Phylogenetic relationships of true butterflies (Lepidoptera: Papilionoidea) inferred from COI, 16S rRNA and EF-1α sequences

The molecular phylogenetic relationships among true butterfly families (superfamily Papilionoidea) have been a matter of substantial controversy; this debate has led to several competing hypotheses. Two of the most compelling of those hypotheses involve the relationships of (Nymphalidae + Lycaenidae) + (Pieridae + Papilionidae) and (((Nymphalidae + Lycaenidae) + Pieridae) + Papilionidae). In this study, approximately 3,500 nucleotide sequences from cytochrome oxidase subunit I (COI), 16S ribosomal RNA (16S rRNA), and elongation factor-1 alpha (EF-1α) were sequenced from 83 species belonging to four true butterfly families, along with those of three outgroup species belonging to three lepidopteran superfamilies. These sequences were subjected to phylogenetic reconstruction via Bayesian Inference (BI), Maximum Likelihood (ML), and Maximum Parsimony (MP) algorithms. The monophyletic Pieridae and monophyletic Papilionidae evidenced good recovery in all analyses, but in some analyses, the monophylies of the Lycaenidae and Nymphalidae were hampered by the inclusion of single species of the lycaenid subfamily Miletinae and the nymphalid subfamily Danainae. Excluding those singletons, all phylogenetic analyses among the four true butterfly families clearly identified the Nymphalidae as the sister to the Lycaenidae and identified this group as a sister to the Pieridae, with the Papilionidae identified as the most basal linage to the true butterfly, thus supporting the hypothesis: (Papilionidae + (Pieridae + (Nymphalidae + Lycaenidae))).     

基于线粒体ND1和16S rRNA基因序列探讨中国蝴蝶12科的系统发育关系(鳞翅目: 双孔次亚目: 蝶类)

对中国12科共32种代表蝶类的ND1基因和16S rRNA 基因进行了序列测定（包括新测30种ND1基因和9种16S rRNA基因）和比较分析, 同时采用邻接法、最大似然法和贝叶斯法构建了12科蝶类的系统发育树, 探讨了其高级分类群的系统发育关系。序列分析的结果显示: 经比对处理后的两个基因总长度为869 bp, 其中保守位点373个, 可变位点496个, 简约信息位点375个; A+T的平均含量为80.2％, 明显高于C+G的平均含量19.8％。分子系统树表明: 蛱蝶科不是单系群; 珍蝶类、斑蝶类和喙蝶类位于蛱蝶科内; 粉蝶科和凤蝶科具有共同祖先。据此建议: 绢蝶科应归入凤蝶科; 蚬蝶科归入灰蝶科; 珍蝶类、斑蝶类和喙蝶类作为蛱蝶科中的亚科, 眼蝶类从蛱蝶科中分离出来独立成科。另外, 环蝶类的系统分类地位还有待于进一步研究。     

Beak marks on the wings of butterflies and predation pressure in the field

Beak marks on butterfly wings have been used as an indicator of predation pressure. The relationship between butterflies and their predators in the field was examined to evaluate the beak mark rate as an indicator of predation pressure. Transect censuses were conducted to measure the beak mark rate on butterflies from May to November, 2010, in Shiga Prefecture, Japan. A total of 1216 butterflies of 42 species from five families were observed during the study period. The beak mark rate in the Papilionidae was significantly higher than those of all other families. Analysis of the monthly fluctuations in the beak mark rate and relative abundance of predators revealed that the higher beak mark rates observed for two butterfly families (Papilionidae and Nymphalidae) were significantly and positively related to predation pressure, while no significant relationship was found for other families (Lycaenidae, Pieridae and Hesperiidae). Beak marks in larger butterflies (Papilionidae and Nymphalidae) can be used as an indicator to evaluate the relative intensity of predation pressure in the field.     

