Romance of the three domains: how cladistics transformed the classification of cellular organisms

Cladistics is a biological philosophy that uses genealogical relationship among species and an inferred sequence of divergence as the basis of classification. This review critically surveys the chronological development of biological classification from Aristotle through our postgenomic era with a central focus on cladistics. In 1957, Julian Huxley coined cladogenesis to denote splitting from subspeciation. In 1960, the English translation of Willi Hennig’s 1950 work, Systematic Phylogenetics, was published, which received strong opposition from pheneticists, such as numerical taxonomists Peter Sneath and Robert Sokal, and evolutionary taxonomist, Ernst Mayr, and sparked acrimonious debates in 1960–1980. In 1977–1990, Carl Woese pioneered in using small subunit rRNA gene sequences to delimitate the three domains of cellular life and established major prokaryotic phyla. Cladistics has since dominated taxonomy. Despite being compatible with modern microbiological observations, i.e. organisms with unusual phenotypes, restricted expression of characteristics and occasionally being uncultivable, increasing recognition of pervasiveness and abundance of horizontal gene transfer has challenged relevance and validity of cladistics. The mosaic nature of eukaryotic and prokaryotic genomes was also gradually discovered. In the mid-2000s, high-throughput and whole-genome sequencing became routine and complex geneologies of organisms have led to the proposal of a reticulated web of life. While genomics only indirectly leads to understanding of functional adaptations to ecological niches, computational modeling of entire organisms is underway and the gap between genomics and phenetics may soon be bridged. Controversies are not expected to settle as taxonomic classifications shall remain subjective to serve the human scientist, not the classified.     

From Asia to Europe: The first record of the blue tiger Tirumala limniace (Cramer, [1775]) (Lepidoptera: Nymphalidae: Danainae) in the European continent

The introduction of species by human action is one of the main causes of rarefaction and extinctions of species. This fact is especially relevant in insular ecosystems. Here, we reported for the first time the arrival of Tirumala limniace (Cramer, [1775]) from the Asian to European continent. A single specimen was recorded in the Balearic Islands, which a large list of introductions around their history. Some reasons could explain the occurrence of this species into the islands, but the most likely are the cargo and nursery trade as well as the release of specimens for weddings and other social events as well as escaped from a private butterfly’s garden. This record highlights the importance of implementing early detection of introduced species in insular ecosystems.     

The relationship between mating and oogenesis in monarch butterflies (Lepidoptera: Danainae)

We studied the relationship between the timing of mating and oogenesis in monarch butterflies (Danaus plexippus) to determine (1) the potential for male nutrient input into eggs and (2) whether mating stimulates egg development. Most females mated soon after they started maturing eggs. One and 2 days after mating, females contained the same number of mature oocytes as virgin females of the same age, while 3 days after mating they contained more mature oocytes than did virgins. These results confirm the potential for male-derived nutrients to augment oocyte production, but indicate that mating is not required for oocyte maturation to occur.     

Biogeography of milkweed butterflies (Nymphalidae: Danainae) and mimetic patterns on tropical Paciflc archipelagos

Distributions of danaine butterfly species and associated mimetic patterns were compared among fifteen archipelagos of the tropical Pacific Ocean, and within five major archipelagos (the Bismarcks, Fiji, East and West Solomon Islands, and Vanuatu). Using both simple and stepwise linear regression analysis, variation in the total number of danaine species and number of mimetic patterns was assessed with respect to island size, isolation and elevation. Relative to interarchipelago distributions, the distribution of danaine species and number of mimetic patterns on islands within archipelagos exhibited less dependence upon interisland distance and island area. Geographical features influencing the number of mimetic patterns were similar to those of danaines as a whole. Analysis of residuals from stepwise linear regression suggested that factors influencing danaine distributions were different from those for non-danaine butterflies. This result is consistent with the hypothesis of enhancement of danaine species establishment through Müllerian mimicry, although other factors such as host plant availability and similar habitat use may also be important.     

