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Chapter
1
Number
and biomass of Earthworms in Several Types of vegetation in the suburb of
Fukuoka City of Kyushu Japan
Chapter
2 Ecological
feature and feeding habit of Pheretima group (Oligochaeta: Megascolecidae)
41
Chapter
4
Chapter
5
Chapter
6
Chapter
11
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Number
and biomass of Earthworms in Several Types of vegetation in the suburb of
Fukuoka City of Kyushu Japan
Introduction
2
Study
Area
2
Method
3
1.
Sampling
3
1.1. Sampling of earthworm 3
1-2. Sampling of earthworm's cocoon
4
2.
The regression coefficient between weight and body volume
4
3.
Weight distribution of earthworms
5
4.
Growth analysis on the population having a bi-modal distribution of weight
5
Result
6
1.
Population densities and weight distributions of earthworms in area H
6
1-1. Pheretima schmardae
6
1-2. Pheretima sp. (H-1)
6
1-3. Pheretima vittata 7
1-4. Pheretima micronaria 7
2.
Population densities and weight distribution of earthworms in area K (Kumano
shrine) 7
2-1. Pheretima irregularis
7
2-2. Pheretima heterochaeta
8
2-3. Pheretima micronaria 8
2-4. Allolobophora japonica 8
3.
Population densities and weight distribution of earthworms in area D
8
3-1. Pheretima schmardae
9
3-2. Pheretima sp. (H-1)
9
3-3. Pheretima vittata 10
3-4. Pheretima heterochaeta
10
3-4. Allolobophora caliginosa
11
4.
Population densities and weight distributions of earthworms in area G
12
4-1. Pheretima sp. (H-1)
12
4-2. Pheretima heterochaeta
12
4-3. Aporrectodea caliginosa 13
Discussion
13
1.
Faunas of earthworms in the suburb of Fukuoka city Kyushu Japan
13
2.
Life history and growth pattern
13
3.
Biomass of earthworm
15
Summary
16
Reference
17
Summary
1) The population studies of earthworms were performed in four areas in
the suburb of Fukuoka city Kyushu Japan.
2) Ten species of Pheretima and 4 species of Lumbricid were founded in
present study. Among them, six Pheretima and two Lumbricid were abundant.
3) Pheretima species founded in the suburb of Fukuoka city could be
grouped into three or four groups with regard to their life history, in
discussion together with the result on IBP Minamata area (Sugi and Tanaka 1978
a, b). Ph.
schmardae and Pheretima sp. (H-1) were founded in litter layer, all the earthworms
appeared in spring, matured in early summer and most disappeared till late
summer (Ia type). Ph.. vittata
and Ph.. irregularis were founded also in litter layer, all the
earthworms appeared in spring, matured in summer but remained till autumn (Ib
type). Pheretima sieboldi and Pheretima
sp. (M-1) were founded in litter layer except winter, all the earthworms
appeared in summer, hibernate at pre-matured stage and matured in summer of next
year (Second type). Ph. heterochaeta
and Ph. micronaria were founded in deeper zone and had a various stage
through a year but most younger appeared in summer, hibernate at pre-matured
stage and matured in early summer of next year (third type).
4) The biomass of each species changed with the difference of vegetation
type. Ph. schmardae
and Pheretima sp. (H-1) were most abundant in old grass field. Ph.
vittata was most abundant in forest
having a rich ground flora. Ph. irregularis was
founded only in small forest. Ph. sieboldi
and Pheretima sp. (M-1) were founded only in forest on mountain slope. Ph.
heterochaeta was most abundant in the
youngest vegetation and Ph. micronaria was most
abundant in the forest having a rich ground flora.
| Chapter One | Chap1FIGTAB |
Ecological
feature and feeding habit of Pheretima group (Oligochaeta: Megascolecidae)
41
Introduction
41
Method
41
1.
Distribution of earthworm in a grass field 41
2.
Eco-morphological feature and behavior of Pheretima 42
3.
Composition of gut content
42
Result
43
1.
The micro habitat distribution of earthworm
43
2.
