椰心叶甲的耐寒力测定

测定椰心叶甲Brontispalongissima(Gestro)的过冷却点和结冰点。各虫态中过冷却点值以卵和蛹最低,分别为-9.8℃和-9.0℃,成虫较高,雌、雄成虫分别为-5.9℃和-5.5℃;幼虫的过冷却点最高,平均为-4.5℃。结冰点由高到低依次为2龄幼虫、5龄幼虫、4龄幼虫、1龄幼虫、3龄幼虫、雄成虫、雌成虫、蛹和卵,分别为-0.3,-0.8,-1.1,-1.3,-1.3,-2.4,-2.7,-5.4和-5.5℃。综合分析认为,蛹、卵的耐寒能力最强。根据广东地区常年平均气温分析了在自然条件下该虫各虫态越冬情况,但由于生物学特性的限制,越冬虫态主要是成虫、蛹。     

甜菜夜蛾在鲁西南地区越冬问题的初步探讨

通过对甜菜夜蛾Spodopteraexigua越冬蛹过冷却点、各虫态发育起点温度、有效积温测定,结合10年来田间调查及越冬观察试验,表明甜菜夜蛾在鲁西南地区一般年份不能安全越冬,得出了每年初代虫源来于外地的结论,这对甜菜夜蛾预测预报和综合防治有着重要的指导作用。     

粘虫在贵州成功越冬的实证

粘虫Mythimna separata(Walker)是一种典型的季节性远距离迁飞害虫,也是我国及其它亚洲和澳洲国家粮食作物上重大害虫。贵州省地处我国西南部,是二代粘虫的常年发生地,但有关粘虫在贵州的越冬问题未有详细报道。为探明粘虫是否可以在贵州越冬,本文对2日龄蛹过冷却点,耐低温能力进行了测定,并在田间进行了越冬笼罩实验和越冬调查试验。结果表明,粘虫的过冷却点为-14.95℃,结冰点为-9.52℃。低温和暴露时间对粘虫的存活率具有显著影响。在10℃下处理30 d后,其存活率高达96.5%,处理90 d后仅有5%个体存活。但在-5℃下处理0.5 h后的存活率为80.00%,而处理8 h就全部死亡。田间笼罩试验结果发现2日龄蛹的存活率随埋蛹时间延长而降低,但至100 d仍有46.67%个体存活;同时,越冬调查发现少量的蛹和老熟幼虫,包括1头老熟幼虫和29头蛹。这些结果表明,粘虫可以以老熟幼虫和蛹等虫态在贵州低海拔地区成功越冬。     

上海地区红棕象甲的耐寒性研究

红棕象甲Rhynchophorus ferrugineus (Olivier)是我国危害棕榈科植物的重要入侵害虫, 为探明其越冬抗寒性, 利用过冷却点测定仪测定了红棕象甲上海种群不同虫态的过冷却点, 在低温箱内测定了该虫不同虫态的耐寒性, 然后结合田间越冬模拟试验、气象资料和寄主分布情况, 初步确定了红棕象甲在我国的越冬北界。结果表明： 红棕象甲过冷却点随虫态的发育程度的升高而下降, 卵、1龄幼虫、5龄幼虫、9龄幼虫和成虫的平均过冷却点分别为-5.92, -6.42, -7.19, -7.43和-11.84℃, 过冷却点由高到低的顺序依次为: 卵>幼虫>成虫。在6, 24, 48和72 h 4个时间处理下, 各虫态在低温与存活率之间呈显著或极显著的logistic回归, 半致死温度（Ltemp₅₀）均随处理时间的延长而上升, 不同虫态在处理72 h 后, 卵、1龄幼虫、5龄幼虫、9龄幼虫和成虫之间的Ltemp₅₀分别为1.61, -1.67, -2.39, -2.40和-0.40℃, 各虫态耐寒性由弱到强的顺序依次为: 卵<成虫<幼虫。红棕象甲不同发育阶段的过冷却点与其耐寒性并不完全相关, 幼虫和成虫均可能是该虫的越冬虫态。连续两年的田间模拟越冬试验表明, 在上海地区, 红棕象甲的幼虫和成虫的越冬存活率均在60％以上, 说明红棕象甲在上海地区是可以越冬的。根据这些结果, 结合寄主分布情况, 初步将红棕象甲在我国的越冬北界定于北纬35°附近, 即1月份0℃等温线左右。     

