Shun-Chern Tsaur | National Taiwan University (original) (raw)

Papers by Shun-Chern Tsaur

Research paper thumbnail of Social parasitism and behavioral interactions between two gall-forming social aphids

Insectes Sociaux, Feb 27, 2008

Research paper thumbnail of The fifth instar nymph of Lophops carinata Kirby (Homoptera: Lophopidae)

Research paper thumbnail of Some Fulgoroids (Insecta: Hemiptera) Collected on Turtle Island, Taiwan

Zoological Studies, 2005

Shun-Chern Tsaur (2005) Some fulgoroids (Insecta: Hemiptera) collected on Turtle Island, Taiwan. ... more Shun-Chern Tsaur (2005) Some fulgoroids (Insecta: Hemiptera) collected on Turtle Island, Taiwan. Zoological Studies 44(1): 1-4. This report deals with a small collection of the Fulgoroidea made using a sweep net by the author during a short visit to Turtle I. in 2003. It represents the most recent insect faunal survey of Turtle I. Among fulgoroid hemipterans, 5 species belonging to 5 genera and 5 families were collected. They are Geisha distinctissima (Walker) (Flatidae), Ugyops vittatus (Matsumura) (Delphacidae), Orthopagus splendens (Germar) (Dictyopharidae), Ricania simulans (Walker) (Ricaniidae), and Tonga westwoodi (Signoret) (Issidae). Ugyops tripunctatus (Kato), although not in the present collection, is also included in order to allow comparison with U. vittatus. Apart from a few widespread or Oriental species, the fauna showed an affinity with that of Taiwan.

Research paper thumbnail of Meenoplidae of Taiwan (Homoptera: Fulgoroidea)

Only 4 meenoplid species belonging to 3 genera, 2 subfamilies recorded in Taiwan previously. The ... more Only 4 meenoplid species belonging to 3 genera, 2 subfamilies recorded in Taiwan previously. The present paper deals with 9 species of adults and 2 species of nymphs which represent subfamilies, 5 genera. (1) The subfamily Kermesiinae includes 3 genera, 3 new species, 2 known species, 1 newly recorded, and 2 species of nymphal stages. (2) Meenoplinae contains 2 genera, 1 new species, 2 known species. Among the total amount of 9 species, 2 genera and 4 species are new to science, 1 species is newly recorded in the fauna of Taiwan, and 4 species are known. Keys are given to the subfamilies, the genera and the species of adults. New genus : Metanigrus Tsaur and Yang. New species : Eponisia macula Tsaur and Yang, Eponisia woodwardi Tsaur and Yang, Eponisiel/a matsumurai Tsaur and Yang. New synonymy: Anigrus nigricans (Matsumuras) and A. metalces Fennah are reduced to Syns. of A. frequens (Matsumura). Nisia australiensis Woodward and N. carolinensis Fennah are new status .

Research paper thumbnail of A Rapidly Evolving Homeobox at the Site of a Hybrid Sterility Gene

Science, Nov 20, 1998

condensed, evenly staining chromosomes were scored as mitotic. At least 300 cells were counted fo... more condensed, evenly staining chromosomes were scored as mitotic. At least 300 cells were counted for each determination. 33. Genomic DNA was purified from cell lysates with the QiaAMP spin blood kit (Qiagen) and used as a substrate for polymerase chain reaction and Southern blot assessment of targeting vector integration. 34. Equal numbers of cells were collected, lysed in Laemmli sample buffer, and subjected t o electrophoresis and protein immunoblotting. Filters were probed with antibodies t o p53 (pAb 1801) and p 2 l (EA10 [W. S. El-Deiry et al., Cancer Res. 55, 2910 (1995)l). Signals were visualized with enhanced chemilunescence (Pierce). 35. Modified from K. Kaufmann et al., Cell Growth. Diff. 8, 1105 (1997). In brief, extracts for in vitro kinase assays were prepared by lysis of washed, centrifuged cells in 50 m M tris-HCI (pH 7.5), 0.5% Nonidet P-40, 10% (vlv) glycerol, 100 m M sodium chloride, 10 m M sodium orthophosphate, 5 m M P-glycerophosphate, 50 m M sodium fluoride, 0.3 m M sodium orthovanadate, 1 m M dithiothreitol, and 1 X complete protease inhibitor cocktail (Boehringer Manheim) for 30 min at 4°C. Kinase complexes were immunoprecipitated by adding monoclonal antibody t o cyclin B1 (150 ng, Santa Cruz) and protein A-Sepharose (Life Technologies). Immune complexes were washed with lysis buffer and incubated in 25 ~1 of a solution containing 20 m M tris-HCI (pH 7.5), 7.5 m M magnesium chloride,

Research paper thumbnail of Revision of the subgenus Stegana (Steganina) from China, with assessment of species delimitation using DNA barcodes (Diptera, Drosophilidae)

