Revision of the Bicyclus ignobilis species-group (Lepidoptera: Nymphalidae: Satyrinae) with descriptions of two new species

The ignobilis-group of the genus Bicyclus Kirby 1871 is revised. The species-group contains six species with a distinct wing pattern, but limited intraspecific variation, distributed across tropical African rainforest. We investigate a set of more than 1000 specimens from a range of museum collections, including some type material, and thoroughly update the biogeographical knowledge for the group. We also describe two new species as members of the group. The included species are: Bicyclus ignobilis (Butler 1870) stat. rev., B. rileyi Condamin 1961, B. maesseni Condamin 1971, B. brakefieldi Brattström 2012, B. ottossoni sp. nov. and B. vandeweghei sp. nov. Due to observing a gradual morphological cline within B. ignobilis without any sharp transitions we suppress the previously identified subspecies B. ignobilis eurini Condamin & Fox 1963 syn. nov. and B. ignobilis acutus Condamin 1965 syn. nov

Systematics and historical biogeography of the old world butterfly subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrinae).

BACKGROUND: Butterflies of the subtribe Mycalesina have radiated successfully in almost all habitat types in Africa, Madagascar, the Indian subcontinent, Indo-China and Australasia. Studies aimed at understanding the reasons behind the evolutionary success of this spectacular Old World butterfly radiation have been hampered by the lack of a stable phylogeny for the group. Here, we have reconstructed a robust phylogenetic framework for the subtribe using 10 genes from 195 exemplar taxa. RESULTS: We recovered seven well supported clades within the subtribe corresponding to the five traditional genera (Lohora, Heteropsis, Hallelesis, Bicyclus, Mycalesis), one as recently revised (Mydosama) and one newly revised genus (Culapa). The phylogenetic relationships of these mycalesine genera have been robustly established for the first time. Within the proposed phylogenetic framework, we estimated the crown age of the subtribe to be 40 Million years ago (Mya) and inferred its ultimate origin to be in Asia. Our results reveal both vicariance and dispersal as factors responsible for the current widespread distribution of the group in the Old World tropics. We inferred that the African continent has been colonized at least twice by Asian mycalesines within the last 26 and 23 Mya. In one possible scenario, an Asian ancestor gave rise to Heteropsis on continental Africa, which later dispersed into Madagascar and most likely back colonised Asia. The second colonization of Africa by Asian ancestors resulted in Hallelesis and Bicyclus on continental Africa, the descendants of which did not colonise other regions but rather diversified only in continental Africa. The genera Lohora and Mydosama are derivatives of ancestors from continental Asia. CONCLUSION: Our proposed time-calibrated phylogeny now provides a solid framework within which we can implement mechanistic studies aimed at unravelling the ecological and evolutionary processes that culminated in the spectacular radiation of mycalesines in the Old World tropics

Conserved patterns of integrated developmental plasticity in a group of polyphenic tropical butterflies

Background: Developmental plasticity is thought to have profound macro-evolutionary effects, for example, by increasing the probability of establishment in new environments and subsequent divergence into independently evolving lineages. In contrast to plasticity optimized for individual traits, phenotypic integration, which enables a concerted response of plastic traits to environmental variability, may affect the rate of local adaptation by constraining independent responses of traits to selection. Using a comparative framework, this study explores the evolution of reaction norms for a variety of life history and morphological traits across five related species of mycalesine butterflies from the Old World tropics. Results: Our data indicate that an integrated response of a suite of key traits is shared amongst these species. Interestingly, the traits that make up the functional suite are all known to be regulated by ecdysteroid signalling in Bicyclus anynana, one of the species included in this study, suggesting the same underlying hormonal regulator may be conserved within this group of polyphenic butterflies. We also detect developmental thresholds for the expression of alternative morphs. Conclusions: The phenotypic plasticity of a broad suite of morphological and life history traits is integrated and shared among species from three geographically independent lineages of mycalesine butterflies, despite considerable periods of independent evolution and exposure to disparate environments. At the same time, we have detected examples of evolutionary change where independent traits show different patterns of reaction norms. We argue that the expression of more robust phenotypes may occur by shifting developmental thresholds beyond the boundaries of the typical environmental variation

Metacognitive Awareness : sebuah Upaya Optimalisasi Kualitas Pembelajaran Akuntansi

Pemahaman dan pembiasaan strategi metakognisi sangat dibutuhkan bagi seseorang yang sedang melakukan proses pembelajaran. Strategi ini memfasilitasi seseorang untuk mampu mengontrol pembelajarannya dalam hal ini merencanakan, memonitoring dan mengevaluasi proses belajarnya agar dapat berjalan efektif dan efisien. Individu yang sudah terbiasa menggunakan strategi ini akan menemukan cara yang tepat dalam memahami sesuatu, mengembangkan cara memecahkan masalah serta kemampuan melakukan self asesmen. Tujuan penulisan artikel ini adalah menawarkan suatu strategi yang mampu membantu pebelajar dalam mengoptimalkan kualitas proses belajarnya. Artikel ini ditulis berdasarkan hasil review literature yang relevan. Artikel ini menyimpulkan bahwa strategi metakognitif dapat dikembangkan melalui pengalaman belajar secara berkelanjutan sejak mahasiswa baru masuk perguruan tinggi sampai mereka lulus