Phylogeny of Polygonia, Nymphalis and related butterflies (Lepidoptera: Nymphalidae): a total-evidence analysis

We investigated the phylogeny of butterflies in the tribe Nymphalini sensu Harvey 1991, comprising the genera Vanessa, Cynthia, Bassaris, Aglais, Inachis, Nymphalis, Polygonia, Kaniska, Antanartia, Hypanartia, Symbrenthia, Mynes and Araschnia . Evidence from the mitochondrial gene ndl, the nuclear gene 'wingless' and from morphology/ ecology/behaviour were used separately and combined to analyse relationships. Phylogenies based on the different types of data agreed in many aspects of basic topology. We show that an analysis of only wing pattern characters (based on Nijhout's homology system) results in a topology broadly similar to the one resulting from analysis of the complete matrix. We found support for a monophyletic Nymphalini, where Hypanartia may be the sister clade to all other genera. Mynes, Symbrenthia and Araschnia together seem to form another basal clade. Evidence presented gives only moderate support for a monophyletic Vanessa in the wide sense, including also Cynthia and Bassaris , but strong support for the monophyly of the largely holarctic clade Aglais + Inachis + Nymphalis + Polygonia + Kaniska + Roddia . Within the latter group there is strong support for a clade consisting of Aglais + Inachis and for a second clade which includes Nymphalis, Polygonia (and its sister clade, the monotypic Kaniska) as well as Roddia l-album (= Nymphalis vaualbum ). As a consequence of this topology, Aglais is recognized as a taxon separate from Nymphalis . We present a hypothesis of species relationships within the focal group of genera. We also analyse and discuss the implications of excluding or including ecological data in phylogenetic tree construction, when the tree is to be used for studies in phylogenetic ecology.     

蛱蝶翅鳞片的超微结构观察

对我国东北地区典型常见蛱蝶科15属20种蝴蝶翅鳞片的超微结构进行了扫描电镜观察。结果显示：蛱蝶翅鳞片形态上可分为窄叶形、阔叶形和圆叶形3种,鳞片长65～135 μm,宽35～85 μm,间距48～112 μm。蛱蝶翅鳞片的超微结构可分为拱桥形、棋盘形和筛孔形3 种。拱桥形结构和棋盘形结构比较接近,二者与筛孔形结构差异较明显。在已观察的种类中,线蛱蝶属红线蛱蝶翅鳞片上的纵肋突起最小（200 nm×300 nm）,闪蛱蝶属柳紫闪蛱蝶翅鳞片上的纵肋突起最大（590 nm×560 nm）。鳞片具有相似的形状、结构和排列,尤其是同属蝴蝶翅鳞片超微结构的形状和尺寸差异较小,表明它们之间的亲缘关系接近。     

Cold resistance in all life stages of two blowfly species (Diptera, Calliphoridae)

1. Freezing was lethal in the eggs, larvae, pupae and adults of the blowflies Calliphora vicina R-D. and Calliphora vomitoria (L.), but varying degrees of supercooling were found. 2. Cold resistance (as determined by their supercooling capacity) was greatest in eggs (to c. -25 degrees C), and moderate in other life stages (range -7 to -13 degrees C), which was evident from the distributions of their supercooling points. 3. Water contents of all life stages varied from 63% to 77% of fresh weight; pupae of C. vomitoria contain the smallest proportion of water. No relationship between the amount of body water and supercooling ability was detected. 4. Glucose was the major compound in all life stages of C. vicina, with a maximum concentration of 8.22 micrograms mg-1 fresh weight in the adults. In addition, five other potential cryoprotectants occurred in concentrations greater than 0.1% of fresh weight. No correlation between solute concentration and supercooling was observed. 5. Both C. vicina and C. vomitoria have sufficient cold hardiness to avoid lethal freezing in sheltered habitats in southern Britain, which may render diapause unnecessary in both species.     