Zootaxa, Mar 7, 2023

A total of 58 (eight known and 50 new) species of the subgenus Stegana (Steganina) from China wer... more A total of 58 (eight known and 50 new) species of the subgenus Stegana (Steganina) from China were surveyed and (re)described: S. (S.) bacilla Chen & Aotsuka, 2004, S. (S.) belokobylskiji Sidorenko, 1997, S. (S.) hirticeps Wang, Gao, & Chen, 2013, S. (S.) izu Sidorenko, 1997, S. (S.) kanmiyai Okada & Sidorenko, 1992, S. (S.) masanoritodai Okada & Sidorenko, 1992, S. (S.) maymyo Sidorenko, 1997, stat. rev., S. (S.) nigripes Zhang & Chen, 2015, S. (S.) alafoliacea Zhang & Chen, sp. nov., S. (S.) baoxing Li & Chen, sp. nov., S. (S.) bibarbata Li & Chen, sp. nov., S. (S.) bimai Cui & Chen, sp. nov., S. (S.) cinereipecta Zhang & Chen, sp. nov., S. (S.) cardua Cui & Chen, sp. nov., S. (S.) cordhirsuta Wang & Chen, sp. nov., S. (S.) cornuta Li & Chen, sp. nov., S. (S.) cucullata Li & Chen, sp. nov., S. (S.) cultella Cui & Chen, sp. nov., S. (S.) curvitabulata Cui & Chen, sp. nov., S. (S.) daiya Cui & Chen, sp. nov., S. (S.) dendrophila Zhang & Chen, sp. nov., S. (S.) flabella Li & Chen, sp. nov., S. (S.) flavipes Li & Chen, sp. nov., S. (S.) formosa Zhang & Chen, sp. nov., S. (S.) fusca Li & Chen, sp. nov., S. (S.) fuscipes Li & Chen, sp. nov., S. (S.) glaucopalpula Cui & Chen, sp. nov., S. (S.) haba Zhang & Chen, sp. nov., S. (S.) hirticlavata Cui & Chen, sp. nov., S. (S.) iaspidea Zhang & Chen, sp. nov., S. (S.) idiasta Cui & Chen, sp. nov., S. (S.) kanda Cui & Chen, sp. nov., S. (S.) labao Li & Chen, sp. nov., S. (S.) lancang Li & Chen, sp. nov., S. (S.) latifoliacea Wang & Chen, sp. nov., S. (S.) liusanjieae Li & Chen, sp. nov., S. (S.) magniflava Cui & Chen, sp. nov., S. (S.) mailangang Li & Chen, sp. nov., S. (S.) marenubila Cui & Chen, sp. nov., S. (S.) menghai Zhang & Chen, sp. nov., S. (S.) menglian Li & Chen, sp. nov., S. (S.) minutiflava Li & Chen, sp. nov., S. (S.) multiprocera Li & Chen, sp. nov., S. (S.) nayun Li & Chen, sp. nov., S. (S.) nigridentata Wang & Chen, sp. nov., S. (S.) nigripalpula Cui & Chen, sp. nov., S. (S.) otphylla Cui & Chen, sp. nov., S. (S.) radiciflava Zhang & Chen, sp. nov., S. (S.) rava Cui & Chen, sp. nov., S. (S.) sciophila Li & Chen, sp. nov., S. (S.) septencolorata Li & Chen, sp. nov., S. (S.) serrata Zhang & Chen, sp. nov., S. (S.) silvestrella Zhang & Chen, sp. nov., S. (S.) simola Cui & Chen, sp. nov., S. (S.) yani Li & Chen, sp. nov., S. (S.) yixiang Zhang & Chen, sp. nov., S. (S.) zaduo Cui & Chen, sp. nov., and S. (S.) zhuoma Cui & Chen, sp. nov. We also provided a complete list of Chinese Steganina species together with their geographical distributions. In addition, the majority of currently available DNA barcode (partial sequence of the mitochondrial cytochrome c oxidase subunit I (COI) gene) sequences of this subgenus (435 sequences of 102 spp.) were employed in a molecular analysis for species delimitation. Taken together, morphology- and molecular-based species delimitation results reached a consensus for an overwhelming majority of these Steganina species (98 of 102 spp.).

Research paper thumbnail of <strong>The rediscovery of the holotype of <em>Kotonisia kanoi </em>Matsumura, 1938 with notes on Matsumura's type specimens of Fulgoroidea (Insecta: Hemiptera: Fulgoromorpha)</strong>

Zootaxa, Dec 21, 2009

Shonen Matsumura (1872–1960), the founder of entomology in Japan, is no doubt among the most infl... more Shonen Matsumura (1872–1960), the founder of entomology in Japan, is no doubt among the most influential and prolific entomologists. He produced a series of works, illustrated lists of the insects and described as many as 1200 new species. Unfortunately, most of the types he named and collected from Taiwan were brought to Japan in the late 1940s, and were hard to access by foreign students until relatively recently.Matsumura&amp;#39;s type depositions contain brief descriptions and, sometimes, only female specimens were available for his types. This prevented sound revisionary work in Taiwan. In addition, Matsumura did not designate holotypes in his early works but simply indicated how many specimens he had.