Diet breadth and host plant diversity of tropical- vs. temperate-zone herbivores: South-East Asian and West Palaearctic butterflies as a case study

1. Data on host plant associations of butterflies (Papilionoidea, excluding Hesperiidae) from two biogeographical regions were used to investigate (1) whether tropical herbivores are more narrowly specialized with regard to host plant choice than those of northern temperate zones, and (2) whether tropical butterflies show a greater diversity of host plant affiliations. 2. There was no evidence for a more restricted diet breadth of tropical butterflies, with diet breadth being measured as number of host plant families used per species. In the families Papilionidae, Pieridae, and Nymphalidae, host plant ranges of West Palaearctic and South-East Asian species are similar, whereas in one speciose group within the Lycaenidae, the Polyommatini, tropical species are significantly more polyphagous. 3. Diet breadth also differs among higher butterfly taxa. While Papilionidae, Pieridae, the nymphalid subfamilies Satyrinae, Morphinae, Libytheinae and Apaturinae, as well as the temperate-zone Polyommatini in the Lycaenidae are composed predominantly of host specialists, the degree of polyphagy is higher among the remaining nymphalid subfamilies and in many lycaenids. These results challenge strongly the view that tropical herbivores are generally more specialized in this regard than herbivores of higher latitudes. Rather, chemical constraints and phylogenetic conservatism shape host plant associations in many taxa in such a way that differences between temperate and tropical representatives are slight. 4. Host plant diversity, measured as the number of plant families used per butterfly family and by application of the log-series model, is much higher in South-East Asian Nymphalidae and Lycaenidae (the two largest families) than in their Western Palaearctic relatives. No such differences are observed in the Papilionidae and Pieridae (the two smaller families). Besides effects of sample size, the strong association of papilionid and pierid butterflies with plants characterized by a small set of classes of secondary plant compounds might generally restrict their capability to utilize a broader taxonomic range of host plants. 5. The results indicate that high floral diversity can be reflected by higher diversity of host plant affiliations of herbivores, but taxonomic idiosyncrasies render it difficult to draw generalized conclusions.     

Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers

Phylogenetic relationships among major clades of butterflies and skippers have long been controversial, with no general consensus even today. Such lack of resolution is a substantial impediment to using the otherwise well studied butterflies as a model group in biology. Here we report the results of a combined analysis of DNA sequences from three genes and a morphological data matrix for 57 taxa (3258 characters, 1290 parsimony informative) representing all major lineages from the three putative butterfly super-families (Hedyloidea, Hesperioidea and Papilionoidea), plus out-groups representing other ditrysian Lepidoptera families. Recently, the utility of morphological data as a source of phylogenetic evidence has been debated. We present the first well supported phylogenetic hypothesis for the butterflies and skippers based on a total-evidence analysis of both traditional morphological characters and new molecular characters from three gene regions (COI, EF-1alpha and wingless). All four data partitions show substantial hidden support for the deeper nodes, which emerges only in a combined analysis in which the addition of morphological data plays a crucial role. With the exception of Nymphalidae, the traditionally recognized families are found to be strongly supported monophyletic clades with the following relationships: (Hesperiidae+(Papilionidae+(Pieridae+(Nymphalidae+(Lycaenidae+Riodinidae))))). Nymphalidae is recovered as a monophyletic clade but this clade does not have strong support. Lycaenidae and Riodinidae are sister groups with strong support and we suggest that the latter be given family rank. The position of Pieridae as the sister taxon to nymphalids, lycaenids and riodinids is supported by morphology and the EF-1alpha data but conflicted by the COI and wingless data. Hedylidae are more likely to be related to butterflies and skippers than geometrid moths and appear to be the sister group to Papilionoidea+Hesperioidea.     

Reconstructing intraordinal relationships in Lepidoptera using mitochondrial genome data with the description of two newly sequenced lycaenids, Spindasis takanonis and Protantigius superans (Lepidoptera: Lycaenidae)

