) were analyzed using an atomic tics were shorter and smaller from base to top (Table 3) absorption spectrophotometer (AA-6800, Autosampler, (Yamamoto et al., 2014), the values of the basal leaves ASC-6100, Shimadzu Co.). The content of each nutrient were used for the leaf and leaflet characteristics. is shown on a dry weight basis. The mean content of pith After the trunk was cut at the nodal position of the per individual plant is shown by the mean value of the lowest surviving leaf, the length from the base to the pith samples collected at five positions from the trunk nodal position was measured as the trunk length. The base to the top. trunk was divided into four equal parts, and at each cut position including the base and the top of the trunk (five Analysis of total sugar and starch positions in total), the diameter was measured. Then, Total sugar and starch were analyzed according to the trunk was cut to a fixed length (log), and the log the method of Murayama et al. (1955) for the dried pith weights were measured with a 100 kg-bar scale. Disks materials. That is, the total sugar was extracted from 0.2 g 2–3 cm thick were taken from the positions of the trunk dried pith materials, which had been pulverized through at which the diameters were measured and around 50 g 100 mesh or less three times with 80% hot ethanol pith samples were radially taken from the center of the (80°C), and the starch was extracted from the ethanol- disks and measured accurately to the nearest 0.1g using extraction residues six times with 4.6 N perchloric acid. a portable electronic balance (HL-200 type, Kyouei Co.). Total sugar and starch were quantified by the anthrone The sampled piths were dried for about two days method as the glucose amount, and the total sugar was in a simple dryer (75–77°C) heated with incandescent expressed as the glucose amount; starch was expressed lamps and brought back to Japan. In Japan, the samples by multiplying 0.9 by the glucose amount. Moreover, were dried again for two days in a ventilated dryer at the mean percentage of the pith per individual plant is 65°C and then the weights were measured. The dried shown by the mean value of the pith samples collected at pith samples were used to analyze macronutrients (N, P, five positions from the trunk base to the top. The starch K, Mg, Ca) and total sugar and starch. The pith weight content (yield) per plant was calculated by the starch of the trunk was calculated as 80% of the trunk (Yatsugi, percentage and the pith dry weight. 1977). Statistical analysis Rutong Village on Ambon Island in 2008 Statistical analysis was performed on the obtained In November 2008, four plants of Ihur and Tuni at data by JMP (Version 7.0) (SAS Institute). harvest stage were sampled in Rutong Village on Am- Results bon Island in the same manner as the method above as being used in Kairatu Village on Seram Island. However, Interview survey no leaf characteristics were measured in this research The results of interview surveys on the except numbers of leaves and leaf scars. characteristics of the four varieties with the sago growers on Seram Islands are shown in Table 1. Growth Interview survey period from sucker planting/emergence to trunk We interviewed the sago growers in the sampled formation and from the trunk formation to harvest sago gardens in Kairatu and Rutong Villages about the (flower bud formation to flowering) were 4–5 and 10–15 characteristics of the major sago palm varieties growing years, respectively, in all four major varieties. Among there. the four varieties, only Molat was non-spiny type. Both 128 Trop. Agr. Develop. 64（3）2020