Research paper thumbnail of Expression Profile and Gene Age Jointly Shaped the Genome-Wide Distribution of Premature Termination Codons in a Drosophila melanogaster Population

Molecular Biology and Evolution, Nov 3, 2014

Widespread premature termination codon mutations (PTCs) were recently observed in human and fly p... more Widespread premature termination codon mutations (PTCs) were recently observed in human and fly populations. We took advantage of the population resequencing data in the Drosophila Genetic Reference Panel to investigate how the expression profile and the evolutionary age of genes shaped the allele frequency distribution of PTCs. After generating a high-quality data set of PTCs, we clustered genes harboring PTCs into three categories: genes encoding low-frequency PTCs (1.5%), moderate-frequency PTCs (1.5-10%), and high-frequency PTCs (410%). All three groups show narrow transcription compared with PTC-free genes, with the moderate-and high-PTC frequency groups showing a pronounced pattern. Moreover, nearly half (42%) of the PTC-encoding genes are not expressed in any tissue. Interestingly, the moderate-frequency PTC group is strongly enriched for genes expressed in midgut, whereas genes harboring high-frequency PTCs tend to have sex-specific expression. We further find that although young genes born in the last 60 My compose a mere 9% of the genome, they represent 16%, 30%, and 50% of the genes containing low-, moderate-, and highfrequency PTCs, respectively. Among DNA-based and RNA-based duplicated genes, the child copy is approximately twice as likely to contain PTCs as the parent copy, whereas young de novo genes are as likely to encode PTCs as DNA-based duplicated new genes. Based on these results, we conclude that expression profile and gene age jointly shaped the landscape of PTC-mediated gene loss. Therefore, we propose that new genes may need a long time to become stably maintained after the origination.

Research paper thumbnail of The rediscovery of the holotype of Kotonisia kanoi Matsumura, 1938 with notes on Matsumura's type specimens of Fulgoroidea (Insecta: Hemiptera: Fulgoromorpha)

Zenodo (CERN European Organization for Nuclear Research), Dec 31, 2009

Tsaur, Shun-Chern (2009): The rediscovery of the holotype of Kotonisia kanoi Matsumura, 1938 with... more Tsaur, Shun-Chern (2009): The rediscovery of the holotype of Kotonisia kanoi Matsumura, 1938 with notes on Matsumura's type specimens of Fulgoroidea (Insecta: Hemiptera: Fulgoromorpha). Zootaxa 2315: 66-68, DOI: 10.5281/zenodo.275420

Research paper thumbnail of Cryptic diversity in the subgenus Oxyphortica (Diptera, Drosophilidae, Stegana)

PeerJ, Oct 29, 2021

Phylogenetic relationships of the subgenus Oxyphortica were reconstructed based on two mitochondr... more Phylogenetic relationships of the subgenus Oxyphortica were reconstructed based on two mitochondrial genes (COI and ND2). The results revealed the paraphyly of Oxyphortica and supported high levels of cryptic diversity within this subgenus. By integrating morphological characteristics and molecular evidence, we identified 17 new species as members of Oxyphortica: S. (O.

Research paper thumbnail of Positive selection driving the evolution of a gene of male reproduction, Acp26Aa, of Drosophila: II. Divergence versus polymorphism

Molecular Biology and Evolution, Aug 1, 1998

The evolution of the gene for a male ejaculatory protein, Acp26Aa, has been shown to be driven by... more The evolution of the gene for a male ejaculatory protein, Acp26Aa, has been shown to be driven by positive selection when nonsibling species in the Drosophila melanogaster subgroup are compared. To know if selection has been operating in the recent past and to understand the details of its dynamics, we obtained DNA sequences of Acp26Aa and the nearby Acp26Ab gene from 39 D. melanogaster chromosomes. Together with the 10 published sequences, we analyzed 49 sequences from five populations in four continents. The southern African population is somewhat differentiated from all other populations, but its nucleotide diversity is lower at these two loci. We find the following results for Acp26Aa: (1) The R : S (replacement : silent changes) ratio is significantly higher in the between-species comparisons than in the within-species data by the McDonald and Kreitman test. Positive selection is probably responsible for the excess of amino acid replacements between species. (2) However, within-species nucleotide diversity is high. Neither the Tajima test nor the Fu and Li test indicates a reduction in nucleotide diversity due to positive selection in the recent past. (3) The newly derived nucleotides in D. melanogaster are at high frequency significantly more often than predicted by the neutral equilibrium. Since the nearby Acp26Ab gene does not show these patterns, these observations cannot be attributed to the characteristics of this chromosomal region. We suggest that positive selection is active, but may be weak, for each amino acid change in the Acp26Aa gene.

Research paper thumbnail of The nucleotide changes governing cuticular hydrocarbon variation and their evolution in <i>Drosophila melanogaster</i>

Proceedings of the National Academy of Sciences of the United States of America, Mar 20, 2001

The cuticular hydrocarbon (CH) pheromones in Drosophila melanogaster exhibit strong geographic va... more The cuticular hydrocarbon (CH) pheromones in Drosophila melanogaster exhibit strong geographic variation. African and Caribbean populations have a high ratio of 5,9 heptacosadiene͞ 7,11 heptacosadiene (the ''High'' CH type), whereas populations from all other areas have a low ratio (''Low'' CH type). Based on previous genetic mapping, DNA markers were developed that localized the genetic basis of this CH polymorphism to within a 13-kb region. We then carried out a hierarchical search for diagnostic nucleotide sites starting with four lines, and increasing to 24 and 43 lines from a worldwide collection. Within the 13-kb region, only one variable site shows a complete concordance with the CH phenotype. This is a 16-bp deletion in the 5 region of a desaturase gene (desat2) that was recently suggested to be responsible for the CH polymorphism on the basis of its expression [

Research paper thumbnail of A new species of Cixius from the United States (Homoptera: Fulgoroidea: Cixiidae)

Research paper thumbnail of The Cixiidae of Taiwan, Part VII: Tribe Pintaliini (Hemiptera: Fulgoroidea)