双委夜蛾不同虫态耐寒性及体内生化物质含量变化

【目的】探索双委夜蛾Athetis dissimilis(Hampson)的耐寒策略及越冬虫态。【方法】以室内双委夜蛾种群为虫源,分别测定3龄、4龄、5龄、6龄和老熟幼虫以及蛹的过冷却点(supercooling point,SCP)、结冰点(freezing point,FP)、含水量以及脂肪、蛋白质、甘油、山梨醇和海藻糖含量。【结果】不同龄期幼虫和蛹之间过冷却点和结冰点存在显著差异,低龄幼虫SCP和FP较高;随着龄期增长,二者均降低,老熟幼虫的SCP和FP最低,分别为-18.20℃和-11.72℃。不同龄期幼虫和蛹的生化物质含量均存在显著差异。随着龄期的增加,含水量和甘油含量呈降低的趋势,蛹的含水量(65.33%)和甘油含量(154.90μg/mg)最低;脂肪和蛋白质含量随龄期的增加呈现增加的趋势,蛹的脂肪和蛋白质含量最高,分别为25.36%和2 298.37μg/mg。老熟幼虫海藻糖含量相对较高(57.28 mg/g),而山梨醇含量较低(15.49 mg/g)。【结论】与其他龄期相比,双委夜蛾老熟幼虫和蛹的耐寒性最强,是最可能的越冬虫态;脂肪、蛋白质和海藻糖可能是双委夜蛾重要的耐寒物质。     

桔小实蝇不同发育阶段过冷却点的测定

对桔小实蝇Bactrocera dorsalis (Hendel)不同发育阶段的过冷却点进行了测定。结果表明：同一虫期个体间的过冷却点出现不同程度的变异,但均服从正态分布。不同虫期的过冷却点差异显著,其中蛹的过冷却点最低（-12.2℃～-15.0℃）。老熟幼虫的过冷却点为-8.1℃。成虫的过冷却点最低值为7日龄雄虫（-10.5℃）和雌虫（-10.1℃）,最高值为60日龄雄虫（-5.9℃）和雌虫（-6.4℃）,但同一发育时期的雌、雄成虫之间的过冷却点没有差异。测定结果提示蛹期最有可能是该虫在温带地区越冬的虫态。     

警纹地老虎Euxoa exclamationis(Linnaeus)的初步研究

1.警纹地老虎在新疆各地与黄地老虎同时混合发生,各虫态又与黄地老虎相近似,因而两种地老虎常常混淆不清,给研究和测报工作带来一定的困难。本文对它与黄地老虎的形态做了比较,生物学特性作了叙述。 2.此虫在南疆莎车地区一年发生二代,以滞育的第二代幼虫在土中越冬。越冬深度较黄地老虎为浅。成虫发生期为4月中旬至9月中旬,其间出现两次高峰,第一次为4月下旬至5月上旬,第二次为7月上中旬。 3.警纹地老虎虽是杂食性害虫,但其寄主植物范围较黄地老虎为狭。第一代幼虫一般生活在藜科杂草上,作物地除苜蓿外很少发现它的踪迹:第二代主要集中苜蓿、甜菜和马铃薯等作物为害。 4.各虫态历期长短随温度而异。卵期当温度为20℃及26℃时,分别为8-10天、4-6天;蛹期在17-18℃及25-27℃时,分别为25-28天、12-14天;幼虫期第—代以灰藜、甜菜、玉米饲养者,分别为26-33天、33天、37-38天。 5.成虫取食马蔺花蜜者产卵量较取食白菜花及清水者高1.29、2.26倍。幼虫取食甜菜者其成虫产卵量较取食玉米者高1.74倍,蛹重以取食灰藜和甜菜者较重,取食马铃薯和玉米者较轻。     