Table 1. Morphological and starch characteristics of the four major sago palm varieties grown on Seram Island, Maluku, Indonesia. Spine Angle of Leaflet Tip part Starch Variety Spine length density leaf crown shape of leaflet productivity color Ihur + 21) 21) wide horizontal droop 11) W, R 2) Tuni + 3 3 wide V-shape non-droop 2 W Makanaru +3) 1 1 narrow V-shape non-droop 3 W, R Molat - - - wide horizontal non-droop 4 W 1) Longer or higher (1) - shorter or lower (3). 2) W: whitish, R: reddish. 3) The tip of spine is black and poisonous. the spine length and density were higher in the order weight, and volume of the four varieties were 21.0 m, of Makanaru>Ihur>Tuni in the spiny type varieties. 11.5 m, 50.1 cm, 1946 kg and 2.3 m3, respectively (Table The angle of leaf crown was narrower in Makanaru 2). Makanaru had the longest plant and trunk lengths than in the other varieties. The differences in leaflet but the shortest trunk diameter. The trunk diameter was characteristics were observed in the leaflet shape and largest in Tuni, and its trunk volume was the largest. tip part of it. The starch productivity was higher in the Ihur had the highest trunk weight. However, no signifi- order of Ihur>Tuni>Makanaru>Molat. The starch color cant differences were found in any of the characteristics was very white in Molat, followed by Tuni, but two types among the varieties. The trunk weight and volume of starch colors, white and reddish, were recognized in were closely related to the diameter (r = 0.720, p<0.05) Ihur and Makanaru. compared to the length (r = 0.317). The two varieties (Ihur and Tuni) found and sampled in Rutong Village on Ambon Island, the charac- Varietal differences of leaf characteristics at harvest stage teristics were mostly same as those of Ihur and Tuni in The mean values of the numbers of leaves, leaf Kairatu Village on Seram Island. scars, their total, and the leaf length of all varieties were 24.1, 100.6, 124.7 and 9.4 m, respectively (Table 4). No Sampling investigation (Seram Island, 2007) significant differences were found among the varieties Varietal differences in growth characteristics at harvest in any characteristic; however, Ihur had the highest val- stage ues for all characteristics. The number of leaves and leaf The mean plant length, trunk length, diameter, lengths were lowest in Tuni. In addition, the mean val-

Table 2. Plant length and trunk characteristics of sago palm varieties at harvest stage grown in Kairatu Village on Seram Island, Maluku, Indonesia.

Plant Trunk No. of Variety Spine length Length Diameter Weight. Volume plants (m) (m) (cm) (kg) (m3) Ihur 3 + 21.8 a 11.9 a 50.6 a 2134 a 2.4 a Tuni 2 + 20.5 a 11.5 a 54.7 a 2012 a 2.6 a Makanaru 3 + 22.2 a 12.7 a 46.1 a 1848 a 2.1 a Molat 2 - 19.5 a 10.1 a 49.1 a 1789 a 1.9 a Mean 21.0 11.5 50.1 1946 2.3 SD 1.1 1.0 3.1 136 0.3 CV (%) 5.1 8.3 6.2 7.0 12.4 SD: Standard deviation. CV: Coefficient of variation. Values in a column followed by the same letter are not significantly different at p< 0.05 by Tukey’s test.

Table 3. Positional differences of leaf and leaflet characteristics of sago palms at harvest stage grown in Kairatu Village on Seram Island, Maluku, Indonesia. Leaf position Characteristie* Basal Middle Top Leaf length (m) 9.4 ± 1.1 8.1 ± 1.6 4.2 ± 1.7 No. leaflets (/leaf) 174 ± 6.6 158 ± 17.1 107 ± 25.3 Max. leaflet length (m) 1.65 ± 0.10 1.61 ± 0.12 1.17 ± 0.17 Max. leaflet width (cm) 11.9 ± 1.0 11.2 ± 1.3 7.3 ± 1.3 Max. leaflet SPAD value 67.0 ± 4.3 69.8 ± 2.7 65.1 ± 6.2 *Mean ± standard deviation of 10 plants of 4 varieties. Yamamoto et al.: Starch productivity of sago palm in Seram and Ambon Islands 129 ues of the number of leaflets per leaf, length, maximum the leaf areas per leaf and plant showed higher positive width, and SPAD value of the longest leaflet of all variet- correlations with leaflet area (r = 0.965, p<0.001) and leaf ies were 172.8, 1.65 m, 11.9 cm, and 66.6, respectively. area per leaf (r = 0.894, p<0.001) than with the number of However, no significant differences were found among leaflets (r = 0.415) and the number of leaves (r = 0.775, the varieties in these characteristics (Table 5). The mean p<0.05), respectively. values of the leaflet area and leaf areas per leaf and per plant of all varieties were 1064 cm2, 18.4 m2 and 445.4 m2, Varietal differences in the dry matter, total sugar, and respectively. None of the areas were significantly differ- starch percentages and starch and macronutrient contents ent among the varieties; however, Ihur had the highest in the pith at harvest stage and Makanaru had the lowest (Table 6). The leaflet area The mean dry matter and total sugar and starch showed a higher positive correlation with leaflet width percentages in the pith of all varieties were 45.6, 5.1, (r = 0.930, p<0.001) than length (r = 0.832, p<0.01), and and 74.7%, respectively; no significant differences were