Zoological Studies, 2003

Shun-Chern Tsaur and Tung-Ching Hsu (2003) The Cixiidae of Taiwan, Part VII: tribe Pintaliini (He... more Shun-Chern Tsaur and Tung-Ching Hsu (2003) The Cixiidae of Taiwan, Part VII: tribe Pintaliini (Hemiptera: Fulgoroidea). Zoological Studies 42(3): 431-443. This study completes the first taxonomic revision of the planthopper family Cixiidae in Taiwan with emphasis on the morphology of the male and female genitalia. In total, 20 genera and 155 species are recognized. A key to the tribes of Bothriocerinae was given in a previous work. In this paper, keys to both the subfamilies and tribes of the Cixiidae are presented. The 5 tribes Pentastirini, Bothriocerini, Bennini, Stenophlepsini, and Cixiini were previously revised by the authors, and the remaining tribe, Pintaliini, is treated here with 6 species. Neocarpia maai represents a new genus and new species. Kirbyana lini, Eucarpia stellata, E. specialis, and E. truncata are described as new species. The male of K. pagana (Melichar) is described and illustrated for the first time.

Research paper thumbnail of <p><strong>Revision of the subgenus <em>Phortica</em> (sensu stricto) (Diptera, Drosophilidae) from East Asia, with assessment of species delimitation using DNA barcodes</strong></p&gt

Zootaxa, Sep 30, 2019

A total of 50 (43 known and seven new) species in the subgenus Phortica (sensu stricto) were surv... more A total of 50 (43 known and seven new) species in the subgenus Phortica (sensu stricto) were surveyed and (re)described from China: P. bicornuta (Chen &amp; Toda, 1997); P. bipartita (Toda &amp; Peng, 1992); P. biprotrusa (Chen &amp; Toda, 1998); P. cardua (Okada, 1977); P. chi (Toda &amp; Sidorenko, 1996); P. conifera (Okada, 1977); P. eparmata (Okada, 1977); P. eugamma (Toda &amp; Peng, 1990); P. excrescentiosa (Toda &amp; Peng, 1990); P. fangae (Máca, 1993); P. flexuosa (Zhang &amp; Gan, 1986); P. foliata (Chen &amp; Toda, 1997); P. gamma (Toda &amp; Peng, 1990); P. gigas (Okada, 1977); P. glabtabula Chen &amp; Gao, 2005; P. hainanensis (Chen &amp; Toda, 1998); P. hongae (Máca, 1993); P. huazhii Cheng &amp; Chen, 2008; P. iota (Toda &amp; Sidorenko, 1996); P. jadete Zhu, Cao &amp; Chen, 2018; P. kappa (Máca, 1977); P. lambda (Toda &amp; Peng, 1990); P. latifoliacea Chen &amp; Watabe, 2008; P. magna (Okada, 1960); P. okadai (Máca, 1977); P. omega (Okada, 1977); P. orientalis (Hendel, 1914); P. pangi Chen &amp; Wen, 2005; P. paramagna (Okada, 1971); P. perforcipata (Máca &amp; Lin, 1993); P. pi (Toda &amp; Peng, 1990); P. protrusa (Zhang &amp; Shi, 1997); P. pseudopi (Toda &amp; Peng, 1990); P. pseudotau (Toda &amp; Peng, 1990); P. psi (Zhang &amp; Gan, 1986); P. rhagolobos Chen &amp; Gao, 2008; P. saeta (Zhang &amp; Gan, 1986); P. setitabula Chen &amp; Gao, 2005; P. subradiata (Okada, 1977); P. tau (Toda &amp; Peng, 1990); P. uncinata Chen &amp; Gao, 2005; P. unipetala Chen &amp; Wen, 2005; P. allomega Gong &amp; Chen, sp. nov.; P. archikappa Gong &amp; Chen, sp. nov.; P. dianzangensis Gong &amp; Chen, sp. nov.; P. imbacilia Gong &amp; Chen, sp. nov.; P. liukuni Gong &amp; Chen, sp. nov.; P. tibeta Gong &amp; Chen, sp. nov.; and P. xianfui Gong &amp; Chen, sp. nov. In addition, seven new synonyms were recognized: P. acongruens (Zhang &amp; Shi, 1997), syn. nov.; P. antillaria (Chen &amp; Toda, 1997), syn. nov.; P. kukuanensis Máca, 2003, syn. nov.; P. linae (Máca &amp; Chen, 1993), syn. nov.; P. shillongensis (Singh &amp; Gupta, 1979), syn. nov.; P. takadai (Okada, 1977), syn. nov.; and P. watanabei (Máca &amp; Lin, 1993), syn. nov. A key to all Asian species (except for the eparmata species complex) of this subgenus was provided. All currently available DNA barcode (partial mitochondrial cytochrome c oxidase subunit I (COI) gene) sequences of this subgenus (217 sequences of 54 species) are employed in a molecular analysis using different species delimitation methods. The results indicate that approximately 68.5% (37 of 54 spp.) of Phortica (s. str.) species could be clearly distinguished from closely related morphospecies or cryptic species.