Lepidoptera is one of the largest insect orders, but the phylogenetic relationships within this order, have yet to be adequately described. Among these unresolved relationships include those regarding the monophyly of the Macrolepidoptera and interfamilial relationships of the true butterflies superfamily Papilionoidea. We present two new mitochondrial genomes (mitogenomes) belonging to the butterfly family Lycaenidae to explore the phylogenetic relationships existing among lepidopteran superfamilies and true butterfly families from a mitogenome perspective, and to evaluate the characteristics of the lepidopteran mitogenomes. Our consensus phylogeny of the Lepidoptera largely supported the superfamilial relationships (((((Bombycoidea + Geometroidea) + Noctuoidea) + Pyraloidea) + Papilionoidea) + Tortricoidea), signifying a lack of support for a traditionally defined Macrolepidoptera. The familial relationships of the true butterflies concordantly recovered the previously proposed phylogenetic hypothesis (((Lycaenidae + Nymphalidae) + Pieridae) + Papilionidae). The test for the effect of optimization schemes (exclusion and inclusion of third codon position of PCGs and two rRNA genes, with and without partitions) on the resolution and relationships within the Lepidoptera have demonstrated that the majority of analyses did not substantially alter the relevant topology and node support, possibly as the result of relatively strong signal in mitogenomes for intraordinal relationships in Lepidoptera.     

Edible Lepidoptera in Mexico: Geographic distribution, ethnicity, economic and nutritional importance for rural people

In this paper, we reported the butterflies and moths that are consumed in Mexico. We identified 67 species of Lepidoptera that are eaten principally in their larval stage in 17 states of Mexico. These species belong to 16 families: Arctiidae, Bombycidae, Castniidae, Cossidae, Geometridae, Hepialidae, Hesperiidae, Lasiocampidae, Noctuidae, Nymphalidae, Papilionidae, Pieridae, Pyralidae, Saturniidae, Sesiidae, and Sphingidae. Saturniidae, Pieridae, Noctuidae and Nymphalidae were the more species consumed with 16, 11, 9, and 8 species, respectively. The genera with the largest numbers of species were: Phassus, Phoebis, Hylesia and Spodoptera, with three species. Their local distribution, corresponding to each state of Mexico, is also presented.     

秦岭国家植物园蝴蝶群落结构与多样性

为了查明秦岭国家植物园蝴蝶资源和多样性状况, 本研究选取了5种生境, 利用样线法进行了3年的蝴蝶多样性观测。结果表明: 秦岭国家植物园蝴蝶共有5科76属134种; 物种丰富度指数、物种多样性指数、均匀度指数和优势度指数分别为1.3295、3.4616、0.7068和0.7656; 其中蛱蝶科属数及物种数最多(33属65种), 凤蝶科最低(5属10种); 蛱蝶科物种多样性指数及物种丰富度指数最高, 分别为2.8525及1.3622; 凤蝶科多样性指数最低(1.4936); 粉蝶科物种丰富度指数最低(0.2790)。5种生境中, 随着植被丰富度的增加, 蝴蝶多样性指数上升。     

广东韶关市郊不同生境蝶类种群结构及多样性的初步研究

20 0 0年 4月～ 2 0 0 1年 1 0月间 ,分别在韶关市郊的西联校园区、芙蓉山公益生态林、沙湖公园、森林公园、沐溪水库区 5个点进行蝴蝶群落结构及多样性的抽样调查研究 ,共收集蝴蝶 1 388只 ,隶属于1 0科 ,35属 ,5 6种。个体数量以凤蝶科最多 ,蛱蝶科次之。种类以蛱蝶科最多 ,凤蝶科次之 ,珍蝶科最少。各样地的多样性指数和物种丰富度排序为 :沙湖公园 >校园绿化生活区 >森林公园 >沐溪水库公益生态林 >芙蓉山公益生态林。     

