Interaction of animal with its environment 1 Structure of earthworm community in the suburbs of Fukuoka City Distribution of environments under study Maximum biomass of earthworm in various vegetation Eco-morphological feature of representative Pheretima Unsettled problem a. Growth analysis on the population having a bi-modal distribution of weight, b. the individual without the male pore, b. The function of intestinal coeca c. Taxonomy problem of Pheretima of Fukuoka outskirts and Minamata 2. Population ecology of representative species Pheretima sp.(H-1) population 1.The earthworm is born, lives, and dies a Number of individuals and weight distribution b. Population metabolism c. Daily fecal pellet production rate, daily food ingestion rate digestive efficiency and assimilation efficiency 2.Relations between individuals of the same species(Movement and dispersion) a mass emergence of Pheretima sp. (H-1) on fine day after rain b The structure of habitat and the distribution pattern 3.Interaction of animal with its environment a c Resource utilization and energy balance b Interaction of animal with its environment The bio-economic life table 1 Coordinates in study area north latitude East longitude north latitude Kumano1968 East longitude Experiment field of university1971-74 Hexagon Quadrangle 33.620157 130.452783 33.627934 130.423611 33.620193 130.453679 33.628488 130.423541 33.619719 130.453716 33.628542 130.423616 33.619635 130.452616 33.628466 130.423772 33.628403 130.424062 33.627863 130.423842 Hakozaki 1968 Quadrangle 33.614841 130.424094 33.614529 130.424459 33.614363 130.42441 33.614515 130.423895 It consists of Grass area and Dicotylendonous area. Refer to slide x. Center of investigation or Sampling Hokanken North 1991 - 2003 33.512697 IBPMinamata 130.499957 1969-1971 32.180103 130.59335 Hokanken South 1991 - 2003 33.512393 130.500784 2 3 Table 1-1 Distribution of environments under study Area G Area D Area H Area K Quercus glauca 100% Imperata cylindrica Solidago altissima Celtis sinensis ca 40 % Artemisia vulgaris Damnacanthus indicus Carex spp. Vicia hirsuta Achyranthes japonica Trachelosperum asiaticum Cover degree Litter layer (cm) A0 horizon (cm) pH in H2O ca 50 % 0 1.7 - 4.2 5.56 100% 0.1 - 1.5 3.0 -12.3 6.05 ca 70 % 2.0 - 4.0 5.0 -40.0 6.2 ca 20 % 3.5 - 5.0 5.0 -10.0 4.7 Primary production 540. ? 1176.5 1010 study year 1971-72 1971-73 1967-68 Tree vegetation Cover degree Ground surface vegetation ? IBP minamata 844 1967-68 1969-1971 4 Change in earthworm's scientific name Old temporary name in field note [2/1/2014] Name of earthworm until 2002 [Abbreviation] Name of earthworm after 2012/7/15 [Abbreviation] aokii Pheretima sp. (H-1) Amynthas tokioensis var. Hakozaki irregularis [Ph. sp. (H-1)] [A. tokioensis var. Hakozaki] heterochaeta Pheretima heterochata Amynthas corticis hupeiensis Pheretima hupeiensis Amynthas hupeiensis robusutus Pheretima robusutus Amynthas robusutus phaselus maculosus? Pheretima phaselus Amynthas phaselus micronaria Pheretima micronaria Amynthas micronarius calfornica Pheretima calfornica Metaphire calfornica vittata Pheretima vittata [Ph. vittata] Metaphire pseudvittatus Pschmardae Pheretima schmardae