Research paper thumbnail of Survey of the genus Stegana Meigen (Diptera, Drosophilidae) from Taiwan, with DNA barcodes and descriptions of three new species

Zoological Studies, Jan 15, 2014

Background: Twelve Stegana species have been reported from Taiwan, yet only four were also record... more Background: Twelve Stegana species have been reported from Taiwan, yet only four were also recorded from Mainland China. This may not reflect the actual fauna between both sides of the strait. This report mainly deals with a fly collection of the genus Stegana during a short visit to Taiwan in 2012. It represents the most recent drosophilid faunal survey of Taiwan associating with bleeding trees. Results: In this study, 17 species were recognized including three new ones and eight new records. They are Stegana (Oxyphortica) convergens (de Meijere, 1911); Stegana (Oxyphortica) nigripennis (Hendel, 1914); Stegana

Research paper thumbnail of Sex in Drosophila mauritiana: A Very High Level of Amino Acid Polymorphism in a Male Reproductive Protein Gene, Acp26Aa

Molecular Biology and Evolution, 2001

Many genes pertaining to male reproductive functions have been shown to evolve rapidly between sp... more Many genes pertaining to male reproductive functions have been shown to evolve rapidly between species, and evidence increasingly suggest the influence of positive Darwinian selection. The accessory gland protein gene (Acp26Aa) of Drosophila is one such example. In order to understand the mechanism of selection, it is often helpful to examine the pattern of polymorphism. We report here that the level of amino acid polymorphism in the Nterminal quarter of Acp26Aa is high in Drosophila melanogaster and is unprecedented in its sibling species Drosophila mauritiana. We postulate that (1) this N-terminal segment may play a role in sperm competition, and (2) D. mauritiana may have been under much more intense sexual selection than other species. Both postulates have important ramifications and deserve to be tested rigorously.

Research paper thumbnail of Gene duplication and speciation in <i>Drosophila</i> : Evidence from the <i>Odysseus</i> locus

Proceedings of the National Academy of Sciences of the United States of America, Aug 10, 2004

The importance of gene duplication in evolution has long been recognized. Because duplicated gene... more The importance of gene duplication in evolution has long been recognized. Because duplicated genes are prone to diverge in function, gene duplication could plausibly play a role in species differentiation. However, experimental evidence linking gene duplication with speciation is scarce. Here, we show that a hybridmale sterility gene, Odysseus (OdsH), arose by gene duplication in the Drosophila genome. OdsH has evolved at a very high rate, whereas its most immediate paralog, unc-4, is nearly identical among species in the Drosophila melanogaster subgroup. The disparity in their sequence evolution is echoed by the divergence in their expression patterns in both soma and reproductive tissues. We suggest that duplicated genes that have yet to evolve a stable function at the time of speciation may be candidates for ''speciation genes,'' which is broadly defined as genes that contribute to differential adaptation between species.

Research paper thumbnail of The phylogeny of closely related species as revealed by the genealogy of a speciation gene, <i>Odysseus</i>

Proceedings of the National Academy of Sciences of the United States of America, Apr 25, 2000

Molecular differentiation between races or closely related species is often incongruent with the ... more Molecular differentiation between races or closely related species is often incongruent with the reproductive divergence of the taxa of interest. Shared ancient polymorphism and͞or introgression during secondary contact may be responsible for the incongruence. At loci contributing to speciation, these two complications should be minimized (1, 2); hence, their variation may more faithfully reflect the history of the species' reproductive differentiation. In this study, we analyzed DNA polymorphism at the Odysseus (OdsH) locus of hybrid sterility between Drosophila mauritiana and Drosophila simulans and were able to verify such a prediction. Interestingly, DNA variation only a short distance away (1.8 kb) appears not to be influenced by the forces that shape the recent evolution of the OdsH coding region. This locus thus may represent a test case of inferring phylogeny of very closely related species. S pecies are delineated by shared reproductive physiology, development, sexual behavior, and morphology (3, 4). Divergence in these systems is manifested as hybrid sterility, hybrid inviability, premating isolation, and morphological differences, respectively. Races are less well defined but members often may cluster by morphological traits. One of the paradoxes concerning race or species differentiation is the common occurrences of ambiguity in distinguishing taxa by molecular means, even when grouping by reproductive or morphological traits is straightforward and clearcut. Human racial differentiation may be a most obvious example in which many morphological characters cluster by geographical origin, whereas almost all molecular polymorphisms are extensively shared among races (5). Morphological distinction among dog breeds is another example (6). In Drosophila, sexual isolation between the Zimbabwe and non-African races of Drosophila melanogaster is clearly determined by many genes spread over the autosomal genome (7), and yet, recent molecular data have failed to show much differentiation at autosomal loci (8, 9). An explanation for the discordance between the ''reproductive'' and ''molecular'' phylogeny is that genomes may be mosaics with respect to molecular genealogy, as illustrated in Fig. 1. Most loci, chosen without regard to their roles in reproductive differentiation, may not reflect the biological divergence in their sequence polymorphism because of either shared ancient polymorphism or gene introgression through secondary contact (Fig. 1b). Ancient polymorphism may persist until present day in species with large population sizes (10, 11), and gene introgression, even at a very low level, may be sufficient to obliterate differentiation (12). In this context, we shall consider separately ''speciation genes,'' defined as loci that contribute directly to some aspects of biological divergence between closely related species (such as gametogenesis, behavior, or morphology). A hypothesis, proposed in various forms (1, 2, 13, 14), is that ''speciation genes'' may record a phylogenetic history more consistent with species' reproductive biology. This is because polymorphism and divergence at these loci should be relatively unaffected by shared polymorphisms or introgressions (see the legend of Fig. 1a). The cloning of the Odysseus (OdsH, H for homeodomain) locus of hybrid male sterility in the Drosophila simulans clade (15) therefore provides an opportunity to test this hypothesis. The sibling species of D. simulans, Drosophila mau-This paper was submitted directly (Track II) to the PNAS office. Data deposition: The sequences reported in this paper have been deposited in the GenBank database (accession nos. AF254805 for D. simulans, AF254806 for D. mauritiana, and AF254807 for D. sechellia).

