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Investigations were conducted both in the field and the laboratory to determine some of the more important ecological factors affecting the efficiency of Trichogramma dendrolimi Matsumura, Telenomus dendrolimi (Matsumura), Anastatus japonicus Ashmead, A. gastropachae Ashmead and Pseudanastatus albitarsis (Ashmead), hymenopterous egg parasites of the pine moth, Dendrolimus spectabilis Butler. Field studies were made in the Japanese black pine forest on the coast of Fukuoka district between 1960 and 1967, and the majority of materials for laboratory studies were also collected from this area. Some biological characteristic s of the five egg parasites, e.g., fecundity, sex ratio, length of developmental period, adult longevity, pattern of ovigenesis etc. were studied. Characteristics of host population in time and space, e.g., seasonal prevalence of host population and spatial distribution of host egg masses in the pine forest were also studied in connection with parasitism by the egg parasites. In addition, hyperparasitism, predation of adult parasites by ants, alternative hosts, interspecific competition between parasites and source of foods for adult parasites were investigated. Conclusions to be derived from the comparative study of these ecological factors affecting the efficiency of the five principal egg parasites of the pine moth as natural enemies can be expressed as follows. /Trichogramma dendrolimi/ This species is the most prolific of the egg parasites of the pine moth (the mean fecundity, 186). There is little variation in the sex ratio, and an extremely high proportion of females (86-88% females) is always recorded. This is probably due to its uniparental reproduction (deuterotokous parthenogenesis). The developmental period of this species is the shortest of the erg parasites of the pine moth (the mean developmental period in 28℃, 8 days). This makes it possible for the parasite to complete three generations per one generation of the host. The above-mentioned characteristics of the parasite indicate that its potential rate of increase is exceedingly high. Consequently, the parasite can respond rapidly to a greater increase in the host population during a host oviposition period. A high level of parasitization at the end of the host oviposition period in the first generation is primarily caused by a rapid increase in parasite populations. This species is pro-ovigenic, and the adult is short-lived (the mean longevity of females fed on honey in 28℃, 11 days). These characteristics are disadvantageous to the parasite in the case of the second host generation in which the host population density is usually very low. The short longevity of the adult females indicates that they can not survive in the period between the first and second host generations. These are the reasons why host eggs in the second generation are rarely attacked by the parasite every year. Although many lepidopterous insects are known to be hosts of the parasite, they are all the inhabitants of other types of forests than pine forests. The absence of alternative hosts in the period between the first and second host generations is another reason for a low level of parasitization in the second host generation. Since there are no overwintering host eggs in the pine forest, the initial