Table 4. Leaf characteristics of sago palm varieties at harvest stage grown in Kairatu Village on Seram Island, Maluku, Indonesia. Number of No. of leaves No. of leaf Leaf length* Variety (a + b) plants (a) scars (b) (m) Ihur 3 26.0 a 106 a 132 a 10.5 a Tuni 2 22.5 a 99 a 121 a 8.4 a Makanaru 3 24.3 a 95 a 120 a 8.9 a Molat 2 23.5 a 103 a 126 a 9.7 a Mean 24.1 101 125 9.4 SD 1.5 4.7 5.6 0.9 CV (%) 6.2 4.6 4.5 10.0 SD: Standard deviation. CV: Coefficient of variation. *Basal leaf length. Values in a column followed by the same letter are not significantly different at p< 0.05 by Tukey’s test.

Table 5. Leaflet characteristics of sago palm varieties at harvest stage grown in Kairatu Village on Seram Island, Maluku, Indonesia.

No. of Longest leaflet* No. of Variety leaflets* length max. width SPAD plants (No. leaf-1) (m) (cm) value Ihur 3 178 a 1.70 a 12.8 a 69.2 a Tuni 2 168 a 1.67 a 11.7 a 63.3 a Makanaru 3 177 a 1.55 a 10.9 a 67.6 a Molat 2 169 a 1.70 a 12.2 a 66.5 a Mean 173 1.65 11.9 66.6 SD 5.3 0.07 0.8 2.5 CV (%) 3.1 4.2 6.7 3.7 SD: Standard deviation. CV: Coefficient of variation. *Measured on the basal leaf. Values in a column followed by the same letter are not significantly different at p< 0.05 by Tukey’s test.

Table 6. Leaf let and leaf area of sago palm varieties at harvest stage grown in Kairatu Village on Seram Island, Maluku, Indonesia. No. of Leaflet area* Leaf area* Leaf area Variety plants (cm2) (m2 leaf-1) (m2 plant-1) Ihur 3 1172 a 20.9 a 543 a Tuni 2 1052 a 17.7 a 401 a Makanaru 3 913 a 16.1 a 396 a Molat 2 1120 a 18.9 a 442 a Mean 1064 18.4 445 SD 112 2.0 68 CV (%) 10.5 11.0 15.2 SD: Standard deviation. CV: Coefficient of variation. *Measured on the basal leaf. Values in a column followed by the same letter are not significantly different at p< 0.05 by Tukey’s test. 130 Trop. Agr. Develop. 64（3）2020

Table 7. Starch contents and its related characteristics of the pith of sago palm varieties at harvest stage grown in Kairatu Village on Seram Island, Maluku, Indonesia. Pith dry Starch No. of Dry matter Total sugar Starch Variety weight content plants (%) (%) (%) (kg plant-1) (kg plant-1) Ihur 3 47.6 a 817 a 4.9 a 71.6 a 585 a Tuni 2 45.3 a 731 a 3.8 a 75.8 a 552 a Makanaru 3 43.8 a 649 a 5.8 a 76.1 a 494 a Molat 2 45.6 a 653 a 5.9 a 75.2 a 491 a Mean 45.6 712 5.1 74.7 530 SD 1.6 80 1.0 2.1 46 CV (%) 3.4 11.2 19.4 2.8 8.6 SD: Standard deviation. CV: Coefficient of variation. Values in a column followed by the same letter are not signifi- cantly different at p< 0.05 by Tukey’s test. found among the varieties in any value (Table 7). The 5 varietal differences of the total sugar percentage in the pith were slightly larger than those of the dry matter * 4 and starch percentages. The variation patterns of these 3 percentages in each variety according to the position position