Research paper thumbnail of Two New Species of Oliarus from Taiwan (Homoptera : Fulgoroidea : Cixiidae), with Proposition and Discussion on 0. Horjshanus Group 1

~.. . . / . Shun-Chern Tsaur (1990) Two new species of Oliarus from Taiwan (Homo­ ptera : Fulgoro... more ~.. . . / . Shun-Chern Tsaur (1990) Two new species of Oliarus from Taiwan (Homo­ ptera : Fulgoroidea : Cixiidae), with proposition and discussion on 0. horishanus group. Bull. Inst. Zoo!., Academia Sinica 29(3): 135-139. Two new species of Oliarus ( 0. hsui and 0. shiaoi) from Taiwan have been added. The 0 . horishanus group proposed here includes five species; 0. horishanus, 0. hsui, 0. nigroner\latus, 0. shiaoi and 0 . yangi.

Research paper thumbnail of Social parasitism and behavioral interactions between two gall-forming social aphids

Insectes Sociaux, Feb 27, 2008

Research paper thumbnail of The fifth instar nymph of Lophops carinata Kirby (Homoptera: Lophopidae)

Research paper thumbnail of Some Fulgoroids (Insecta: Hemiptera) Collected on Turtle Island, Taiwan

Zoological Studies, 2005

Shun-Chern Tsaur (2005) Some fulgoroids (Insecta: Hemiptera) collected on Turtle Island, Taiwan. ... more Shun-Chern Tsaur (2005) Some fulgoroids (Insecta: Hemiptera) collected on Turtle Island, Taiwan. Zoological Studies 44(1): 1-4. This report deals with a small collection of the Fulgoroidea made using a sweep net by the author during a short visit to Turtle I. in 2003. It represents the most recent insect faunal survey of Turtle I. Among fulgoroid hemipterans, 5 species belonging to 5 genera and 5 families were collected. They are Geisha distinctissima (Walker) (Flatidae), Ugyops vittatus (Matsumura) (Delphacidae), Orthopagus splendens (Germar) (Dictyopharidae), Ricania simulans (Walker) (Ricaniidae), and Tonga westwoodi (Signoret) (Issidae). Ugyops tripunctatus (Kato), although not in the present collection, is also included in order to allow comparison with U. vittatus. Apart from a few widespread or Oriental species, the fauna showed an affinity with that of Taiwan.

Research paper thumbnail of Meenoplidae of Taiwan (Homoptera: Fulgoroidea)

Only 4 meenoplid species belonging to 3 genera, 2 subfamilies recorded in Taiwan previously. The ... more Only 4 meenoplid species belonging to 3 genera, 2 subfamilies recorded in Taiwan previously. The present paper deals with 9 species of adults and 2 species of nymphs which represent subfamilies, 5 genera. (1) The subfamily Kermesiinae includes 3 genera, 3 new species, 2 known species, 1 newly recorded, and 2 species of nymphal stages. (2) Meenoplinae contains 2 genera, 1 new species, 2 known species. Among the total amount of 9 species, 2 genera and 4 species are new to science, 1 species is newly recorded in the fauna of Taiwan, and 4 species are known. Keys are given to the subfamilies, the genera and the species of adults. New genus : Metanigrus Tsaur and Yang. New species : Eponisia macula Tsaur and Yang, Eponisia woodwardi Tsaur and Yang, Eponisiel/a matsumurai Tsaur and Yang. New synonymy: Anigrus nigricans (Matsumuras) and A. metalces Fennah are reduced to Syns. of A. frequens (Matsumura). Nisia australiensis Woodward and N. carolinensis Fennah are new status .

Research paper thumbnail of A Rapidly Evolving Homeobox at the Site of a Hybrid Sterility Gene

Science, Nov 20, 1998

condensed, evenly staining chromosomes were scored as mitotic. At least 300 cells were counted fo... more condensed, evenly staining chromosomes were scored as mitotic. At least 300 cells were counted for each determination. 33. Genomic DNA was purified from cell lysates with the QiaAMP spin blood kit (Qiagen) and used as a substrate for polymerase chain reaction and Southern blot assessment of targeting vector integration. 34. Equal numbers of cells were collected, lysed in Laemmli sample buffer, and subjected t o electrophoresis and protein immunoblotting. Filters were probed with antibodies t o p53 (pAb 1801) and p 2 l (EA10 [W. S. El-Deiry et al., Cancer Res. 55, 2910 (1995)l). Signals were visualized with enhanced chemilunescence (Pierce). 35. Modified from K. Kaufmann et al., Cell Growth. Diff. 8, 1105 (1997). In brief, extracts for in vitro kinase assays were prepared by lysis of washed, centrifuged cells in 50 m M tris-HCI (pH 7.5), 0.5% Nonidet P-40, 10% (vlv) glycerol, 100 m M sodium chloride, 10 m M sodium orthophosphate, 5 m M P-glycerophosphate, 50 m M sodium fluoride, 0.3 m M sodium orthovanadate, 1 m M dithiothreitol, and 1 X complete protease inhibitor cocktail (Boehringer Manheim) for 30 min at 4°C. Kinase complexes were immunoprecipitated by adding monoclonal antibody t o cyclin B1 (150 ng, Santa Cruz) and protein A-Sepharose (Life Technologies). Immune complexes were washed with lysis buffer and incubated in 25 ~1 of a solution containing 20 m M tris-HCI (pH 7.5), 7.5 m M magnesium chloride,