parasitization in the first host generation is probably due to the parasites which happen to enter the pine forest from outside areas. It is, therefore, apparent that the absence of alternative hosts in the pine forest is an important factor limiting the efficiency of the parasite in both of the two host generations. A high degree of gregarious parasitism inherent in this species (mean number of parasites produced in each host egg, 15-29) decreases the efficiency of the parasitism, so that each female parasite can parasitize only nine host eggs on an average during its life span. Spatial distribution of host egg masses in the pine forest influences t he efficiency of this parasite; the more the large-sized egg masses are deposited in a pine tree, the heavier the parasitization of the egg masses occurs, because large-sized egg masses are more frequently deposited in taller pine trees especially in their higher crowns which correspond to the preferential range of the female parasite. Hyperparasitism by Pacheneuron sp. is very rare and is not an important factor limiting the efficiency of the parasite. /Telenomus dendrolimi/ In spite of the relatively low fecundity of this species (the mean fecundity, about 70), its potential rate of increase is considerably high, because the developmental period is so short (the mean developmental period in 28℃, 12 days) that two generations of the parasite can be produced per one generation of the host. Consequently, the parasite is able to increase in number rapidly and to attain to its high population density at the end of the host oviposition period. This species exhibits arrhenotokous parthenogenesis. Male progeny produced by unmated females may reduce the proportion of females in the parasite population. However, such a phenomenon rarely occurs in the field, and a great preponderance of females (75-86% females) is usually recorded there. This species is pro-ovigenic, but the adult females fed on honey can live long (the mean longevity in 28℃, about 50 days). The combination of these two desirable characteristics as control agents suggests a high power of host discovery of the parasite followed by a high efficiency of egg deposition. Furthermore, the great longevity of the female parasite suggests that she can survive in the period between the first and second generations of the host during which no host is available. For these reasons, the high adult parasite population, which has already grown at the end of the host oviposition period of the first generation, can attack host eggs in the second generation period, and a high level of parasitization often occurs in the host generation regardless of the host population density. High host populations are occasionally observed in the second host generation, while this host generation is usually produced partially in the study area. Such a great abundance of hosts would favour the efficiency of the parasite, because the parasite can cause a severe mortality of the pine moth eggs in the second generation irrespective of the host population density. In contrast to Tr. dendrolimi, this parasite tends to attack the host egg masses in the lower crown of a pine tree. Such a failure to fully occupy a host-inhabited area may reduce the efficiency of the parasite as in the case of Tr. dendrolimi, but it prevents the former from decreasing its