Body size, body form and other ecological feature of Pheretima species
44
3.
Composition of gut content
44
Discussion
46
1.
Ecological feature of Pheretima group relating to their distribution
46
2.
Feeding habitat of Pheretima group in relation to their abundance
47
3.
Habitat, feeding habit and life history of Pheretima group
48
Summary
50
Reference
51
Summary
1) Ecological distribution of earthworm was investigated in the
experimental field of the Department of biology Kyushu University, which was
composed by two vegetation type. Litter dwellers: Ph.
schmardae, Pheretima
sp. (H-1), Ph.
vittata distributed
within the area having a plenty ground flora, thick A0 horizon. On the other
hand, Soil dwellers: Ph.
heterochaeta
distributed in whole area, regardless vegetation type and habitat structure near
soil surface.
2) Some eco-morphological feature such as body size, body form mobility
and the developmental degree of intestinal coeca, and feeding habit were
investigated on seven representative Pheretima species: Ph.
schmardae, Pheretima
sp. (H-1), Ph. vittata,
Ph. irregularis, Pheretima sp.
(M-3), Ph. heterochaeta and Ph. micronaria.
3) Soil dweller: Ph. heterochaeta and Ph.
micronaria had in a small body size
and slender body form, and showed a sluggish mobility compared with those of
litter dwellers. Among litter dwellers, Ph.
schmardae had the smallest body size,
and Pheretima sp. (M-3) had the
slenderest body form. It was discussed that the eco-morphological feature of
each species were largely reflected on their life history, their hibernating
behavior and distributions, in relation to the structure of their habitats.
4) There was large difference between every species in the gut material.
The main component of gut material were large particle of raw humus in Ph.
vittata, of small particle of raw
humus in Pheretima sp. (H-1), of small
particle of raw humus and organic rich material in Ph. irregularis, of
organic rich material in Pheretima
schmardae and Pheretima sp. (M-3),
and or organic rich soil and mineral material in Ph.
micronaria and Ph. heterochaeta. There
were observed a linear relation between the decayed degree of gut material and
the developmental degree of intestinal coeca, of each species. The feeding habit
of each species was discussed in relation to the quantity and quality of the
food resource presenting in their habitats.
5) It was discussed that Pheretima species found in suburb of Fukuoka
city had a different niche with each other, along three axes: life history,
habitat preference and food preference.
| Chapter Two | Chap2FIGTAB |
Population
metabolism of Earthworm populations
Introduction 59
Method 60
1
Measurement of various metabolic rate 60
1-1 Energy equivalent of animal tissue
60
1-2 Measurement of respiration
60
2.
Energy conversion of biomass and respiration rate
60
Result 62
1.
Measurement of various metabolic rate
62
1-1. Energy equivalent of animal tissue
62
1-2 Respiration rate
62
2.
Biomass, production and respiration of abundantly populations
63
Pheretima sp. (H-1) in area H
1968:
63
Pheretima sp. (H-1) in area D
1971:
63
Pheretima sp. (H-1) in area D
1972:
63
Pheretima sp. (H-1) in area G
1972:
64
Pheretima vittata
in area H 1968:
64
Pheretima irregularis
in area K 1968:
64
Pheretima heterochaeta
in area D 1971-1973:
64
Pheretima heterochaeta
in area G 1971-1972:
64
Discussion
65
Summary
67
Reference
67
Summary
1) The energy equivalent of animal tissue of Pheretima
sp. (H-1) was 21.133 ± 1.213 KJ g-1 dry wt. Respiration rate of Pheretima
sp. (H-1) were 58.5 - 111.23 CO2 for an individuals having one wet weight.
2) The energy budgets of several populations of Pheretima
species were estimated. The assimilation (P+R) of Pheretima
sp. (H-1) were 224.011 KJ m-2 in area H 1968, 356.008 KJ m-2 in area D 1971,
831.917 KJ m-2 in area D 1972 and 68.810 KJ m-2 in area G 1972. The assimilation
of Ph. vittata
was 194.765 KJ m-2 in area H 1968. The assimilation of Ph.
irregularis was 61.212 KJ m-2 in area
K 1968. The assimilation of Ph. heterochaeta
were 268.322 KJ m-2 in area D 1971-1973 and 759.304 KJ m-2 in area G 1971-1972.