Butterflies (Lepidoptera, Rhopalocera) in the Arctic fauna

The Arctic fauna includes 106 species of diurnal butterflies: Papilionidae (6 species), Pieridae (20), Lycaenidae (18), Nymphalidae (30), Satyridae (27), and Hesperiidae (5). Among them, representatives of the family Nymphalidae predominate as to the features characterizing the biological progress in the Arctic, as well as to the number of the most strongly pronounced arctic forms. The family Satyridae shares the first place with Nymphalidae by the number of species, but differs from the latter in the uneven or local distribution. The family Pieridae demonstrates a wide distribution of polyzonal and boreal species in the tundra zone. The distribution patterns of Lycaenidae are different in the Eurasian and Beringian-American sectors. Species of Papilionidae and Hesperiidae occur only in the southern part of the tundra zone. Each family is characterized by specific distribution in the Arctic subzones and landscapes and by latitudinal trends in its specific ratio in the faunas. There are 30 to 40 arctic species, including arctic proper (euarctic and hemiarctic) and hypoarctic, arctoalpine, arctomontane, and arctoboreal species. The species developing successfully under high-latitude conditions are Boloria chariclea, B. polaris, B. improba, Colias nastes, C. hecla, and Erebia fasciata; the first two species can be considered true euarctic forms. Specific features of the latitudinal and longitudinal distribution of the butterfly species in different parts of the Arctic are discussed.     

阿勒泰山蝴蝶种类调查及其垂直分布的研究

对阿勒泰山南坡中国境内的蝴蝶进行了采集调查,共鉴定整理出７科、６１属、１００余种。其中凤蝶科１属１种,绢蝶科１属６种,粉蝶科８属２２种,眼蝶科１６属２３种,蛱蝶科１６属３２种,灰蝶科１２属１５种,弄蝶科７属１０种。分析了这些种类的垂直分布特点,将其划分为４个垂直分布带,即低山半荒漠带、山地灌木草地带、山地森林草原带和亚高山草原－草甸草原－高山垫状植被带。     

Genome size variation in butterflies (Insecta,Lepidotera, Papilionoidea): a thorough phylogenetic comparison

Butterflies have been of great interest to naturalists for centuries, and the study of butterflies has been an integral part of ecology and evolution ever since Darwin proposed his theory of natural selection in 1859. There are > 18 000 butterfly species worldwide, showing great diversity in morphological traits and ecological niches. Compared with butterfly diversity, however, patterns of genome size variation in butterflies remain poorly understood, especially in a phylogenetic context. Here, we sequenced and assembled the mitogenomes of 68 butterflies and measured the genome sizes (C-values) of 67 of them. We also assembled 10 mitogenomes using reads from GenBank. Among the assembled 78 mitogenomes, those from 59 species, 23 genera and one subfamily are reported for the first time. Combining with published data of mitogenomes and genome size, we explored the patterns in genome size variation for 106 butterfly species in a phylogenetic context based on analyses of mitogenomes from 264 species covering six families. Our results show that the genome size of butterflies has a 6.4-fold variation ranging from 0.203 pg (199 Mb) (Nymphalidae: Heliconius xanthocles) to 1.287 pg (1253 Mb) (Papilionidae: Parnassius orleans). Within families, the largest variation was found in Papilionidae (5.9-fold: 0.22–1.29 pg), followed by Nymphalidae (4.8-fold: 0.2–0.95 pg), Pieridae (4.4-fold: 0.22–0.97 pg), Hesperiidae (2.2-fold: 0.3–0.66 pg), Lycaenidae (2.6-fold: 0.39–1.02 pg) and Rioidinidae (1.8-fold: 0.48–0.87 pg). Our data also suggest that butterflies have an ancestral genome size of c. 0.5 pg, and some ancestral genome size increase or decrease events along different subfamilies or tribes produce the diversity of genome size variation in diverse butterflies. Our data provide novel insights into patterns of genome size variation in butterflies and are an important reference for future genome sequencing programmes.     

陕西延安市区蝶类多样性初探

延安市区有蝶类6科25属36种,其中凤蝶科2属3种;粉蝶科5属8种;眼蝶科3属5种;蛱蝶科7属10种;灰蝶科7属9种;弄蝶科1属1种。陕西省新记录5种。区系分析表明,该地区以广布种为优势种。     

宁夏六盘山国家自然保护区蝶类群落结构和多样性

对宁夏六盘山国家自然保护区蝴蝶群落的结构组成和多样性进行了研究.结果表明：六盘山保护区有蝴蝶5科68属130种,其中,蛱蝶科的属数和物种数最多,为33属68种;凤蝶科最少,为2属2种.灰蝶科的物种丰富度和均匀度指数最高,分别为1.75和0.84;蛱蝶科的多样性指数最高,为3.01;粉蝶科的优势度指数最高,为0.90;凤蝶科的物种丰富度、多样性和优势度指数均最低,分别为0.37、0.40和0.蛱蝶科中,眼蝶亚科的属数和种数最多,为13属24种;喙蝶亚科最少,仅1属1种;闪蛱蝶亚科的物种丰富度和均匀度指数最高,分别为1.11和0.84;线蛱蝶亚科的多样性指数最高,为2.04;眼蝶亚科的优势度指数最高,为0.96.     