Revision of the Bicyclus sciathis species group (Lepidoptera: Nymphalidae) with descriptions of four new species and corrected distributional records

In this paper we present a thorough revision of the sciathis species group of the butterfly genus Bicyclus (Kirby). Type materials are discussed and in several cases lectotypes are assigned to specimens from original type series. Four new, and morphologically distinct, species are described (B. elishiae Brattström sp.n., B. heathi Brattström sp.n., B. sigiussidorum Brattström sp.n. and B. subtilisurae Brattström sp.n.), along with a comprehensive molecular phylogeny that includes exemplar taxa of all currently recognized species. We also investigate the types of all previously synonymized taxa and in the process invalidate the name B. ewondo Libert. This was done after finding the previously missing holotype of B. makomensis (Strand), which clearly belongs to the same species and thereby gives the older name priority. The phylogeny showed that some distinctly different species were surprisingly closely related, suggesting a high rate of morphological evolution in parts of the sciathis group. The distributional records for the group are updated after investigating over 1700 specimens kept in a range of museum collections. Many species previously thought to be broadly sympatric were found to have much more restricted ranges, with the previous overestimations probably based on misidentified specimens. The higher level of allopatry now established will make identification of many morphologically similar species easier. The fact that species often have smaller ranges than previously known, meaning that the level of endemism for African butterflies is likely to be higher than current estimates, has important implications for conservation management. An identification key for males of all 13 currently recognized species in the species group is included

Out of the Orient: Post-Tethyan transoceanic and trans-Arabian routes fostered the spread of Baorini skippers in the Afrotropics

The origin of taxa presenting a disjunct distribution between Africa and Asia has puzzled biogeographers for more than a century. This biogeographic pattern has been hypothesized to be the result of transoceanic long-distance dispersal, Oligocene dispersal through forested corridors, Miocene dispersal through the Arabian Peninsula or passive dispersal on the rifting Indian plate. However, it has often been difficult to pinpoint the mechanisms at play. We investigate biotic exchange between the Afrotropics and the Oriental region during the Cenozoic, a period in which geological changes altered landmass connectivity. We use Baorini skippers (Lepidoptera, Hesperiidae) as a model, a widespread clade of butterflies in the Old World tropics with a disjunct distribution between the Afrotropics and the Oriental region. We use anchored phylogenomics to infer a robust evolutionary tree for Baorini skippers and estimate divergence times and ancestral ranges to test biogeographic hypotheses. Our phylogenomic tree recovers strongly supported relationships for Baorini skippers and clarifies the systematics of the tribe. Dating analyses suggest that these butterflies originated in the Oriental region, Greater Sunda Islands, and the Philippines in the early Miocene c. 23 Ma. Baorini skippers dispersed from the Oriental region towards Africa at least five times in the past 20 Ma. These butterflies colonized the Afrotropics primarily through trans-Arabian geodispersal after the closure of the Tethyan seaway in the mid-Miocene. Range expansion from the Oriental region towards the African continent probably occurred via the Gomphotherium land bridge through the Arabian Peninsula. Alternative scenarios invoking long-distance dispersal and vicariance are not supported. The Miocene climate change and biome shift from forested areas to grasslands possibly facilitated geodispersal in this clade of butterflies.Directorate for Biological Sciences. Grant Numbers: DEB‐1541500, DEB‐1541560.Peer reviewe

Miocene climate and habitat change drove diversification in Bicyclus, Africa’s largest radiation of satyrine butterflies

Compared to other regions, the drivers of diversification in Africa are poorly understood. We studied a radiation of insects with over 100 species occurring in a wide range of habitats across the Afrotropics to investigate the fundamental evolutionary processes and geological events that generate and maintain patterns of species richness on the continent. By investigating the evolutionary history of Bicyclus butterflies within a phylogenetic framework, we inferred the group’s origin at the Oligo-Miocene boundary from ancestors in the Congolian rainforests of central Africa. Abrupt climatic fluctuations during the Miocene (ca. 19-17 Ma) likely fragmented ancestral populations, resulting in at least eight early-divergent lineages. Only one of these lineages appears to have diversified during the drastic climate and biome changes of the early Miocene, radiating into the largest group of extant species. The other seven lineages diversified in forest ecosystems during the late Miocene and Pleistocene when climatic conditions were more favourable–warmer and wetter. Our results suggest changing Neogene climate, uplift of eastern African orogens, and biotic interactions might have had different effects on the various subclades of Bicyclus, producing one of the most spectacular butterfly radiations in Africa

Conserved ancestral tropical niche but different continental histories explain the latitudinal diversity gradient in brush-footed butterflies.

The global increase in species richness toward the tropics across continents and taxonomic groups, referred to as the latitudinal diversity gradient, stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluate several of these hypotheses to explain spatial diversity patterns in a butterfly family, the Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal, and also the extent to which these processes differ among regions at the same latitude. We generate a time-calibrated phylogeny containing 2,866 nymphalid species (~45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the latitudinal diversity gradient among regions because temporal diversification dynamics differ greatly across longitude. The Neotropical diversity results from low extinction rates, not high speciation rates, and biotic interchanges with other regions are rare. Southeast Asia is also characterized by a low speciation rate but, unlike the Neotropics, is the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, combined with tropical niche conservatism, played a major role in generating the modern latitudinal diversity gradient of nymphalid butterflies