Duplodicodrilus schmardae hilgendorfi agrests noting (H-1) heterochata 5 earthworm in various vegetation g fresh wt m-2 Area H Area K IBP Minamata Area Area G Pheretima schmardae 6.648 1.877 11.045 122.964 43.945 0.679 24.912 29.24 Pheretima sp. (H-1) Pheretima vittata Pheretima irregularis Pheretima sieboldi Pheretima sp. (M-3) Pheretima heterochaeta 60.512 Pheretima micronaria Allolobophora caliginosa 7.298 Area D 10.552 2.312 17.122 34.39 Other fewer species 0.182 2.494 2.809 11.14 Allolobophora jaoinica Pheretima hupeiensis Two unknown Pheretima Bimastos parvus 5.111 3.762 0.471 Pheretima hupeiensis Two unknown Pheretima Bimastos parvus Four unknown Pheretima One unknown Lumbricidae Bimastos parvus Bimastos parvus 6 Table 2-3 Eco-morphological feature of representative Pheretima Species Life history Pheretima Pheretima Pheretima Pheretima Pheretima Pheretima Pheretima Pheretima schmardae sp. (H-1) vittata irregularis sieboldi sp. (M-3) heterochaeta micronaria Newly born hatch out Newly born hatch out Newly born hatch out in Many younger appear in in spring, mature in in spring, mature in summer, mature till next summer, mature till next summer and disappear June and some remained summer, and disappear summer, and some remain till August till November. till August. till winter. . 6 months Life time 8 months age composition Main habitat 12 months One generation Compost Two or three generation Old grass Old grass Edge of Ever green forest in field edge of forest mountain site Inhabit layer yellow dark reddish dark brown dark pigmentation green brown with reddish yellow band brown Number of pair of spermatica Activity two Deeper soil layer deep black purple light grey light brown purplish brown three four two Active Body form Older vegetation vegetation Litter - A -Soil Dry dingle Body Intestinal coeca Younger Litter Hibernate site Body size over 12 months sluggish Plumply slenderly 474 1922 6006 2206 30500 5 6-7 8 6 Most complex 1434 799 564 Simple showing a conical form pairs of finger projection Composition of gut content Organic rich Small raw Large raw Organic rich Litter from Organic rich Organic rich soil matter humus humus matter with tree leaves matter and mineral soil small raw humus 7 Total density 160 Body weight frequency 80 N 0 (b) Density of each generation 80 8 N 00 G 72/7/19 1000 125 (a) Body weight of each generation 0 A O D F A J A O D F mg Fig. 1-15. Seasonal change in (a) body weight and (b) density of each generation of Pheretima heterochaeta in area G. Vertical lines in upper figure indicate one standard deviation. 8 Table 7 - 1 Microbial density in gut materials of Pheretima sp. (H-1) coeca 6-7 finger shape litter dweller 4-5 individuals per a measurement The average of 5 measurement (1972/5/15 - 6/25) Body D Fore gut Mid gut Hind gut weight × 10 × 10 × 10 Average weight 5 2.9 6.1 9.4 2722.6 6 4.0 7.6 11.2 GWW 7 7.7 15.4 27.9 Table 7 - 2 The number of observed colonies in gut materials of Pheretima heterochaeta coeca conical form soil dweller 4 individuals per a measurement The average of 5 measurements (1971/10/21-12/8) Body D Fore gut Mid gut Hind gut length 3 13.1 12.3 21.7 10cm から 4 3.6 3.1 6.6 12cm 5 0.9 0.9 1.7 9 FIG. 3.2-5. Seasonal changes in average population density (a), mean body size (b) 10 and number of dead worms observed (c). Taxonomical problem of earthworm collected in Fukuoka pref. • Young earthworm cannot be identified. • All of Pheretima vittata and the majority of Pheretima sp.