Natural mortality of currant clearwing,Synanthedon salmachus (Lepidoptera: Sesiidae), in New Zealand

Abstract

In a wide-ranging survey of 11 blackcurrant blocks, mortality of currant clearwing, Synanthedon salmachus (L.), was found to range from 1.3% to 9.7% for larvae and from 5.3% to 23.0% for pupae. At least 51% of the mortality was caused by the fungal pathogens Beauveria bassiana, Paecilomyces farinosus, and Verticillium lecanii. One unidentified predatory carabid larvae was collected, and two parasites—Xanthocryptus novozealandicus and Diadegma sp.—were reared from clearwing larvae.     

The phylogenetics and biochemistry of host-plant specialization in Melitaeine butterflies (Lepidoptera: Nymphalidae)

Butterflies in the tribe Melitaeini (Lepidoptera: Nymphalidae) are known to utilize host plants belonging to 16 families, although most host-plant records are from four families. Of the 16 host-plant families, 12 produce secondary plant metabolites called iridoids. Earlier studies have shown that larvae of several melitaeine species use iridoids as feeding stimulants and sequester these compounds for larval defense. I investigate the evolutionary history of host-plant use in the tribe Melitaeini by testing a recent phylogenetic hypothesis of 65 species representing the four major species groups of the tribe. By simple character optimization of host-plant families and presence/absence of iridoids in the host plants, I find that plant chemistry is a more conservative trait than plant taxonomy. The ancestral host plant(s) of the entire tribe most likely contained iridoids and were likely to be in the plant family Plantaginaceae. A major host shift from plants containing iridoids to plants not containing iridoids has happened three times independently. The results show that the evolution of host-plant use in melitaeines has been (and still is) a dynamic process when considering plant taxonomy, but is relatively stable when considering host-plant chemistry.     

韩国波尺蛾亚科三种幼虫记述（鳞翅目：尺蛾科）

本文详细地记述了波尺蛾亚科Eupithecia abietaria debrunneata Staudinger, E. spadix Inoue 及Telenomeuta punctimarginaria (Leech) 幼虫的形态特征.     

Phylogenetic relationships among genera of danaine butterflies (Lepidoptera: Nymphalidae) as implied by morphology and DNA sequences

Cladistic relationships among genera and subtribes of Danaini (the milkweed butterflies) were inferred by analysis of data combined from five sources: morphology of adults and immature stages, and DNA sequences from three gene regions. The results corroborate and greatly increase support for prior hypotheses based on morphology alone. A new index summarizing incongruence among data partitions, the Partition Congruence Proportion (PCP), is introduced. The significance of the inferred pattern of phylogenetic relationships for comparative chemical ecology of milkweed butterflies is briefly discussed.     

Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera: Nymphalidae)

The historical biogeography of insects in South America is largely unknown, as dated phylogenies have not been available for most groups. We have studied the phylogenetic relationships and historical biogeography of a subtribe of butterflies, Phyciodina in the family Nymphalidae, based on one mitochondrial gene (COI) and two nuclear gene regions (EF-1alpha and wingless). The subtribe comprises 89 species mainly found in tropical South America, with a few species in North America and the Greater Antilles. We find that the enigmatic genus Antillea is sister to the rest of Phyciodina, and suggest that it should be included in the subtribe. Several genera are found to be polyphyletic or nested within another genus, and are proposed to be synonymised. These are Dagon, Castilia, Telenassa and Janatella, which we propose should be synonymised with Eresia. Brazilian "Ortilia" form an independent lineage and require a new genus name. The diversification of Phyciodina has probably taken place over the past about 34 MYA. The ancestral phyciodine colonised South America from North America through a possible landspan that connected the Greater Antilles to South America about 34MYA. A vicariance event left the ancestral Antillea on the Greater Antilles, while the ancestral 0e on South America colonised the Guyanan Shield and soon after the Brazilian Shield. We hypothesise that the Brazilian Shield was an important area for the diversification of Phyciodina. From there, the ancestor of Anthanassa, Eresia and Tegosa colonised NW South America, where especially Eresia diversified in concert with the rising of the Andes beginning about 20 MYA. Central America was colonised from NW South America about 15 MYA by the ancestors of Anthanassa and Phyciodes. Our study is the first to use a dated phylogeny to study the historical biogeography of a group of South American species of butterflies.     