Pb2+在甜菜夜蛾体内的积累及对种群的影响

重金属的污染及其治理是目前倍受关注的生态学问题之一.甜菜夜蛾是我国长江流域及以南地区蔬菜、棉花等作物上的主要害虫,近几年突发趋势明显.本文以甜菜夜蛾为靶标对象,设计5个浓度(0.3、1.2、4.8、19.2、76.8 mg · kg^-1)重金属Pb²⁺处理,探讨处理后甜菜夜蛾各虫态体内Pb²⁺的积累和排泄,并根据甜菜夜蛾各虫态的生长与存活情况,构建了该虫连续3个世代的种群生命表.结果表明: 取食含有不同浓度Pb²⁺的人工饲料后,各虫态甜菜夜蛾体内Pb²⁺的浓度在同一世代不同处理间存在显著差异,且呈显著的剂量 反应关系.各虫态体内Pb²⁺的浓度基本上随胁迫世代数的延长而增加.受Pb²⁺胁迫后,各虫态体内Pb²⁺的浓度顺序为5龄幼虫＞成虫＞蛹;该虫可通过粪便、预蛹蜕和蛹壳排泄体内部分重金属,且粪便和预蛹蜕中的Pb²⁺浓度远大于蛹蜕中的浓度;根据Pb²⁺不同浓度条件下甜菜夜蛾各虫态的存活率、雌性比和产卵量,构建的种群生命表表明,低浓度Pb²⁺胁迫可促进甜菜夜蛾的种群增长,而高浓度胁迫使种群增长衰退;且随着重金属的持续胁迫,其种群衰退更明显.研究结果为长期、客观评价重金属的危害性以及对重要农业害虫的种群影响提供了借鉴.     

美国白蛾过冷却点的影响因子

美国白蛾是一种重要的国际检疫害虫,作为外来入侵生物在中国不断扩散蔓延。过冷却点是研究其潜在适生区重要的生理指标之一。本文研究了不同虫态、寄主树木、低温驯化时间和地理种群与过冷却点的关系。结果表明:美国白蛾不同虫态过冷却点间存在显著差异(P0.001),蛹期过冷却点最低(-15.91℃),5龄幼虫的过冷却点最高(-6.97℃),过冷却点随虫态龄期增加而升高;用悬铃木饲养的美国白蛾蛹的平均过冷却点为-16.09℃,显著低于复叶槭饲养的美国白蛾蛹的平均过冷却点-14.61℃(P=0.002);在6℃下分别驯化美国白蛾非滞育蛹24、48和72 h,美国白蛾非滞育蛹过冷却点无显著差异(P=0.846);武邑、内黄、濮阳、东营、寿光和沈阳6个不同地理种群美国白蛾蛹的过冷却点间存在极显著差异(P0.001),其中内黄地区美国白蛾过冷却点最低(-21.39℃),东营地区过冷却点最高(-15.35℃)。     

食用菌厉眼蕈蚊Lycoriella ingenua在温度逆境下的存活率

以食用菌重要害虫-厉眼蕈蚊属的Lycoriella ingenua为研究对象,研究了不同逆境温度下各虫态存活率的差异。结果表明温度对不同虫态存活率的影响存在显著性差异。成虫和蛹对高温具有一定适应性,卵对高温的适应性最低但具有一定的低温耐受力。在35-40℃高温下,相同暴露时间,各虫态的存活率随温度的升高而降低,但成虫的存活率均高于其它虫态。38℃下全部死亡的暴露时间分别为:成虫48 h,蛹24 h,幼虫12 h,卵2h。在40℃达到全部死亡的暴露时间分别为:成虫24 h、蛹12 h、卵和幼虫0.5 h。低温处理下,各虫态随着暴露时间的延长存活率逐渐降低。5℃低温下,相同暴露时间,成虫存活率均高于其它虫态,但在0℃低温下暴露24 h后全部死亡,显著高于其它虫态;幼虫在低温下取食活动减弱,5℃下暴露24 h后停止取食,相同暴露时间下随着温度的降低存活率下降;蛹对0℃的耐受力高于成虫和幼虫,暴露4 h后可以正常羽化,48 h后全部死亡;卵对短时间的低温具有一定耐受力,但随着暴露时间的延长存活率降低幅度大。     

Reproduction-flight relationship in the beet armyworm, Spodoptera exigua (Lepidoptera: Noctuidae)