(H-1) have the male pore. • In Fukuoka and Minamata, there are a lot of nameless earthworms. . . The 18th section Rough sketch of Pheretima sp.(H-1) 11 1972 7 15 0 0 0 0 6 0 2 0 0 7 4 2 2 6 0 0 010 6 1 7 0 0 5 6 1 0 0 0 7 11 4 2 1 0 9m Fig. 2-3a. Distribution maps of earthworm in the experimental field. Black letter indicate the density of Pheretima sp. (H-1) (25 x 25 cm-2) 12 13 140 120 Pheretima sp.(H-1) 100 80 60 40 20 0 F A J A O D F A J A O D F A Fig. 1-10. Seasonal change in population density of Pheretima sp.(H-1) in area D. Vertical lines indicate one standard deviation. 14 Pheretima sp.(H-1) Body fresh weight (mg wt) 3375 D 72/2/23 Ph.H 1000 D 72/3/12 Ph.H D 72/3/28 Ph.H D 72/5/23 Ph.H D 72/6/8 Ph.H D 72/4/12 Ph.H D 72/2/14 Ph.H 125 0 Pheretima sp.(H-1) 3375 1000 125 D 72/4/27 Ph.H D 72/5/10 Ph.H D 72/6/19 Ph.H 0 013826 20 40 60 Frequency (%) D 72/6/30 Ph.H D 72/7/15 Ph.H D 72/7/29 Ph.H Fig. 1-11. Seasonal change in body weight frequency of Pheretima sp. (H-1) in area D( 1972). Open bars indicate the frequency of immature and black bars indicate the frequency of mature. 15 Density and average weight 1000 3 CO 2 mm h 120 140 100 120 -1 100 80 80 60 60 40 100 40 20 20 0 0 4000 10 Production 2000 現存量と呼吸量 0 Respiration 呼吸速度 生産と異化 -2000 Elimination 1 -4000 10 100 1000 10000 J F M A M J J A S 16 Table 3-1 Production, respiration, assimilation and ecological ratios Pheretima sp. (H-1) Species Year Area 1968 Area H 1971 Area D 1972 Area D Ph. Vittata 1972 Area G 1972 Clay sand 1968 Area H Ph.irregularis Ph. sieboldi 1968 Area K Ph. Heterochaeta 1970 1971 1971 1971 1973 1972 IBP Area D Area G Net production g dry wt m-2 11.489 242.642 4.075 7.688 1.174 6.368 3.217 0.827 1.59762 4.7243 KJ m -2 Cocoon production 85.772 162.339 24.686 134.575 68.199 17.573 5.23 33.765 99.839 g dry wt m-2 0.0018 0.0021 0.0022 KJ m -2 Respiration 0.007554 0.008648 0.009205 KJ m -2 Assimilation 138.239 193.669 589.275 44.124 263.885 126.566 43.639 6.527 234.467 659.465 KJ m -2 Average biomass 224.011 356.008 831.917 68.81 398.46 194.765 61.212 11.757 268.232 759.304 g dry wt m-2 Maximum biomass 0.779 1.729 4.368 0.268 1.72 0.883 0.226 0.09 0.695 1.632 g dry wt m-2 R/A 2.982 3.324 0.617 0.544 P/B 5.218 4.446 P/B MAX 1.363 2.313 8.378 0.708 2.63 1.371 0.76 4.352 1.964 0.725 0.157 1.368 2.889 0.641 0.662 0.65 0.713 0.555 0.874 0.869 4.381 3.702 3.643 3.659 2.649 2.3 2.895 1.545 1.463 1.638 1.141 1.584 1.168 1.635 17 Food ingestion rate and Fecal pellet production rate -1 (mg dry wt d ) 10000 1000 100 at 20℃. Body weight (mg fresh wt) 10 10 food consumption rate 100 1000 10000 fecal pellet production Y = 545.15 W 0.4382 at 25 ºC F = 1419.12 W 0.7015 at 25 C Y = 481.55 W 0.5280 at 20 ºC F = 1405.93 W 0.9594 at 20 ºC Y = 338.13 W 0.8764 at 15 ºC F = 986.86 W 0.92840 at 15 ºC Y:Food consumption per day W:Body weight F:Fecal pellet production per day 18 12 10 8 6 4 2 0 Carbon 0 (22.5 mg C g 50 -1 dry wt) 100 (222.9 mg C