along the trunk from base to top were almost the same; r unk T 2 the mean variation patterns of the varieties by the trunk position are shown in Fig. 2. Although there were small 1 differences in each percentage depending on the posi- 0 10 20 30 40 50 60 70 80 tion of the trunk, the dry matter and starch percentages Dry matter, total sugar and starch (%) were somewhat lower at the bottom and top of the trunk. Fig. 2. Positional differences of dry matter, total sugar and The mean pith dry matter weight of all the varieties at starch percentages in pith of sago palms at harvest stage grown in Kairatu Village on Seram Island, harvest stage was 712 kg, and no significant differences Maluku, Indonesia. were found among the varieties; however, it was highest Mean values of 10 palms of 4 varieties. for Ihur and lowest for Makanaru and Molat (Table 7). *Basal (1) to top (5) position of trunk. ●Dry matter, ○Total sugar, △Starch The mean starch content (yield) of all varieties was 530 kg, and no significant differences were found among the varieties. The maximum was 585 kg for Ihur, followed by the other varieties. Looking at the difference of contents 552 kg for Tuni; as compared with these two varieties, of these macronutrients according to the positions along the starch content of Makanaru and Molat was lower the long axis of trunk, the positional pattern of each ― around 490kg (Table 7). Focusing on the individual nutrient content was almost same among the varieties, palms, the highest starch yield, 747 kg, was recorded in so the mean values of N, P, Ca, and Mg of all varieties are the individual of Ihur and that of Makanaru also showed shown in Fig. 3. They tended to be higher from base to a high yield of 620 kg. The starch content showed a high top, particularly, the contents of the top N and Ca were significant positive correlation (r = 0.957, p<0.001) with higher than those at other positions, while the K content the pith dry weight, but not with the starch percentage tended to decrease from the base to the top. (r = 0.339). The mean contents of macronutrients in pith of all Sampling investigation (Ambon Island, 2007) the varieties were N: 1.96; P: 0.65; K: 5.01; Ca: 2.16; and The growth characteristics, the starch content, and Mg: 0.94 g kg-1. There were no significant differences the related characteristics of Ihur and Tuni in Rutong in the N and Ca contents among the varieties (Table 8). Village on Ambon Island were compared with those However, the P content was significantly higher in Ihur of the values of the same varieties on Seram Island than in the other varieties, and the K content was sig- (Kairatu Village) described above. Ihur and Tuni on nificantly lower in Molat than in Ihur and Tuni. Also, the Ambon Island had significantly longer plant lengths Mg content was significantly higher in Makanaru than and more leaf scars and significantly fewer leaves than in Ihur. The K content was highest for all varieties, fol- the same varieties on Seram Island (Table 9). As a lowed by N or Ca, and lowest in Mg for Ihur and in P for result, the numbers of leaves + leaf scars were almost Yamamoto et al.: Starch productivity of sago palm in Seram and Ambon Islands 131

Table 8. Macronutrient contents in the pith of sago palm varieties at harvest stage grown in Kairatu Village on Seram Island, Maluku, Indonesia. No. of Macronutrient (g kg-1) Variety plants N P K Ca Mg Ihur 3 2.28 a 1.17 a 5.63 ab 2.01 a 0.77 b Tuni 2 1.92 a 0.53 bc 5.62 ab 2.07 a 0.87 ab Makanaru 3 1.97 a 0.58 bc 4.86 bc 1.96 a 1.12 a Molat 2 1.68 a 0.33 c 3.95 c 2.59 a 1.01 ab Mean 1.96 0.65 5.01 2.16 0.94 SD 0.25 0.36 0.80 0.29 0.16 CV (%) 12.5 55.5 15.9 13.5 16.5 SD: Standard deviation. CV: Coefficient of variation. Values in a column followed by the same letter are not signifi- cantly different at p< 0.05 by Tukey’s test.