To evaluate the relationship between reproduction and migratory flight, we examined ovaries, tethered flight, and egg laying in the beet armyworm, Spodoptera exigua (Hübner). Ovarian development in female S. exigua was classified into five stages in relation to the postemergence age of the moths. Oviposition peaked at ovarian stages III and IV. Mating accelerated ovarian development in this species, and thus the mated females developed faster than the virgin moths in ovaries. In virgin adults, flight capacity increased from the first day to the third day after emergence and remained at a high level until the moths were 7 d old. In contrast, the mated moths reached the maximum flight capacity in 2 d after emergence and maintained a high level until the age of 7 d. However, mating status could not influence the flight capacity of the beet armyworm, although there were general differences in flight capacity between virgin and mated adults at the same age. Moreover, significant correlations between the number of eggs laid and the flight capacity were not performed by the mated moths. These results showed that mating status and ovarian development did not seem to influence flight capacity in the beet armyworm, which was not in agreement with the definitions of the oogenesis-flight syndrome observed in many other migratory insects.     

Does migration promote or restrict circumpolar breeding ranges of arctic birds?

Aim Migration has been suggested to promote large breeding ranges among birds because of the greater mobility of migratory compared with non‐migratory species, but migration has also been suggested to restrict breeding ranges because of evolutionary constraints imposed by the genetically based migration control programme. We aim to investigate the association between migration and the breeding ranges of both land birds and pelagic birds breeding in the Arctic region. Location The Arctic region. Methods Information on breeding and wintering ranges and migratory status of bird species breeding in the arctic tundra biome was compiled from the literature. The association between breeding range, migration distance and primary winter habitat was tested using multivariate generalized linear models and pair‐wise Mann–Whitney U‐tests. Phylogenetic effects were tested for using Mantel’s permutation tests. Results We found different relationships depending on the species’ major winter habitat. Among birds that are pelagic during winter, long‐distance migrants have the largest breeding ranges, while among terrestrial birds, residents and short‐distance migrants have the largest breeding ranges. Breeding ranges of coastal birds of all migratory distance classes are comparatively restricted. Main conclusions As a new explanation for this pattern we suggest that the possibility of colonizing large winter ranges is a key factor for the subsequent expansion of breeding ranges in arctic bird communities and possibly also in bird communities of other regions of the world. Because of the reversal in the relative extent of continents and oceans between the hemispheres, longitudinally wide winter ranges are more likely for long‐distance than short‐distance migrants among pelagic birds, while the reverse holds true for birds that use terrestrial winter habitats. For coastal birds both continents and oceans form barriers restricting colonization of extensive winter quarters and consequently also of extensive breeding ranges, regardless of the distance to the winter quarters.     

甜菜夜蛾飞行肌中与能量代谢有关的酶活性

通过生化方法测定了参与飞行能源物质代谢的5种酶活力的变化,即3-磷酸甘油醛脱氢酶(GAPDH)、3-磷酸甘油脱氢酶(GDH)、乳酸脱氢酶(L DH)、柠檬酸合酶(CS)和3-羟酰辅酶A脱氢酶(HOAD)。结果表明,1、3、5日龄处女蛾的GAPDH和CS酶活力显著高于7日龄蛾;但HOAD的活性恰恰与之相反,其酶活力随日龄的增加逐渐增强。GDH初羽化时活性较低,但到3日龄后增加到最大值,以后随日龄的增加活性逐渐下降,L DH的活性一直比较低,且不同日龄的酶活力差异不显著。交配状况和幼虫期饥饿处理对成虫的代谢酶活力没有显著影响。在第7代选育家系中,强飞家系的GAPDH酶活力显著高于短飞和不飞;而在第8代中强飞家系的上述5种酶的活性均高于短飞家系,而第10代中仅有CS的活性在强飞和短飞家系中存在显著差异,其他均差异不显著。这说明甜菜夜蛾在初始阶段主要利用糖类作为能源物质,而后期飞行又可通过脂类代谢提供能量,且不同日龄的处女蛾和选择家系的飞行能力与代谢酶活性呈现一定的正相关关系。     

Relationship of flight and reproduction in beet armyworm, Spodoptera exigua (Lepidoptera: Noctuidae), a migrant lacking the oogenesis-flight syndrome