g 12 10 8 6 4 2 0 -1 dry wt) Nitrogen 0 (1.64 mg N g 50 -1 dry wt) 100 (13.1 mg N g -1 dry wt) 60 50 mature immature 40 30 20 10 0 0 50 100 Fig.4-6. Nutrient contents of the materials in alimental canal of the pre-mature collected in field. worms a, Carbon content; b, Nitogen content; c; The ratio of food consumption to fecal pellt production, of the 19 reared worms. In Lowest figure, the open area indicates the contribution by immature worms and the black area indicates the contributionby mature worms. 60 Carbon Assimilation efficiency (%) 40 Carbon餌の質 20 0 0 20 40 60 80 100 -20 32.7 31.3 35.4 42.6 41.8 41.6 The ratio of food consumed to 29 total material consumed (%) 49.5 -40 -60 24.6 60 -80 Nitrogen Assimilation efficiency (%) 40 20 Nitrogen 0 0 20 40 60 80 100 -20 -40 -60 Fig. 5-2a. The relation between the assimilation efficiency and the ratio of food consumed to total material consumed. 20 Table 5-1 Digestive efficiency of Phererima sp. (H-1) Efficiency (%) Food consumption Pellet production (%) Reared worms (A=C-F) Carbon Nitrogen 8.124 7.392 35.5 Reared worms (A=P+R) 25℃ 20℃ 15℃ 2.31 1.98 2.22 35.5 Carbon Nitrogen 1.895 9.559 24.77 38.86 Field Population 21 10 Pheretima vittata 1972 5 0 White bars : Movement individual Black bars: Death individual Pheretima sp. (H-1) 1972 40 20 0 35 Air temperature 30 1972 25 100 20 15 50 Precipitation 10 5 0 0 M J J Mass movement of earthworm on fine after rain 22 35 30 100 -1 25 Rain fall (mm d ) Soil surface temperature ( ℃) Water content of soil (%) 40 20 15 50 10 5 0 0 Percipitation temperature Water content of soil soil respiration -2 -1 CO2 evolution (mg CO2 m h ) 100 50 0 50 Control 25 0 5/7 5/14 5/21 5/28 6/4 6/11 6/18 23 Soil respiration rate 24 4 2 2.0 L CO 2 10 individuals / 2 replicate 1.5 L CO 2 25 individuals / 5 replicate 0 6 4 2 0 2 1.0L CO 2 15 individuals /3 replicate 0 0.667 L CO 1 2 5 individuals / 1 replicate 0 0.5 L CO 2 2 40 individuals /8 replicate 1 0 0.25 L CO 1 0 1 0.125 L CO 2 2 20 individuals /4 replicate 10 individuals /2 replicate 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Fig. 7-7a Number of earthworm showing the rashing out behavior to various concentration of Carbon dioxide gases. 25 The death of worm on bare area Expantion of distribution area of earthworm The aggregation is weakened. The worm can aboid the food shortage condition. Solar radiation Abruptly dispersion of earthworm Increase of activity of soil organism. Increase of CO2 concentration in habitat. Wormcast became to be bad habitat, for worms Fine day after rain Conformation of a aggregated distribution of worms, The resistency of worm to dry condition increase. On the other hand, the food shortage condition for worms occure. Decrease of CO 2 expiration rate of wormcast. Wormcast became to confortable condition as shelter for earthworms. dry condition No rain Number increase of filamentous fungi, Rest pause of bacteria. Increase of water holdency of worm cast Fecal pellet expiration Number increase of micro-organism in earthworm intestine. Metabolism of Earthworm in habitat. Fig7-9 Population meaning of the simultanously death of earthworm; Pheretima sp. (H-1) on fine days after rain 26 2.0 4000 1.5 同化 2000 