5 the spine among the varieties were different from the results of Sastrapradja and Mogea (1977), Flach (1980),

* 4 Yamamoto (1999), and Schuiling (2009). It was not clear what caused the differences in the spine characteristics 3 position among the varieties observed between the two results, but the results may fluctuate by the palm ages when r unk T 2 the spine characteristics were evaluated (Sastrapradja, 1986). Moreover, the differences in the other morpho- 1 0123456 logical leaf characteristics such as leaf shape and droop- Macro-nutrient (g kg-1) ing of the leaf tip, and in the starch color were coincided Fig. 3. Positional differences of macronutrient contents in with the results of Flach (1980) and Yamamoto (1999). pith of sago palms at harvest stage grown in Kairatu However, the starch productivity among the varieties Village on Seram Island, Maluku, Indonesia. were not entirely coincided with the results of the Mean values of 10 palms of 4 varieties. previous reports of Flach (1980), Louhenapessy (1992), *Basal (1) to top (5) position of trunk. ●N, ○P, △K, ◇Ca, □Mg Yamamoto (1999), and Schuiling (2009). Considering this point, it needs a further detailed study on the differ- the same, and the difference in years until harvest was ences in the characteristics among the varieties. estimated to be small. There was no significant varietal In the following, we discussed about the results of difference in trunk growth characteristics between the sampling survey. The growth characteristics of the the two islands. On the other hand, the dry matter major varieties (Tables 2 and 10) were in the same range and starch percentages in the pith of Ihur and Tuni on as high-yielding and late-flowering varieties around Lake Ambon Island were significantly lower than those on Sentani near Jayapura, Papua (Yamamoto et al., 2020), Seram Island; conversely, the total sugar content was and showed higher values compared with those of sago significantly higher (Table 10). In particular, Ihur’s dry palms in Sarawak (Yamamoto et al., 2003a) and the matter percentage in the pith on Ambon Island was low varieties Rotan and Tuni in , while and the total sugar content was high. However, there showing value levels comparable to those of Molat in was no significant difference in the starch contents of Southeast Sulawesi (Yamamoto et al., 2010). According Ihur and Tuni between the two islands. to an interview with sago growers at Kairatu Village, the main sago palm varieties there, Tuni, Molat, Ihur, and Discussion Makanaru, take four to five years from sucker planting According to the interview results with the sago to trunk formation and approximately 10–15 years from growers on Seram and Ambon Islands, the four major trunk formation to flowering, and the varietal differences varieties on Seram and Ambon Islands were recognized in these period are small. These periods are similar with there by spiny or non-spiny; length and density of the those of the late-flowering varieties around Lake Sentani spine; angle of leaf crown; leaflet shape (the attaching near Jayapura, Papua (Yamamoto et al., 2020). features of leaflets to the rachis: horizontal or V-shaped); The leaf and leaflet characteristics of the major drooping of the leaflet tip; starch productivity; starch varieties (Tables 4–6) were superior to those of the sago color, etc. The differences in length and density of palms in Sarawak (Flach and Schuiling, 1991; Nakamura 132 Trop. Agr. Develop. 64（3）2020

Table 9. Plant length and trunk characteristics of sago palm varieties at harvest stage grown on Seram (Kairatu Village) and on Ambon (Rutong Village) Islands, Maluku, Indonesia. Plant Number of Trunk Researched No. of Variety length leaves leaf scars Length Diameter Weight. Volume site plants (a + b) (m) (a) (b) (m) (cm) (kg) (m3) Seram 3 21.8 26.0 106 132 11.9 50.6 2134 2.4 Ihur Ambon 4 26.2 18.0 114 132 14.9 52.9 2672 3.3 a Seram 2 20.5 22.5 99 121 11.5 54.7 2012 2.6 Tuni Ambon 4 23.1 21.3 111 132 12.3 47.2 1822 2.2 Ihur 7 24.0 a 22.0 a 110 a 132 a 13.4 a 51.8 a 2403 a 2.9 a Variety Tuni 6 21.8 a 21.9 a 105 a 127 a 11.9 a 50.9 a 1917 a 2.4 a Seram 5 21.1 B 22.0 A 107 B 129 A 12.6 A 51.4 A 2160 A 2.6 A Research site Ambon 8 24.7 A 19.7 B 113 A 132 A 13.6 A 50.1 A 2247 A 2.8 A ANOVA Research site * * * ns ns ns ns ns Variety ns ns ns ns ns ns ns ns Research site × Variety ns ns ns ns ns ns ns ns * indicate significant difference at p<0.05. Values with the same letter within the same column are not significantly different at p<0.05 by Tukey’s test.