The beet armyworm, Spodoptera exigua, undertakes long-distance migration. We used flight mills to investigate the interaction between flight and reproduction in this species given the apparent absence of the oogenesis-flight syndrome. This syndrome, common in many migratory insects, is characterized by a suite of traits including migration during the pre-oviposition period followed by a switch to oogenesis. No negative effects of inter-ovipositional flight on lifetime fecundity were observed. Instead, adult reproductive output suffered when female flight was initiated the first day after eclosion and before oviposition, suggesting that migratory flight overlaps with the oviposition period rather than being confined to the pre-oviposition period. Mating status of both females and males had no negative influence on their flight performance except that flight distance and flight duration of 7-day-old mated females were significantly less than in unmated females. Furthermore, the number of eggs produced and mating frequency of females less than 7 days of age were not significantly correlated with flight performance, suggesting reproductive development paralleled and was independent of migratory behavior. This independent relationship between flight and reproduction of adults is consistent with the very short pre-oviposition period in this species, and suggests that resources are partitioned between these activities during pupal development. Together, our results uncovered neither obvious trade-offs nor mutual suppression between flight and reproduction in S. exigua, which indicates the lack of an oogenesis-flight syndrome for coordination of these two energy-intensive processes. We propose a conceptual model of migration for this species based on the current and previous studies.     

温、湿度对粘虫蛾飞行能源物质利用的影响

系统研究了不同温、湿度对粘虫蛾飞行能源物质消耗的影响,结果表明,温、湿度对成虫飞行能源物质消耗有显著影响,在适宜的温、湿度下飞行时,粘虫蛾主要飞行能源物质（甘油三酯和糖原）消耗最少,其飞行单位距离所需的甘油三酯也最少,即能源利用效率最高,随着温,湿度从适宜到不适宜,甘油三酯消耗有逐渐增多的趋势,但温,湿度对成虫飞行能源物质消耗会因蛾龄及性别的不同而异,在所测试的几种温,湿度下（高温35℃除外）,5日龄成虫在飞行中消耗的甘油三酯最多,低龄和高龄时消耗均较少,高温下飞行时,雄蛾比雌蛾消耗的能源物质多,而在高温或低湿条件下,雌蛾比雄蛾消耗的多,进一步对影响成虫飞行能源物质消耗的海藻糖酶活力及呼吸强度研究表明,温、湿度对海藻糖酶活性有明显影响,在适宜的温,湿度下,酶的活性高,反之,酶的活性降低,其中高温对酶活性的抑制作用比低温明显,但高湿和低湿对酶活性的抑制作用基本一致,成虫呼吸强度随温度的升降而升降,飞行初期成虫呼吸强度急剧增加,但0.5h后开始下降并维持在一个较稳定的水平,根据所获结果对温、湿度与粘虫蛾飞行能源物质消耗之间的内在联系进行了讨论.     

粘虫产卵前期的遗传特征

从田间采集的粘虫 Mythimna separata (Walker)种群在室内相同条件下饲养时 ,其后代雌蛾的产卵前期有较大程度的分化 ,最短的为 3d,最长的达 8d。通过亲子回归法 (雌性子代和亲本产卵前期的回归分析 )估算的粘虫蛾产卵前期遗传力值为0 .2 899,表明粘虫产卵前期受遗传因素的影响较小。对单雌系的双向选择所产生的选择响应表明 ,在一定的选择压力下 ,选择有利于产卵前期长的个体 ,即产卵前期长的个体进化速度要比短的个体快 ,但连续选择的响应较低。进一步通过家系间交互杂交来研究雌蛾产卵前期的遗传基础表明 ,产卵前期在很大程度上受雄性亲本的影响 ,即可能受 X-性染色体上的基因调控     

Host plant resistance in romaine lettuce affects feeding behavior and biology of Trichoplusia ni and Spodoptera exigua (Lepidoptera: Noctuidae)