1.0 0 異化 -2000 0.5 -4000 0.0 J F M Production 巌のM star A M J Elimination J A S Surviving rate 12 4.0 10 3.0 8 6 森下のIδ 2.0 4 1.0 2 0 0.0 J F M A M M star J Iδ J A S Figure8-2 Distribution pattern of individuals 27 -2 Density (N / 25×25 cm ) 25 30 20 25 72/6/19 73/4/28 20 15 15 10 10 5 5 0 0 0 2 1973/4/20 1973/5/10 4 A 24.18 14.89 (21.38) 1972/6/8 1973/6/15 1972/6/19 1972/6/30 1972/7/15 4.86 6.1 6.01 5.08 9.92 6 1/A 0.414 0.07 (0.037) 0.2058 0.1639 0.1664 0.1967 0.1008 0 B 0.0157 0.143 (0.221) 0.499 0.665 0.7658 1.1167 0.8306 0.5 1 1.5 r 0.889 0.834 (0.739) 0.849 0.812 0.911 0.767 0.257 The regression coefficient between number of earthworm and thickness of wormcast28 Hot dry condition after late July Re-aggregation of matured individual Aggregation of cocoon oviposited Aggregatness of earthworm Aggregation in cocoon stage Hatch out Dispersion of young worm to search a food resource Dry condition in May Aggregation of pre-matured individuals Warm-wetter condition in June Abruptly dispersion of pre-matured and matured individuals Disappearance of population Feb. March April May June July August Fig. 8-7. The seasonal change in the degree of aggregation of Pheretima sp. (H-1) in relation to own's life history. 29 Decrease of habitatness Dispersion of young worm to search a foor resource. Decrease of aggregatness. Random distribution of loose colonies with random deposition of individuals within colonies. B) Pre-matured stage in dry season Abruptly dispersion, Simultanously death on fine day after rain. Uniform distribution of small colony (copulation) within clump. Random distribution of clump. D) Post matured state Dry Aggregation of worm Conformation of the compact colony Increase of thickness of worm cast. Increase of resistency of earthworm to dry condition. food shortage in habitat End of wetter-summer season dry condition Re-aggregation of matured individual, Conformation of compact colony. Then, the aggregated distribution of cocoon Disappearance of population Fig. 8-8. The structure of habitat and the distribution pattern of earthwormPheretima ; sp. (H-1). 30 Distribution range range Litter Distribution fall fall Soil asLitter food resouce as food CaloricSoil content of resouce litter, and soil Caloric content of litter, and soil Soil surface tenperature Soil surface tenperature Population density Body weight Population density distribution Body weight Litter distribution consumption rate Fecal pellet Litterrate consumption rate production Fecal pellet production rate Assimilation efficiency in rearing condition Assimilation efficiency in activity rearing condition Feeding of food competitor Feeding activity of food competitor Maximum quantity of Caloric content in the utilizable resource Caloric content in the utilizable resource Total quantity of fecal pellet produced Total quantity of fecal pellet produced Quality of food resource Quality of food resource Assimilation efficiency in field Assimilation Utilizable efficiency energy in field Resouce utilization intensity Resouce utilization intensity Resource division among competitor Resource division among competitor wormcast presenting in fieldMaximum quantity of wormcast presenting in field