3) The R/A ratio of Pheretima were ranged between 0.544 and 0.713 for
litter dweller Pheretima and 0.869-0.874 of soil dweller Pheretima. The values
of litter dweller Pheretima were slightly smaller than those of Lumbricidae
species.
4) P/B ratio, and P/Bmax of litter dweller Pheretima were ranged between
2.631 and 5.218, and between 1.141 and 2.313, respectively. Also, the values of
soil dweller Pheretima were 2.300-2.895 and 1.168-1.635, respectively. The
values of litter dweller Pheretima were larger than those of Lumbricidae
species.
| Chapter Three | Chap3FIGTAB |
Feeding
ecology of Pheretima sp. (H-1) (Oligochaeta:
Megascolecidae)
Introduction
79
Method
80
1.
Cultivation of earthworm 80
2.
Nutrient content and energy equivalent of the material used for the cultivation
81
3.
Nutrient contents of the materials contained in alimental canal of the
earthworms collected in field
81
Result
81
1.
Growth rate 81
2.
Food consumption and fecal pellet production
82
3.
Nutrient contents of the material contained in alimental canal of field
population
85
4.
Feeding ecology of Pheretima vittata 86
Discussion
87
Summary
89
Reference
89
Summary
1) Pheretima sp. (H-1) were
cultivated under three temperature condition, and fecal pellet production, food
consumption rate growth rate were determined.
2) The cultivated earthworms required 34 - 64 days for their maturation
against to 100 days of maturation period for the field earthworms. Also, the
maximum weight of the cultivated earthworms was attained to 4 - 4.5 g fresh wt
against 1.7 - 2.5 g fresh wt of the field earthworm.
3) Fecal pellet production of the cultivated earthworms were 1054 - 1540
mg dry wt for an individuals having one gram body wet weight.
4) Litter consumption of the cultivated earthworms were 374 - 590 mg dry
wt for an individuals having one gram body wet weight. Still, the litter
consumption rate of mature earthworms was smaller than the rate of immature
earthworms.
5) The quality of the material consumed by the field earthworms were
investigated with determining Carbon and Nitrogen content in the gut content of
the field earthworms. The carbon content in material was regarded as the more
fundamental indicator for the consideration on the metabolic rate of field
earthworms. Carbon content in the gut content of the field earthworm was 72.15
mg g-1 dry wt in average. On the other hand, Carbon content in the materials
consumed by the cultivated earthworm was estimated at 89.91 mg g-1 dry wt. From
these results, it was estimated that the food consumption of the field earthworm
was attained to 73.64 % of the rate of the cultivated earthworms.
6) It was thought that the lower growth rate and the lower weight of the
field earthworms than those of the cultivated earthworms might be due to the
lower resource consumption rate of the field earthworm than that of the
cultivated earthworms.
| Chapter Four | Chap4FIGTAB |
Digestive
efficiency of Pheretima sp. (Oligochaeta:
Megascolecidae)
Introduction
99
Method
99
1.
Digestive efficiency (consumption minus rejection) 99
2．Assimilation
efficiency 100
Result
101
1．Digestive
efficiency and the ratio of the food to total quantity consumed
101
2．The
relation between digestive efficiency and weight 101
3．The
effect of digestive efficiency on growth rate 101
4．Assimilation
efficiency of Production plus Respiration
102
Discussion
102
1．The
digestive efficiency of the cultivated earthworms 102
2．Several
figure of digestive efficiency
103
3．The
speculation on the excreta
103
Summary
105
Reference
105
Summary
1) The digestive efficiency of the cultivated earthworms was estimated
with the difference in the quantity of nitrogen and carbon between consumed
minus ejected. The estimated efficiency was in wide range from -200 % to 80 %.