Lettuce quality and yield can be reduced by feeding of several lepidopterous pests, particularly cabbage looper, Trichoplusia ni (Hübner), and beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae). Host plant resistance to these insects is an environmentally sound adjunct to conventional chemical control. In this study we compared the survival, development, and feeding behavior of cabbage looper and beet armyworm on two romaine lettuce cultivars, resistant 'Valmaine' and susceptible 'Tall Guzmaine'. Larval mortality of both species was significantly higher on resistant Valmaine than on susceptible Tall Guzmaine. The average weight per larva after feeding for 1 wk on Tall Guzmaine plants was 6 times (beet armyworm) and 2 times (cabbage looper) greater than that of larvae feeding on Valmaine plants. Significant reduction in larval growth on Valmaine compared with that on Tall Guzmaine resulted in a 5.9- (beet armyworm) and 2.6-d (cabbage looper) increase in larval duration and almost a 1-d increase in pupal duration. Average pupal and adult weights and successful pupation of cabbage looper and beet armyworm were reduced on Valmaine compared with Tall Guzmaine. The sex ratio of progeny did not deviate from 1:1 when larvae were reared on either Valmaine or Tall Guzmaine. The fecundity of cabbage looper and beet armyworm adults that developed from larvae reared on Valmaine was about one-third that of adults from Tall Guzmaine, but adult longevity did not significantly differ on the two lettuce cultivars. The two insect species showed different feeding preferences for leaves of different age groups on Valmaine and Tall Guzmaine. Cabbage loopers cut narrow trenches on the leaf before actual feeding to block the flow of latex to the intended site of feeding. In contrast, beet armyworms did not trench. The different feeding behavior of the two species on Valmaine may explain the superior performance of cabbage looper compared with beet armyworm.     

Is the range size of migratory birds constrained by their migratory program?

Aim Intuitively, species in which the individuals migrate long distances between summer and winter quarters should be more likely to disperse and colonize new breeding areas than resident species. However, it has repeatedly been noted that many bird species with large ranges are residents. This paradox was tested on land birds breeding in the boreal forest of the Palaearctic, the largest uninterrupted stretch of habitat on earth. Methods The longitudinal distribution of two land bird communities on each side of the Eurasian continent, in Scandinavia and eastern Siberia, were used to test whether migratory birds indeed have a lower colonization success than resident birds. Results The migratory species are significantly less likely than resident species to have a range including both regions. The pattern held true even after controlling for latitudinal effects and local abundance, and was also observed at the level of genus and family. Main conclusions The relatively low colonization success of migratory species into new breeding areas may be because these new areas require novel migratory programs (migratory distance, direction and timing) in order for the birds to reach suitable wintering grounds.     

The role of migration for spatial turnover of arctic bird species in a circumpolar perspective

Several different factors may determine where species range limits are located within regions of otherwise continuously available habitat and suitable climate. Within the Arctic tundra biome many bird species are migratory and their breeding distributions are affected by migration routes that are in turn limited by factors such as suitable winter habitat, migratory stopover sites, geographical barriers and historical routes of colonization. We identified longitudinal zones in the circumpolar Arctic of pronounced changes in the avian species composition (high species spatial turnover; ‘species divides’). We tested for the association between migratory status and the geographical location and numbers of such species divides for species with non‐breeding habitats mainly within terrestrial, pelagic and coastal ecosystems. Our results demonstrate that migration is of profound importance for both the number and locations of species divides in the Arctic. Long‐distance migration is associated with a large number of divides among terrestrial and coastal arctic birds but with a reduced number of divides among pelagic birds. We suggest that long‐distance migration permits pelagic but not terrestrial and coastal birds to colonize large winter ranges, which in turn causes expansion of breeding ranges, with more homogenous communities and reduction of species divides as consequences, among the long‐distance migrants of pelagic but not of terrestrial and coastal birds. Furthermore, the divides among long‐distance migrants are situated in two main regions, the Beringia and Greenland zones, while divides among short‐distance migrants are more evenly spaced throughout the circumpolar Arctic. The Beringia and Greenland divides result largely from inter‐continental colonization of new breeding ranges but retainment of original winter quarters in a process of evolution through extension of migration routes, leading to aggregated divides in the meeting zones of major global flyways.     

Exploring intraspecific variation in migratory destinations to investigate the drivers of migration

Various benefits (e.g. tracking of resources and of climate niche) and costs (e.g. distance travelled) are hypothesized to drive seasonal animal migrations. Until now, these potential factors have been investigated together at the species level, but migratory movements are made at the individual level, leading to intraspecific variability. Here, we use ringing/recovery data from 1308 individuals belonging to thirteen North American bird species to analyse patterns in intraspecific variability of migratory destinations in order to investigate which factors underpin bird migration and how individuals trade-off among multiple factors. Our results suggest that migratory destinations have been shaped by access to resources (most important during the breeding season) and climatic niche tracking (during winter, mostly). However, the benefits of resource surpluses and climate niche tracking appear to be traded off against the cost of migratory distance, which seems to strongly constrain where individuals migrate.