Decaying rate of wormcasts Decaying rate of wormcasts Distribution pattern of individuals Distribution pattern of individuals utilizable area utilizable area Fig. 9-1 The estimating procedure of energy absorbed by earthworms Fig. 9-1 The estimating procedure of energy absorbed by earthworms 31 Copy Distribution range Litter fall Soil as food resouce Caloric content in the utilizable resource Caloric content of litter, and soil Quality of food resource Soil surface tenperature Population density Body weight distribution Litter consumption rate Fecal pellet production rate Assimilation efficiency in field Total quantity of fecal pellet produced Resouce utilization intensity Assimilation efficiency in rearing condition Feeding activity of food competitor Resource division among competitor Maximum quantity of wormcast presenting in field Decaying rate of wormcasts Distribution pattern of individuals utilizable area Fig. 9-1 The estimating procedure of energy absorbed by earthworms 32 Three process of resource utilization food requirement 62416.494 KJ m-2 (=10283.43×0.887+2963.66×17.983) (cultivated earthwom) the absorbed energy 892.531 KJ m-2 the absorbable energy The procedure for estimation are shown in side22 978.638 KJ m-2 (P+R=832.198 KJ) 33 Copy Three process of resource utilization food requirement 62416.494 KJ m-2 (=10283.43×0.887+2963.66×17.983) (cultivated earthwom) the absorbed energy 892.531 KJ m-2 the absorbable energy The procedure for estimation are shown in side22 978.638 KJ m-2 (P+R=832.198 KJ) 34 Table 11-1 Bio-economic life table ofPheretima sp. (H-1) in area D 1972 Date Jan. N B 0.0 B.W 0.0 P F.R A.H E 0.0 F.R A.V F.C A.H F.C A.V L-C R.W *_ m/m * m A 0.0 0.0 0.0 0.0 1176.5 0.2 0.0 0.1 0.0 1175.3 1.7 6.2 0.4 0.0 0.3 0.0 1170.2 1.9 11.0 0.6 0.1 0.4 0.1 1162.2 1.7 11.1 3.8 0.3 2.6 0.3 1118.7 1.5 11.0 5.9 0.5 4.1 0.5 1049.7 1.6 10.8 19.3 1.5 13.4 1.4 827.7 1.4 9.7 24.2 30.7 3.1 21.4 2.8 531.3 1.3 8.6 14.9 33.0 6.4 23.0 5.9 68.4 1.5 9.5 32.8 5.1 22.9 4.7 0.0 15.2 1.3 7.1 27.0 4.9 18.9 4.5 15.2 2.7 25.6 5.6 17.9 5.1 15.2 1.6 7.5 5.1 19.3 8.6 13.4 7.9 15.2 1.5 5.9 9.9 9.1 4.9 6.3 4.5 15.2 3.9 7.1 0.0 0.0 0.0 0.0 15.2 15.8 Feb.14 60.8 379.6 6.2 26.6 Feb.23 83.2 619.2 7.4 18.5 Mar.12 92.8 933.7 10.1 57.3 Mar.28 Apr.12 Apr.27 May 10 May 23 Jun. 8 Jun.19 108.8 100.5 99.8 93.4 88.3 76.8 68.0 1850.9 2457.3 4924.5 6821.0 7892.4 8378.2 8326.0 17.0 51.9 11.5 166.2 1.7 152.5 26.1 114.2 31.9 101.7 71.3 87.9 92.6 24.5 49.3 73.0 89.4 109.1 122.4 4.9 10.4 6.01(6.10) 87.2 Jun.30 66.6 7368.0 110.6 156.0 Jul.15 51.8 5028.5 97.1 232.5 Jul.29 26.2 1803.5 68.8 60.1 Aug.28 0.0 0.0 0.0 N:density (N m-2 ), B:Biomass (dry wt m-2 ), B.W:Body dry weight of individual, P:Production (dry wt w-2 ), E:Elimination (dry wt w-2 ) F.R A.H:Litter requirement ofAmynthas sp.