2) The relation between the digestive efficiency and the quality of the
material consumed was approximated by following equation A = 0.58028 X - 12.624,
where X is the ratio of the food consumed to total material consumed. The
average ratio of the food consumed to total material consumed was estimated as
35.5 % in cultivation condition. The digestive efficiency of the cultivated
earthworms was calculated at 7.397 %.
3) Assimilation efficiency of the cultivated earthworm can be estimated
another process {A = (P+R)/C}. Average efficiency from (P+R)/C was calculated at
2.17 %. The values from (C-F)/C were far larger than that from (P+R)/C.
Particularly, the excess absorption (total absorption minus basal metabolism)
was larger in younger earthworm than in elder earthworm. It was discussed that
the excess energy assimilated by younger earthworms must be expended for their
mobility and their specific dynamic action to gain the larger weight increase.
4) The digestive efficiency of the field earthworm (A=(C-F)/C) could be
estimated at 1.895 %, the value of which was deduced from Carbon content in the
gut material of the field earthworms. This value was far lower than the average
value of the cultivated earthworm. It was discussed that the lower digestive
efficiency of the field earthworms than that of the cultivated earthworms might
be another reason why the weight and the growth rate were far lower in field
than in cultivation.
5) The equation between digestive efficiency and the quality of the
material consumed indicate that the earthworm consuming the nutrient poor
resource showed the minus digestive efficiency. From this, it was discussed on
the hidden production arising from the excretion of the nitrogenous substance
such as digestive fluid, muco protein and gut lining.
| Chapter Five | Chap5FIGTAB |
Production
of earthworms in Several Types of Vegetation
Introduction
114
Method
114
Result
115
1.
Seasonal change of biomass and respiration of entire earthworm
115
Area H 115
Area K 116
Area D 116
Area G 116
2.
Total assimilation of entire earthworms
117
3.
Litter consumption and fecal pellet production of earthworms
117
Discussion
117
1.
The biomass of earthworms in the suburb of Fukuoka city Kyushu Japan
117
2.
Biomass and activity of earthworm activity in various region 118
3.
Role of earthworms as energy consumer, litter decomposer and ploughman
119
Summary
120
Reference
121
Summary
1) Total assimilation of entire earthworms was estimated by using the
average ratio of R/A of Pheretima species.
2) The maximum biomass and total assimilation of earthworms were
0.926-10.653 g dry wt m-2, and 98.069-1573.184 KJ m-2y-1 respectively, in the
suburb of Fukuoka City Kyushu Japan. Earthworms were more plentiful in grass
field than in old forest. The maximum biomass of entire earthworms was largely
depend on the structure of soil surface, and the quality of food resource.
3) There were some difference in the size order between the maximum
biomass and the total assimilation. The earthworms in younger grass field show
larger total assimilation than those in elder grass field, in spite of smaller
maximum biomass in earlier than in later. This was due to the difference in the
age composition of dominant species between two habitats.
4) It was discussed on the contribution of entire earthworms to ecosystem
as energy consumer, litter decomposer and ploughman.
| Chapter Six | Chap6FIGTAB |
The
mass emergence of Earthworm Pheretima
sp. (Oligochaeta: Megascolecidae) on Fine Day after Rainy Day
Introduction
128
Method
129
1.
Counting the number of dead earthworm
129
2.
Measurement of soil respiration
129
3.
Respiration of wormcast and of soil-litter mixture 130
4.
Microbial increase in earthworm intestine 130
5.
Responses of earthworm to CO2 gas and N2 gas 131
Result
132
1.
Daily change of the number of death individual
132
2.
Daily fluctuation in soil respiration
133
3.
CO2 expiration rate of the soil-litter mixture and the wormcast
134
4.
Micro organisms in earthworm intestine
135
5.
The response of earthworm to various gases
136
Discussion
136
1.
The factor bringing about the mass emergence of earthworm
136
2.
How concentration of CO2 gas induce the behavior response of earthworm
137
3.
CO2 concentration in soil 138
4.
Why is the soil respiration high in the day of the rainy rising fine?
138
5.