东莞莲花山自然保护区蝴蝶群落多样性研究

记述了东莞莲花山自然保护区的蝴蝶9科36属50种;分析了不同季节蝴蝶群落的特点和在科、属、种级分类下的多样性指数[H′(GS)、H′(G)、H′G(S)]．结果表明,物种丰富度以6月份最高为34,9、4、12月则分别为18、14、2;多样性指数也是以6月份最高为2．9041,9、4、12月则分别为2．5547、2．4842、0.1085;均匀度指数以4月份最高为0.9413,9、6、12月则分别为0.8827、0.8235、0.1565：优势种以12月份最明显,报喜斑粉蝶的优势度指数高达0.9773,其次,9月份为宽边黄凤蝶(0.2571)、6月份为巴黎凤蝶(0.2263)、4月亦为巴黎凤蝶(0.2174);各科蝴蝶物种丰富度、科、属的多样性指数和均匀度指数均以蛱蝶科为最高,分别为12、2．3992、1．9231、0.9655,凤蝶科、弄蝶科次之,个体数量以斑蝶科最大,粉蝶科、凤蝶科次之,而属内种的多样性指数则以凤蝶科最高(1．5456),蛱蝶科次之(0.4761)。     

福州国家森林公园蝶类名录(Ⅱ)

本文鉴定出福州国家森林公园蝶类共计 1 0科、1 2 0属、2 0 4种 ,分载名录 ( )、( )篇。名录 ( )作为前者的续篇 ,包含 4科、6 6属、86种。名录 ( )已见本刊 1 0 ( 2 )。红蛱蝶属 Vanessa Fabricius1 1 9小红蛱蝶 V. cardui ( Linnaeus)成虫 6、9月可见。寄主 :荨麻属、水麻属、飞廉属等。1 2 0大红蛱蝶 V.indica( Herbst)成虫 2、5- 6、9- 1 2月可见。寄主 :苎麻、密花苎麻、荨麻属植物等。琉璃蛱蝶属 Kaniska Moore1 2 1琉璃蛱蝶 K.canace( Linnaeus)成虫 1 - 5、9- 1 2月可见。寄主 ;尖叶菝葜、圆锥菝葜、菝葜、马甲菝葜、穿鞘菝…     

Complete mitochondrial genome of the Teinopalpus aureus guangxiensis (Lepidoptera: Papilionidae) and related phylogenetic analyses

In this study, we sequenced the complete mitochondrial genome of Teinopalpus aureus guangxiensis (Lepidoptera: Papilionidae), which is considered as an endemic species in China. It is listed as a vulnerable species by International Union for Conservation of Nature and Natural Resources Red List and also a first class endangered species in China. The complete mtDNA from T. aureus guangxiensis was 15,235 base pairs in length and contained 13 protein-coding genes (PCGs), 2 rRNA genes, 22 tRNA genes, and a control region. The T. aureus guangxiensis genes were in the same order and orientation as the completely sequenced mitogenomes of other lepidopteran species. All PCGs of T. aureus guangxiensis mitogenome start with a typical ATN codon and terminate in the common stop codon TAA, except that ND1 gene uses TTA, ND3 gene uses ATT, and ND4 and ND4L gene use TAA. The phylogenetic relationships were reconstructed with the concatenated sequences of the 13 PCGs of the mitochondrial genome, and phylogenetic results confirmed that Nymphalidae, Lycaenidae, Papilionidae, Pieridae are monophyletic clades.     

Mud-puddling behavior in tropical butterflies: in search of proteins or minerals?