Table 10. Starch contents and its related characteristics of sago palm varieties at harvest stage grown on Seram (Kairatu Village) and on Ambon (Rutong Village) Islands, Maluku, Indonesia. Pith dry Starch No. of Dry matter Total sugar Starch Variety Researched site weight content plants (%) (%) (%) (kg plant-1) (kg plant-1) Seram 3 47.6 817 4.9 71.6 585 Ihur Ambon 4 34.5 734 11.2 66.2 489 Seram 2 45.3 731 3.8 75.8 552 Tuni Ambon 4 41.8 601 6.9 69.7 419 Ihur 7 41.4 a 776 a 8.0 a 68.9 a 537 a Variety Tuni 6 43.6 a 666 a 5.3 b 72.7 a 486 a Seram 5 46.5 A 721 A 6.7 B 70.8 A 511 A Research site Ambon 8 38.2 B 668 A 9.1 A 68.0 B 454 A ANOVA Variety ns ns * ns ns Research site * ns * * ns Variety × research site ns ns ns ns ns * indicate significant difference at p<0.05. Values with the same letter within the same column are not significantly different at p<0.05 by Tukey’s test.

et al., 2004; 2009), South Sulawesi (Osozawa, 1990), There was no significant variation among the major and a variety of Rotan in Southeast Sulawesi, Indonesia varieties in the pith dry weight or the starch percentage (Yamamoto et al., 2014). In addition, the values of these and, consequently, in the starch content (yield) characteristics were comparable to the values of a (Table 7). The pith dry matter, total sugar, and starch variety of Tuni on Seram Island as reported by Flach percentage values of the major varieties were almost and Schuiling (1991) and a variety of Molat of Southeast the same as those of sago palms at harvest as reported Sulawesi as reported by Yamamoto et al. (2014). In previously (Yamamoto et al., 2003a; Yamamoto et al., particular, as the leaf area per plant was considered 2010). Schuiling (2009) reported that the starch density to be closely related to the starch productivities of the of pith (kg m-3) beyond the early vegetative phase were sago palm varieties (Yamamoto et al., 2014; 2016), it was almost same among the varieties, Molat, Ihur, Tuni, and presumed that the starch yields of sago palm varieties Makanaru on Seram Island. The positional differences in growing on Seram Island were higher than those of sago the pith dry matter, total sugar, and starch percentages palms growing in Sarawak, and that of a variety of Rotan along the trunk base to the top were small in all varieties in Southeast Sulawesi, and almost the same as that of (Fig. 2) (Yamamoto et al., 2003a). However, the starch a variety of Molat in Southeast Sulawesi, which had content was more affected by the pith dry weight than 400–500m2 leaf area per plant. by the starch percentage (Yamamoto et al., 2010). As Yamamoto et al.: Starch productivity of sago palm in Seram and Ambon Islands 133 a result, the starch content tended to be higher in the necessarily identical to these results (Table 8). However, order of Ihur>Tuni>Makanaru≒Molat, although there it was agreed that the K content was extremely high as was no significant varietal difference, it was close to the compared with other nutrients, and Yamamoto et al. interview results mentioned above and of Yamamoto (2003b) estimated that the K in the pith of sago palm (1999). The Ihur and Tuni on Abmon Island surveyed plays an important role in starch synthesis. Significant in this research might not have reached the appropriate varietal differences in P, K, and Mg contents were harvest stage (Jong, 1995; Yamamoto et al., 2003a), as observed in this research and the N, P, and K contents they showed lower dry matter and starch percentages of the major varieties were higher than those in the sago and higher total sugar percentages in the pith than did palms grown in Sarawak (Yamamoto et al., 2003b). In those of the same varieties on Seram Island (Tables addition, Yamamoto et al. (2003b) showed that the N, 7 and 10). Therefore, if they were harvested at a time P, Ca, and Mg contents were higher from the base to when the dry matter percentages in the pith were further the top of the trunk, but the positional difference of the increased, it was considered