Meaning of mass emergence and large amount of the death in the population study
139
Summary
140
Reference
141
Summary
1) The observation on mass emergence of Pheretima
sp. (H-1) were carried out on the bare land in the experimental field of
Department of Biology Kyushu University in the period from May to July for two
years 1971 and 1972. The mass emergence of Pheretima
sp. (H-1) occurred frequently on fine day after rain. Several individuals was
killed by solar radiation on bare land. Few individuals died on other days.
2) The response of Pheretima sp. (H-1) to harmful gases was examined under experimental
condition. Earthworm showed the abruptly creeping out behavior even to the small
increase of CO2 tension. They showed no behavior response by the continuously
exposing of pure Nitrogen gas.
3) The soil respiration rate on fine day after rain was two fold of the
rate on rainy days or dry days. The increase of soil respiration rate in soil
air on fine days after rain was the factor inducing the creeping our behavior of
earthworm.
4) Bacteria increased to 3-5 fold during the passage of materials through
the intestine of Pheretima sp. (H-1). It was inferred that the wormcast had more
plentiful flora of microbial population than the control soil.
5) Worm cast showed higher soil respiration rate and larger water holding
capacity than those of control materials. The higher respiration rate of
wormcast might be due to the high microbial activity besides the high water
holding capacity of the materials. The warm and wet condition on fine days after
rain stimulated the activity of microbial populations contained in wormcast and
then the soil respiration rate increased.
6) CO2 expiration rate was higher in wormcast (earthworm's habitat) than
in other areas. By the reason that earthworm piled up the pellets on soil
surface most abundantly in early summer, soil respiration in earthworm's habitat
might be highest in this season through a year. This might be the reason why
mass emergence of earthworm occurred only in this season.
7) Earthworm adjusted their density to the adaptive density by the
dispersion of individuals on fine days after rain. As the biological
conditioning of the earthworm’s habitat caused the mass emergence of
earthworm, the mass emergence of earthworm on fine days after rain was regarded
as the self regulation.
| Chapter Seven | Chap7FIGTAB |
The
distribution Pattern and the Habitat structure of Earthworm Pheretima
sp. (Oligochaeta: Megascolecidae)
Introduction
153
Method
154
1.
Seasonal change of distribution pattern of field population
154
2.
Regression coefficient between number of individuals and depth of wormcast layer
154
3.
The evaluation of earthworm to wormcast
155
4.
The disappearance rate of wormcast 155
Result
155
1.
Distribution pattern of individuals
155
1-1
The seasonal change of the value of ＊m/-m
index 155
The 1972 year population
156
1-2 ＊m/-m index value and mean crowding index value (＊m)
156
1-3 A common feature in distribution pattern of animals in field
157
2.
The regression coefficient between number of earthworm and thickness of wormcast
157
3.
The habitat preference of earthworm
159
4.
The disappearance rate of wormcast 160
Discussion
160
1.
Structure of habitat and distribution pattern of earthworm
160
Young 1 stage: 161
Young 2 stage in dry season: 161
Matured stage: 161
Post matured stage;
162
2.
Ecological meaning of wormcast relating to the life history of earthworm
162
Summary
163
Reference
164
Summary
1) The distribution pattern of Pheretima
sp. (H-1) were studied in relation to the structure of habitat through a life
span.
2) The spatial distribution passes a highly aggregated phase in spring.
Through the dispersed phase in later spring, the second aggregated phase
followed in late spring-early summer. When the wetter season come, earthworms
dispersed abruptly. After wetter season, earthworm distributed in patch.
3) There were linear relation ship between the population density and the
thickness of the wormcast. This relation could be approximated by following
equation Y = AX + B, where Y was the thickness of the wormcast and X was the
number of individual per unit area. It was though that the values of 1/A
represented the number of individual per one cm thickness of the wormcast. The
value of 1/A decreased through a life span, in roughly. Still, there was
tendency that the value of 1/A was high in dry days and low in wetter days.
4) The role of the wormcast as the shelter and as the food resource, for
earthworms was examined. The fundamental significance of the wormcast to the
life of earthworm could be summarized in three.
