Evolution of swarm-founding and polygyny in South American epiponine wasps

这项研究是在1998/1999年完成的上一个项目的基础上进行的，旨在进一步推进南美洲埃皮波尼族黄蜂群体建立和永久一夫多妻制(多皇后社会制度)的进化研究。为了从遗传学的角度了解社会结构，这对理解多皇后社会系统的进化是实质性的，该项目侧重于估计给定群蜂Polybia Paulista(1)中成年和未成熟个体之间的遗传联系。为了与上述高度发达的社会制度相比较，对米氏田鼠羽化前的群体进行了社会学和生态学的观察。，他们是另一个部落Mischcyttarini的独立创始人(2)。此外，为了了解巢作为黄蜂社会生活基础的重要性，我们分析了成年黄蜂口腔分泌物中含有的化学成分，如氨基酸，以检测Polybia pau…。更多的Lista，3种，和一些其他的Polistine种(3)。口腔分泌物用来结合黄蜂从巢外觅食的巢材料。巢的另一个方面，热特性，在两个巢的保利斯塔和西洋参(4)。在这些研究的基础上，发表了两篇评论文章：(1)对圣保罗州Ribeirao Preto的圣保罗大学15个群体的等位酶和DNA分析表明，每个群体的遗传亲缘关系很低，0-0.2。因此，整个群体的成年个体和分开摘除梳子的未成熟个体实际上是无关的。这意味着许多蜂王甚至在蜂群周期的后期也在产卵。这一结果与West-Eberhard(1978)提出的“周期性寡头雌体”假说不符，必须发展另一种假说。(2)调查了两种泥蜂30多个羽化前群体的雌性个体，以了解雌性个体数量与雌性个体大小(体重)之间的关系。(3)用高效液相色谱方法分析了黄蜂与外来物质结合的口分泌物的氨基酸组成，并与已有的研究结果进行了比较。氨基酸组成在同一物种内是恒定的，在同一属的物种间非常相似，但在属间有较大的差异。这表明所产生的氨基酸的种类和相对数量反映了系统发育关系。(4)对紫背水蜂的巢温度进行了两次逐日测量，一次是在成蜂超过500只的活跃状态下进行的，另一次是在所有成蜂被移走后进行的。对2500多只成虫的活跃巢进行了一次Diel测量。巢温度，尽管放置热电偶的位置不同，但相对独立于环境温度。这可能是由于封闭式多层梳状结构具有较高的隔热能力。结果还表明，成年黄蜂实际上没有主动的体温调节。较少

This research project was carried out in succession of a previous project done in 1998/1999, and aimed to advance further the study of evolution of swarm-founding and permanent polygyny (multi-queen social system) in wasps of tribe Epiponini in South America. To know the social structure from genetic point of view, which is substantial to understand the evolution of multi-queen social systems, the project put emphasis on the estimation of genetic relatedness among adult and immature individuals in given colonies of an epiponine wasp, Polybia paulista (1). To compare with the above highly advanced social system, some sociological and ecological observations were made on pre-emergence colonies of Mischocyttarus spp. , which are independent-founders of another tribe Mischocyttarini (2). Further, to know the significance of nests as a basis for wasps' social life, chemical components, e. g. , amino acids, contained in the oral secretion secreted by adult wasps were analyzed for Polybia pau … More lista, 3 Mischocyttarusu species, and some other polistine species (3). The oral secretion is used to bind nest materials foraged by wasps from the outside of nest. Another aspect of nests, the thermal characteristics, was examined in two nests of Polybia paulista and P. occidentalis (4). On the basis of these studies, two reviews "Multi-queen social systems in eusocial insects, particularly among the Polistinae" (Y. Ito, 2002), and "Recent advances in a study of Polistinae" (S. Miyano, 2003) were published.(1) Allozyme and DNA analyzes of 15 Polybia paulista colonies taken on the campus of Sao Paulo University in Ribeirao Preto, Sao Paulo State revealed that in every colony the genetic relatedness was very low, 0-0.2. Thus, adult individuals of the whole colony and also immatures from separately removed combs were virtually non-related. This means that many queens were laying eggs even in the later phase of colony cycle. This result is not consistent with a hypothesis of "cyclical oligogyny" proposed by West-Eberhard (1978), and another hypothesis has to be developed.(2) Founding females of more than 30 pre-emergence colonies of two Miscocyttarus species were censused to know the relation between the number of founding females and the body size (weight) of adult females emerged from the first batch of brood reared by the foundresses. The distribution pattern of the number of females were also analyzed with a "Zero-truncated model".(3) Amino acid composition of the oral secretion used for binding extraneous nest materials by wasps was analyzed with a HPLC for three Mischocyttarus species and some other polistine species, Polybia paulista, Protopolybia exigua, etc. , and the results were compared with those so far studied until now. The amino acid composition was constant within the same species, and closely similar among species of the same genus, but fairly different among genera. This suggests that the kinds and relative amounts of amino acids produced reflect the phylogenetic relations.(4) Two diel measurements of nest temperature were carried out for a nest of Polybia paulista, one was done in an active state with more than 500 adult wasps and the other done after all the adults were removed. One diel measurement was done for an active nest of P. occidentalis with more than 2500 adults. Nest temperatures, though different at positions at which thermocouples were set were relatively independent from the ambient temperature. This is possibly due to high heat insulation ability of an enveloped and multi-layered-comb structure. The result also showed virtual absence of active thermoregulation by adult wasps. Less