that both the pith dry weight K content was small. The results of this research were and the starch content increased, and the starch content consistent for all macronutrients except K (Fig. 3). also improved accordingly. This may confirm that there Possible causes for these disagreements between the were no significant differences in the starch contents of present research results and the results of Yamamoto et Ihur and Tuni grown on Seram and Ambon Islands. al. (2003b) may include differences in the soil and water In addition, the starch yields of the four major vari- environment in which sago palms grow and differences eties were higher than those of sago palms in Sarawak in the sago palm varieties investigated. A detailed study (Yamamoto et al., 2003a; 2003c) and in the western areas is needed on the relationships between the mineral of Indonesia (Yamamoto, 2015), as well as varieties of nutrient contents in the pith and the growth and starch Tuni and Rotan in Southeast Sulawesi. Moreover, they productivity of sago palms. were comparable to some individuals of a variety of In conclusion, there were no significant differences Molat (Yamamoto et al., 2010) in Southeast Sulawesi in growth characteristics, leaf and leaflet characteristics, and those of the late-flowering varieties around Lake starch content (yield), and the related characteristics Sentani, Papua (Yamamoto et al., 2020). Especially, as among the four major sago palm varieties on Seram mentioned above, the starch content of Molat in this Island. The trunk growth characteristics and starch con- research showed the same levels as some individuals of tent of Ihur and Tuni on Ambon Island were comparable the same name variety in Southeast Sulawesi; however, to those on Seram Island. In addition, the starch produc- that of Tuni in this research had about 250 kg higher tivities of the four major sago palm varieties, 500–600 kg content than did that of the same name variety in South- plant-1, were superior to those of sago palms in Malaysia east Sulawesi, mainly due to the difference in the pith and in the western areas of Indonesia. They showed the dry weight (Yamamoto et al., 2010). It is necessary to same high starch content as those of the late-flowering examine the causes from the viewpoints of differences varieties of the Indonesian territory on New Guinea. in the genetic background and environmental conditions Acknowledgments in both regions. In addition, the starch productivities of other varieties are also expected to differ depending on In conducting this survey, Professor Maris Elder environmental conditions such as soil, water, and culti- Theodoor Hetharia, Faculty of Agriculture, Pattimura vation management methods. Therefore, it is necessary University, Maluku, Indonesia, provided various to investigate a greater number of plants for each variety conveniences for the survey. He and Professor Emeritus, in wider areas of investigation. Dr. Julius Elseos Louhenapessy, the former Vice There have been very few research reports on the President of the University, gave us valuable information contents of mineral nutrients in the pith of sago palm. about sago palms in the Maluku Islands, Indonesia. We Flach and Schuiling (1991), Matsumoto et al. (1998), express our gratitude. We thank Mr. Charles Niak, a and Yamamoto et al. (2003b) reported that the contents sago farmer, for his kind support and contributions to of macronutrients in the pith of sago palm from flower our research. This survey was supported by a Tropical bud formation to the young fruit stage in Sarawak Biological Resources Research Grant from the Japan were higher in the order of K>Ca>N>Mg>P; the mean Society for the Promotion of Science (JSPS) and by a values of all varieties in this research were consistent JSPS KAKENHI Grant (Grant No. 19405023). We would with those results, but the individual varieties were not like to express our gratitude. 134 Trop. Agr. Develop. 64（3）2020

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