(a)
Wormcast have a porosity character, by which earthworms can move easily in soil
surface layer. Wormcast have a high water holding capacity. Then, wormcast
become a good shelter for earthworms to avoid a drying in dry days.
(b)
Wormcast have a plentiful flora of microbial population. These microbial
population induce a high soil respiration and then bring about the mass
emergence of earthworms on fine days after rain.
(c)
The decaying wormcast become to the alternatively food resource or earthworm.
5)
The value of 1/A inducing from the linear equations between the number of
individuals and thickness of wormcast was regarded as the integrated evaluation
of earthworm to wormcast as shelter and food resource.
| Chapter Eight | Chap8FIGTAB |
Resource
utilization and energy balance of Earthworm Pheretima
sp. (Oligochaeta : Megascolecidae) in old grass field
Introduction
175
1.
First estimation of the resource utilization 175
2.
Secondary estimation of the resource utilization 176
3.
Third estimation of the resource utilization 177
Discussion
178
1.
Do the absorbed energy satisfy the energy consumption?
178
2.
Three aims of ecological feeding studies 178
Summary
179
Summary
1) Three process of the resource utilization of animal were concerned for
the population of Pheretima sp. (H-1) in area D. And the energy balance of this
population was discussed.
2) If animal can exhibit owns ability of intrinsic food requirement also
in field, the energy required was estimated as 609.32 % of the energy
consumption (P + R). Then, this assumption might not reflect the true process of
feeding activity in field.
3) If animal share the available resource from litter supply and other
thing among members, the available energy was 117.6 % of the energy consumption.
4) The absorbed energy was estimated by integrating the data on the
fecal pellet production rate of field population and on the nutrient content of
gut content of the earthworm in field. The absorbed energy was attained to 107.3
% of the energy consumption.
5) The comparison on three process for feeding activity show that the
available energy and the absorbed energy, both would reflect the true process of
the resource utilization of animals under the field.
| Chapter Nine | Chap9FIGTAB |
Population
study of Earthworm Pheretima sp. (Oligochaeta:
Megascolecidae) under the artificial condition
Introduction
182
Study
area 182
Method
183
Result
183
1.
Average weight of individual and population density 183
2.
Metabolism of the experimental population
184
3.
Dispersing earthworm and dead earthworms
184
4.
Aggregation degree of the experimental population 184
Conclusion
185
Summary
185
Summary
1) The experimental population of Pheretima
sp. (H-1) was established on the clay sand area in the experimental field of
Kyushu University.
2) Maximum weight of individuals, and biomass, annual production and
total assimilation of population, these were far lower in the experimental
population than in the field population. The difference in bio-economic items
between two populations may be due to the difference in the quantity of the food
resource between each habitat. The experimental population showed the more
dispersed distribution of individuals than the distribution pattern of the field
population. The difference in distribution pattern between two populations may
be due to the difference in the uniformity of the environmental condition
between two habitats.
| Chapter Ten | Chap10FIGTAB |
The
bio-economic life table of earthworm Pheretima
sp.
-The
interaction between animal and environment-
Introduction
189
1.
The bio-economic life table of Pheretima
sp. (H-1)
190
1-1. The relation among resource supply, resource requirement,
aggregation pattern, population change and production.
191
1-2 The factors determining the abundance of earthworms 193
2.
The relation among each items
194
Summary
198
Reference
198
Summary
1)
The bio-economic life table of earthworm: Pheretima
sp. (H-1) was constructed. The bio-economic faces and the population faces were
concerned with mutual viewpoint by using the bio-economic life table.
2)
The relation among the items composing the bio-economic life table were arranged
in one schema, under the assumption that, in the mutual interaction between
animal and environment, the subject was animal themselves.
3)
The purpose for the construction of the schema on the mutual interaction between
animal and environment, the content of the items composing the schema, and the
way of the mutual interaction were characterized by comparing with the concept
of “Life system” advocated by Clark et al (1967).
| Chapter Eleven | Chap11FIGTAB |