Research paper thumbnail of Revision of the subgenus Stegana (Steganina) from China, with assessment of species delimitation using DNA barcodes (Diptera, Drosophilidae)

Zootaxa, Mar 7, 2023

A total of 58 (eight known and 50 new) species of the subgenus Stegana (Steganina) from China wer... more A total of 58 (eight known and 50 new) species of the subgenus Stegana (Steganina) from China were surveyed and (re)described: S. (S.) bacilla Chen &amp; Aotsuka, 2004, S. (S.) belokobylskiji Sidorenko, 1997, S. (S.) hirticeps Wang, Gao, &amp; Chen, 2013, S. (S.) izu Sidorenko, 1997, S. (S.) kanmiyai Okada &amp; Sidorenko, 1992, S. (S.) masanoritodai Okada &amp; Sidorenko, 1992, S. (S.) maymyo Sidorenko, 1997, stat. rev., S. (S.) nigripes Zhang &amp; Chen, 2015, S. (S.) alafoliacea Zhang &amp; Chen, sp. nov., S. (S.) baoxing Li &amp; Chen, sp. nov., S. (S.) bibarbata Li &amp; Chen, sp. nov., S. (S.) bimai Cui &amp; Chen, sp. nov., S. (S.) cinereipecta Zhang &amp; Chen, sp. nov., S. (S.) cardua Cui &amp; Chen, sp. nov., S. (S.) cordhirsuta Wang &amp; Chen, sp. nov., S. (S.) cornuta Li &amp; Chen, sp. nov., S. (S.) cucullata Li &amp; Chen, sp. nov., S. (S.) cultella Cui &amp; Chen, sp. nov., S. (S.) curvitabulata Cui &amp; Chen, sp. nov., S. (S.) daiya Cui &amp; Chen, sp. nov., S. (S.) dendrophila Zhang &amp; Chen, sp. nov., S. (S.) flabella Li &amp; Chen, sp. nov., S. (S.) flavipes Li &amp; Chen, sp. nov., S. (S.) formosa Zhang &amp; Chen, sp. nov., S. (S.) fusca Li &amp; Chen, sp. nov., S. (S.) fuscipes Li &amp; Chen, sp. nov., S. (S.) glaucopalpula Cui &amp; Chen, sp. nov., S. (S.) haba Zhang &amp; Chen, sp. nov., S. (S.) hirticlavata Cui &amp; Chen, sp. nov., S. (S.) iaspidea Zhang &amp; Chen, sp. nov., S. (S.) idiasta Cui &amp; Chen, sp. nov., S. (S.) kanda Cui &amp; Chen, sp. nov., S. (S.) labao Li &amp; Chen, sp. nov., S. (S.) lancang Li &amp; Chen, sp. nov., S. (S.) latifoliacea Wang &amp; Chen, sp. nov., S. (S.) liusanjieae Li &amp; Chen, sp. nov., S. (S.) magniflava Cui &amp; Chen, sp. nov., S. (S.) mailangang Li &amp; Chen, sp. nov., S. (S.) marenubila Cui &amp; Chen, sp. nov., S. (S.) menghai Zhang &amp; Chen, sp. nov., S. (S.) menglian Li &amp; Chen, sp. nov., S. (S.) minutiflava Li &amp; Chen, sp. nov., S. (S.) multiprocera Li &amp; Chen, sp. nov., S. (S.) nayun Li &amp; Chen, sp. nov., S. (S.) nigridentata Wang &amp; Chen, sp. nov., S. (S.) nigripalpula Cui &amp; Chen, sp. nov., S. (S.) otphylla Cui &amp; Chen, sp. nov., S. (S.) radiciflava Zhang &amp; Chen, sp. nov., S. (S.) rava Cui &amp; Chen, sp. nov., S. (S.) sciophila Li &amp; Chen, sp. nov., S. (S.) septencolorata Li &amp; Chen, sp. nov., S. (S.) serrata Zhang &amp; Chen, sp. nov., S. (S.) silvestrella Zhang &amp; Chen, sp. nov., S. (S.) simola Cui &amp; Chen, sp. nov., S. (S.) yani Li &amp; Chen, sp. nov., S. (S.) yixiang Zhang &amp; Chen, sp. nov., S. (S.) zaduo Cui &amp; Chen, sp. nov., and S. (S.) zhuoma Cui &amp; Chen, sp. nov. We also provided a complete list of Chinese Steganina species together with their geographical distributions. In addition, the majority of currently available DNA barcode (partial sequence of the mitochondrial cytochrome c oxidase subunit I (COI) gene) sequences of this subgenus (435 sequences of 102 spp.) were employed in a molecular analysis for species delimitation. Taken together, morphology- and molecular-based species delimitation results reached a consensus for an overwhelming majority of these Steganina species (98 of 102 spp.).