efficiency by avoiding the multiparasitic competition with the latter. Because this species is characterized by a lower degree of gregarious parasitism than Tr. dendrolimi (mean number of the parasites produced in each host egg, 5-7) in spite of its low fecundity, each female of the former can parasitize a few more host eggs than that of the latter during her life. Hyloicus caligineus Butler, only one alternative host of this parasite in the pine forest, is always very scarce and seems of little importance except for a brief period of spring when no pine moth eggs are available in the forest. /Anastatus japonicus, A. gastropachae and Pseudanastatus albitarsis/ Some factors common to these three related species influence their efficiency. These species are all synovigenic, and the adults are long-lived ( the mean longevity of females fed on honey in 28℃, 53 days in A. japonicus, 45 days in P. albitarsis and 28 days in 1. gastropachae respectively). It follows from this that the relatively high fecundity (the mean fecundity, e.g., 153 or more in. A. japonicus) and the great longevity of these species are largely dependent on the availability of the source of foods for adult parasites such as nectar-bearing plants and honeydew-producing insects in the pine forest. Therefore, the great scarcity of these kinds of food sources in the pine forest is an important factor limiting the efficiency of these parasites, The synovigenic gravid females of these species are characterized by a phenomenon of ovisorption. Low host densities would be advantageous to these species because of conservation of reproductive material based on ovisorption. However, high host densities attained in a short time during the host oviposition period as in the first host generation are disadvantageous to these synovigenic species because of their preoviposition period and of limited egg deposition in a brief period. The long developmental period of these species (the mean developmental period in 28℃, about 20 days in A. japonicus and A. gastropachae and 16 days in P. albitarsis respectively) indicates that they are unable to have more than one generation during one generation of the host and can not respond rapidly to a remarkable change in the host density during the host oviposition period. This is the main reason why a large number of host eggs in the first generation are usually able to escape from the parasitization by these species. These species exhibit arrhenotokous parthenogenesis. Male progeny produced by unmated females may reduce the proportion of females in the parasite population. Furthermore, in the laboratory the proportion of male progeny of A. japonicus and A. gastropachae increases toward the end of the life of females. However, such phenomena rarely occur in the field, and a great preponderance of females (84% females in A. japonicus, 79-86% females in A. gastropachae and 75-79% females in P. albitarsis) is usually recorded there. These species are all solitary in the pine moth egg. Undoubtedly, this parasitic habit gives the parasitism a greater efficiency as compared with gregarious parasites, Tr. dendrolimi and Tel. dendrolimi. These species may live the whole year on only pine moth egg without alternative hosts as far as the study area is concerned. Neverthless, there is very little doubt that the absence of alternative hosts in the pine forest is reducing the efficiency of these species.