The phylogenetic pattern of speciation and wing pattern change in neotropical Ithomia butterflies (Lepidoptera: nymphalidae)

Species level phylogenetic hypotheses can be used to explore patterns of divergence and speciation. In the tropics, speciation is commonly attributed to either vicariance, perhaps within climate-induced forest refugia, or ecological speciation caused by niche adaptation. Mimetic butterflies have been used to identify forest refugia as well as in studies of ecological speciation, so they are ideal for discriminating between these two models. The genus Ithomia contains 24 species of warningly colored mimetic butterflies found in South and Central America, and here we use a phylogenetic hypothesis based on seven genes for 23 species to investigate speciation in this group. The history of wing color pattern evolution in the genus was reconstructed using both parsimony and likelihood. The ancestral pattern for the group was almost certainly a transparent butterfly, and there is strong evidence for convergent evolution due to mimicry. A punctuationist model of pattern evolution was a significantly better fit to the data than a gradualist model, demonstrating that pattern changes above the species level were associated with cladogenesis and supporting a model of ecological speciation driven by mimicry adaptation. However, there was only one case of sister species unambiguously differing in pattern, suggesting that some recent speciation events have occurred without pattern shifts. The pattern of geographic overlap between clades over time shows that closely related species are mostly sympatric or, in one case, parapatric. This is consistent with modes of speciation with ongoing gene flow, although rapid range changes following allopatric speciation could give a similar pattern. Patterns of lineage accumulation through time differed significantly from that expected at random, and show that most of the extant species were present by the beginning of the Pleistocene at the latest. Hence Pleistocene refugia are unlikely to have played a major role in Ithomia diversification.     

Patterns of Amazonian area relationships based on raw distributions of papilionid butterflies (Lepidoptera: Papilioninae)

Diversifications within a biota are due to several factors. Although some of these are untestable with current analytical methods, hierarchical congruence obtained with different cladistic methods and based on independent taxa are undoubtedly important. In the recent past, most hypotheses of historical biogeography (e.g. refugial, riverine, disturbance, vicariance) were tested on the Amazonian biota, selecting a number of diverse organisms such as plants, anurans, lizards, butterflies, birds and monkeys. In this study we used parsimony analysis of endemicity to infer historical relationships among 16 interfluvial areas in the Amazonian lowlands based on raw distributions of 114 Papilioninae (Lepidoptera). The analysis yielded two most parsimonious trees of area relationships. One tree was characterized by two main clusters of areas which showed a separation of Guyanan + south-east Amazonian interfluvial areas from western Amazonian interfluvial areas. The second tree showed the Guyanan interfluvial areas basal to a cluster which included all the other interfluvial areas. This latter cluster was subdivided into two main groups of areas separating the south-east Amazonian and the western Amazonian interfluvial areas. This result is discussed in the light of previous hypotheses obtained with the same method using some vertebrate taxa in the Amazonian lowlands. Likewise, comparisons with other hypotheses on lineages of birds, mammals and butterflies obtained applying cladistic biogeographical methods are made. The two alternative vicariant patterns presented for papilionid butterflies are strictly congruent with those for birds.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 345–357.     

毒蛾科幼期的鉴别

报道了毒蛾科幼期形态特征及 1 6属 5 1种毒蛾幼虫系统检索表。     

Phylogeny of flatfishes (Pleuronectiformes): comparisons and contradictions of molecular and morpho-anatomical data

Several species of flatfishes (Pleuronectiformes) have been studied by isoenzyme electrophoresis and DNA/DNA hybridization to establish the evolutionary relationships of this group of fishes. This molecular data-based phylogeny has been compared to previously described phylogenies based on morpho-anatomical criteria. A number of discrepancies are observed and discussed. The high degree of genetic divergence observed within the families of Pleuronectiformes indicates that their origin seems to be much older than suggested by their striking morphological similarity.