We experimentally investigated the attraction of adult butterflies to moist soil and dirt places (a behavior termed `mud-puddling') in two species-rich tropical communities on the island of Borneo. At a rain forest site, 227 individuals (46 species) were attracted to the baits, compared to 534 individuals (54 species) at a farmland site. With one single exception, all attracted butterflies were males. Of various salt and amino acid solutions, only sodium was accepted, but overall, albumin solutions turned out to be the most attractive puddling resource. Butterfly families differed consistently in their resource preferences. Representatives of the families Papilionidae and Pieridae more often visited NaCl solutions, but still accepted albumin, whereas representatives of the Nymphalidae, Hesperiidae and, in particular, Lycaenidae preferred the protein resource. In experiments using decoys prepared from pinned butterfly specimens, representatives of the Papilionidae and Pieridae were more strongly attracted to baits provided with decoys made from conspicuous, medium-sized yellow Eurema species (Pieridae), whereas dummies made from small, cryptically colored lycaenids (Prosotas and Caleta species) were ineffective. Decoys did not influence the attraction of lycaenid butterflies towards baits. Hence, visual cues play an important role in locating puddling resources for papilionids and pierids, while for lycaenid butterflies searching for nitrogen sources, olfactory cues emitted by decaying organic matter are more likely to be important. The strong attraction of male butterflies to nitrogen-rich resources suggests that, as in the case of sodium, these nutrients may increase reproductive success. Received: 5 October 1998 / Accepted: 7 December 1998     

Convergent evolution of neuroendocrine control of phenotypic plasticity in pupal colour in butterflies

Phenotypic plasticity in pupal colour occurs in three families of butterflies (the Nymphalidae, Papilionidae and Pieridae), typically in species whose pupation sites vary unpredictably in colour. In all species studied to date, larvae ready for pupation respond to environmental cues associated with the colour of their pupation sites and moult into cryptic light (yellow–green) or dark (brown–black) pupae. In nymphalids and pierids, pupal colour is controlled by a neuroendocrine factor, pupal melanization-reducing factor (PMRF), the release of which inhibits the melanization of the pupal cuticle resulting in light pupae. In contrast, the neuroendocrine factor controlling pupal colour in papilionid butterflies results in the production of brown pupae. PMRF was extracted from the ventral nerve chains of the peacock butterfly Inachis io (Nymphalidae) and black swallowtail butterfly Papilio polyxenes (Papilionidae). When injected into pre-pupae, the extracts resulted in yellow pupae in I. io but brown pupae in P. polyxenes. These results suggest that the same neuroendocrine factor controls the plasticity in pupal colour, but that plasticity in pupal colour in these species has evolved independently (convergently).     

菜粉蝶线粒体基因组的全序列测定和分析

目前关于蝶类线粒体基因组全序列及其分子进化的研究还不多见。本研究通过长PCR和引物步移法对菜粉蝶Pieris rapae Linnaeus线粒体基因组全序列进行了测定和初步分析。结果表明：菜粉蝶线粒体基因组全长15 157 bp, 包含13个蛋白编码基因、22个tRNA和2个rRNA基因以及1个非编码的控制区域, 它们的长度分别是11 196 bp, 1 474 bp, 2 093 bp和393 bp。37个基因的位置与已报道的其他蝶类基本一致, 共有10对基因间存在总共59 bp的重叠, 重叠碱基数在1~35 bp之间; 基因间隔序列共计13处120 bp, 间隔长度1~46 bp不等, 最大的基因间隔46 bp, 位于tRNA^Ile和tRNA^Gln基因之间。另外, 基于13个蛋白质编码基因的氨基酸序列, 重建了基于蛋白质编码基因序列数据的11种代表性蝶类的NJ和MP系统树。结果表明:凤蝶类(包括凤蝶和绢蝶)为一大支系, 粉蝶类、 灰蝶类与蛱蝶类(包括蛱蝶、 珍蝶)构成另一大支系。结果不支持粉蝶科与凤蝶科（包括凤蝶类和绢蝶类）构成单系群, 却显示粉蝶科、 灰蝶科和蛱蝶科的组合为单系群。