(H-1), F.R A.V:Litter requirement ofAmynthas vittatus , F.C A.H: Litter consumption of Amynthas sp. (H-1), F.C.A.V: Litter consumption ofAmynthas vittatus L-C:Litter supply - Litter consumption, R.W: Resource supply from wormcast, (m/m, m, A: The degree of Aggregation of individuals, see Chapter 8) 35 Table 11-1 Bio-economic life table ofPheretima sp. (H-1) in area D 1972 Date Jan. N B 0.0 B.W 0.0 P F.R A.H E 0.0 Copy F.R A.V F.C A.H F.C A.V L-C R.W *_ m/m * m A 0.0 0.0 0.0 0.0 1176.5 0.2 0.0 0.1 0.0 1175.3 1.7 6.2 0.4 0.0 0.3 0.0 1170.2 1.9 11.0 0.6 0.1 0.4 0.1 1162.2 1.7 11.1 3.8 0.3 2.6 0.3 1118.7 1.5 11.0 5.9 0.5 4.1 0.5 1049.7 1.6 10.8 19.3 1.5 13.4 1.4 827.7 1.4 9.7 24.2 30.7 3.1 21.4 2.8 531.3 1.3 8.6 14.9 33.0 6.4 23.0 5.9 68.4 1.5 9.5 32.8 5.1 22.9 4.7 0.0 15.2 1.3 7.1 27.0 4.9 18.9 4.5 15.2 2.7 25.6 5.6 17.9 5.1 15.2 1.6 7.5 5.1 19.3 8.6 13.4 7.9 15.2 1.5 5.9 9.9 9.1 4.9 6.3 4.5 15.2 3.9 7.1 0.0 0.0 0.0 0.0 15.2 15.8 Feb.14 60.8 379.6 6.2 26.6 Feb.23 83.2 619.2 7.4 18.5 Mar.12 92.8 933.7 10.1 57.3 Mar.28 Apr.12 Apr.27 May 10 May 23 Jun. 8 Jun.19 108.8 100.5 99.8 93.4 88.3 76.8 68.0 1850.9 2457.3 4924.5 6821.0 7892.4 8378.2 8326.0 17.0 51.9 11.5 166.2 1.7 152.5 26.1 114.2 31.9 101.7 71.3 87.9 92.6 24.5 49.3 73.0 89.4 109.1 122.4 4.9 10.4 6.01(6.10) 87.2 Jun.30 66.6 7368.0 110.6 156.0 Jul.15 51.8 5028.5 97.1 232.5 Jul.29 26.2 1803.5 68.8 60.1 Aug.28 0.0 0.0 0.0 N:density (N m-2 ), B:Biomass (dry wt m-2 ), B.W:Body dry weight of individual, P:Production (dry wt w-2 ), E:Elimination (dry wt w-2 ) F.R A.H:Litter requirement ofAmynthas sp.(H-1), F.R A.V:Litter requirement ofAmynthas vittatus , F.C A.H: Litter consumption of Amynthas sp. (H-1), F.C.A.V: Litter consumption ofAmynthas vittatus L-C:Litter supply - Litter consumption, R.W: Resource supply from wormcast, (m/m, m, A: The degree of Aggregation of individuals, see Chapter 8) 36 Next food chain Atmosphere Solar radiation Eco-physiological feature Niche segregation along with time, food and habitat among equivalent species Life history Habitat preference Food preference Enlargement of distribution area Space deposition of individuals Space requirement m/m Attractive effect among individuals Negative chemotaxis of earthworm to carbon dioxide F.H P.H Abruptly deispersion of worm on fine days after rain E Elimination m Aggregated distribution in dry days Respiration Death of worm B Biomass N Number B.WBody weight Food requirement Food shortage in limitted area Escape from food shortage condition P Food consumption rate under rearing condition Assimilation ability * Production Environmental value of wormcast as food Assimilation Energy-material balance Assimilation Food efficiency consumption in field in field F.C Persistence of individual and species P.H Quality of resource Food consumption of competitor Fecal pellet expiration Litter supply L-C Food requirement of competitor F.R P.V Decaying wormcast Water holdency of worm cast Porously character of wormcast Microbial density in worm cast D.W Wormcast as habitat condition Soil surface temperature Fig. 11-1. Inter relation ship between environment and earthworm 37 Next food chain Atomosphere Life requirement Eco-physiological feature Inter-species relation ship Inner-species relation ship Space requirement Child care Cocoon production or breeding Sexual passion Food requirement Persistence of individuals and species Population Metabolism 餌に対する関係 Food resource Habitat structure Environment Habitat condition Fig. 11-3. Inter relation ship between environment and animals. Vegetation Equivalent species 38
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