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マツカレハの卵寄生蜂の主要種であるキイロタマゴバチ,マツケムシクロタマゴバチ,フタスジタマゴバチ,マツケムシハネミジカヤドリバチおよびシロオビタマゴバチの5種について,これらの天敵としての有効性に関与する生態学的諸要因を決定するため,1960年から1967年まで福岡付近のクロマツ海岸林で野外調査を実施するとともに室内実験を重ねた.これら卵寄生蜂の天敵としての有効性に関与する要因として,まず,各寄生蜂種の生態的特性,寄主マツカレハ卵個体群の時間的,空間的分布特性,およびこれら以外の要因に大別し,さらに各細かな要因に分析して比較検討した.その結果は次の通りである. /1.キイロタマコバチ/ この種の産卵能力はマツカレハの卵寄生蜂中最も高い.その性比は変動が少なく,雌の割合が常に高いが,これは産雌雄性単為生殖を行なうためと考えられる.1世代の所要日数は他種にくらべて最も短かく,寄生の1世代に対して3世代を繰り返すことができる.以上述べた2,3の特性は本種がきわめて高い増殖能力を持つていることを示唆しており,このためマツカレハの産卵期間中の著しい寄主密度の増加に対応することが可能である.第1世代のマツカレハの産卵末期にしばしばみられる高率の寄生はこの蜂の個体群密度の急速な増加によるものである.この種はpro-ovigenicな卵形成を示し,同時に成虫の寿命も非常に短かい.このような特性は寄主第2世代のように寄主密度が著しく低い場合にはきわめて不利に作用する.また,寿命が短かいので寄主の第1世代と第2世代の間の時期をこの蜂の雌成虫は生きのびることができない.これらの理由で,例年寄主第2世代はほとんどこの蜂の寄生を受けることがない.多数の鱗翅目昆虫がこの蜂の寄主としてこれまで知られているが,それらはすべてマツ林外に生息しているもので,マツ林内には代替寄主を欠いている.寄主の第1世代と第2世代の間の時期にマツ林内に代替寄主がないことも,第2世代でこの蜂の寄生が低い理由の1つに数えられる.この蜂の越冬寄主もマツ林内にはなく,寄主第1世代当初の寄生は林外からたまたま飛来する蜂によるものと思われる.したがつて,マツ林内にこの蜂の代替寄主が欠如していることが,寄主の両世代を通じてこの種の有効性を限定する重要な要因であるといえる.この種は極度な多寄生が正常なので,この習性はこの蜂の寄生の効率を著しく低いものにしている.マツ林における寄主卵塊の空間的分布はこの蜂の有効性に影響する.すなわち,大型の寄主卵塊が多く産まれたマツの単木ほどそれらの卵塊は高率の寄生を受けるが,これは大型の寄主卵塊がこの蜂の雌の選好域である高い木の特にその樹冠上方部によく産まれるためと考えられるからである.二次寄生蜂であるPachyneuron sp.の寄生率はきわめて低く,その影響は無視できる. /2.マツケムシクロタマゴバチ/ この種の産卵能力は比較的低いけれども,その増殖能力はかなり高い.なぜなら,寄主の1世代に対して2世代を繰り返せるほどその発育期間が短かいからである.このために,この蜂は第1世代のマツカレハの産卵末期には高い密度に到達することができる.この蜂は産雄性単為生殖を行なうから,未交尾雌による雄の子孫の産出が蜂の個体群中の雌の比率を低下させることも考えられるが,野外ではそのようなことは稀で,本種の性比は雌の割合がきわめて高いのが普通である.この種はpro-ovigenicな卵形成を示すにもかかわらず,成虫の寿命はきわめて長い.これら2つの天敵としての望ましい特性が組み合わさるため,この蜂は寄主探索の能率も高く,また一度寄主を発見するとその産卵能力を短期に効率よく発揮できると思われる.また,成虫の寿命も長いため,寄主の第1世代と第2世代の間の時期を雌成虫は生きのびることができる.したがつて,既に第1世代のマツカレハの産卵末期に達していた高密度を代替寄主なしに寄主第2世代まで維持でき,さらに寄主密度の低いこの世代においても高率の寄生を示すことができる.寄主第2世代は第1世代のごく一部の発育の進んだ個体に由来するもので,その密度は例年著しく低いが,年によつては高密度の個体群が出現することがある.この蜂は寄主第2世代には寄主密度の高低にかかわりなく高率の寄生を示すから,そのようにこの世代の寄主密度が高い年には必然的にこの蜂の有効牲は増進されることになる.キイロタマゴバチとは逆に,この蜂はクロマツの樹冠下方部の寄主卵塊に好んで寄生する傾向がある.このように,寄主の生息域の一部にしか蜂が寄生しないことは,キイロタマゴバチの場合と同様,その有効性が限定されることにもなるが,一方,キイロタマゴバチとの共寄生を避けることによつて両種間での競争から生ずる有効性の低下を防止していることにもなつている.この種の多寄生の程度はキイロタマゴバチにくらべてずつと低いため,より効率の高い寄生が可能で,その産卵能力が低いのにもかかわらず,キイロタマゴバチを上回る数の寄主に寄生できる.クロスズメはこの種の唯一の代替寄主としてマツ林に生息しているが,その卵密度は年間を通じて非常に低いので,越冬後の蜂が春季の一時期,マツカレハ卵の林内にない時期に寄主として利用する場合を除けば,それほど重要ではないと思われる. /3.フタスジタマゴバチ,マツケムシハネミジカヤドリバチ,シロオビタマゴバチ/ これら3種の近縁な寄生蜂はその有効姓に関与する要因が共通している点が多いので,一括して述べる.これらの種はすべてsynovigenicな卵形成を示し,成虫の寿命は長い.このことから,その比較的高いと思われる産卵数も,また著しく長い寿命も,花蜜を出す植物や甘露を分泌する昆虫など成虫の食物源となるものがマツ林内部に確保されているか否かによつて大いに左右される.したがつて,実際にはマツ林の中にこれらの食物源がきわめて少ないことがこれらの蜂の有効性を制限する重要な要因になつている.これらの種の雌成虫にはその卵巣が成熟するとともに卵吸収という現象がみられる.この現象によつて一定の産卵能力を常に保持できるので,寄主の密度が低い場合はこれらの種に有利であると考えられる.しかしながら,寄主第1世代のように,産卵期が短かく,しかも高密度に達するような場合には,これらsynovigenicな種は産卵前期間を持ち,また短期にはその産卵能力が十分発揮できないため不利である.これらの種の発育期間はいずれも長いので,蜂は寄主の1世代に対し1世代しか送れない.そこで,マツカレハの産卵期間内の寄主密度の著しい変化に対応することもできない.このことが寄主の第1世代では通常大量の寄主卵がこれらの蜂の寄生を免がれるということの主要な理由である.これらの種はいずれも産雄性単為生殖をするから,未交尾雌による雄の産出が蜂の個体群中の雌の比率を低下させる可能性がある.また,室内ではフタスジタマゴバチやマツケムシハネミジカヤドリバチで雌がその生存期間の後半になるにしたがつて雄の子孫を多く産出することがわかつている.しかしながら,野外ではこれらの現象はほとんどみられず,これらの種の性比は普通雌の割合がきわめて高い.これらの種はすべて単寄生をする.このことは多寄生をするキイロタマゴバチやマツケムシクロタマゴバチにくらべて寄生の効率からみればはるかに勝つており,これらの種の天敵としての有利な特性の1つとみられる.これらの種は調査地域に関する限り,代替寄主がなくとも,マツカレハ卵だけで年間を通じ世代を続けているようである.しかし,やはりマツ林内に代替寄主がないことが,これらの蜂の有効性を制限していることも疑いのない事実と思われる.
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