Research paper thumbnail of <strong>The rediscovery of the holotype of <em>Kotonisia kanoi </em>Matsumura, 1938 with notes on Matsumura's type specimens of Fulgoroidea (Insecta: Hemiptera: Fulgoromorpha)</strong>

Zootaxa, Dec 21, 2009

Shonen Matsumura (1872–1960), the founder of entomology in Japan, is no doubt among the most infl... more Shonen Matsumura (1872–1960), the founder of entomology in Japan, is no doubt among the most influential and prolific entomologists. He produced a series of works, illustrated lists of the insects and described as many as 1200 new species. Unfortunately, most of the types he named and collected from Taiwan were brought to Japan in the late 1940s, and were hard to access by foreign students until relatively recently.Matsumura&amp;#39;s type depositions contain brief descriptions and, sometimes, only female specimens were available for his types. This prevented sound revisionary work in Taiwan. In addition, Matsumura did not designate holotypes in his early works but simply indicated how many specimens he had.

Research paper thumbnail of Expression Profile and Gene Age Jointly Shaped the Genome-Wide Distribution of Premature Termination Codons in a Drosophila melanogaster Population

Molecular Biology and Evolution, Nov 3, 2014

Widespread premature termination codon mutations (PTCs) were recently observed in human and fly p... more Widespread premature termination codon mutations (PTCs) were recently observed in human and fly populations. We took advantage of the population resequencing data in the Drosophila Genetic Reference Panel to investigate how the expression profile and the evolutionary age of genes shaped the allele frequency distribution of PTCs. After generating a high-quality data set of PTCs, we clustered genes harboring PTCs into three categories: genes encoding low-frequency PTCs (1.5%), moderate-frequency PTCs (1.5-10%), and high-frequency PTCs (410%). All three groups show narrow transcription compared with PTC-free genes, with the moderate-and high-PTC frequency groups showing a pronounced pattern. Moreover, nearly half (42%) of the PTC-encoding genes are not expressed in any tissue. Interestingly, the moderate-frequency PTC group is strongly enriched for genes expressed in midgut, whereas genes harboring high-frequency PTCs tend to have sex-specific expression. We further find that although young genes born in the last 60 My compose a mere 9% of the genome, they represent 16%, 30%, and 50% of the genes containing low-, moderate-, and highfrequency PTCs, respectively. Among DNA-based and RNA-based duplicated genes, the child copy is approximately twice as likely to contain PTCs as the parent copy, whereas young de novo genes are as likely to encode PTCs as DNA-based duplicated new genes. Based on these results, we conclude that expression profile and gene age jointly shaped the landscape of PTC-mediated gene loss. Therefore, we propose that new genes may need a long time to become stably maintained after the origination.

Research paper thumbnail of The rediscovery of the holotype of Kotonisia kanoi Matsumura, 1938 with notes on Matsumura's type specimens of Fulgoroidea (Insecta: Hemiptera: Fulgoromorpha)

Zenodo (CERN European Organization for Nuclear Research), Dec 31, 2009

Tsaur, Shun-Chern (2009): The rediscovery of the holotype of Kotonisia kanoi Matsumura, 1938 with... more Tsaur, Shun-Chern (2009): The rediscovery of the holotype of Kotonisia kanoi Matsumura, 1938 with notes on Matsumura's type specimens of Fulgoroidea (Insecta: Hemiptera: Fulgoromorpha). Zootaxa 2315: 66-68, DOI: 10.5281/zenodo.275420

Research paper thumbnail of Cryptic diversity in the subgenus Oxyphortica (Diptera, Drosophilidae, Stegana)

PeerJ, Oct 29, 2021

Phylogenetic relationships of the subgenus Oxyphortica were reconstructed based on two mitochondr... more Phylogenetic relationships of the subgenus Oxyphortica were reconstructed based on two mitochondrial genes (COI and ND2). The results revealed the paraphyly of Oxyphortica and supported high levels of cryptic diversity within this subgenus. By integrating morphological characteristics and molecular evidence, we identified 17 new species as members of Oxyphortica: S. (O.

Research paper thumbnail of Positive selection driving the evolution of a gene of male reproduction, Acp26Aa, of Drosophila: II. Divergence versus polymorphism

Molecular Biology and Evolution, Aug 1, 1998

The evolution of the gene for a male ejaculatory protein, Acp26Aa, has been shown to be driven by... more The evolution of the gene for a male ejaculatory protein, Acp26Aa, has been shown to be driven by positive selection when nonsibling species in the Drosophila melanogaster subgroup are compared. To know if selection has been operating in the recent past and to understand the details of its dynamics, we obtained DNA sequences of Acp26Aa and the nearby Acp26Ab gene from 39 D. melanogaster chromosomes. Together with the 10 published sequences, we analyzed 49 sequences from five populations in four continents. The southern African population is somewhat differentiated from all other populations, but its nucleotide diversity is lower at these two loci. We find the following results for Acp26Aa: (1) The R : S (replacement : silent changes) ratio is significantly higher in the between-species comparisons than in the within-species data by the McDonald and Kreitman test. Positive selection is probably responsible for the excess of amino acid replacements between species. (2) However, within-species nucleotide diversity is high. Neither the Tajima test nor the Fu and Li test indicates a reduction in nucleotide diversity due to positive selection in the recent past. (3) The newly derived nucleotides in D. melanogaster are at high frequency significantly more often than predicted by the neutral equilibrium. Since the nearby Acp26Ab gene does not show these patterns, these observations cannot be attributed to the characteristics of this chromosomal region. We suggest that positive selection is active, but may be weak, for each amino acid change in the Acp26Aa gene.

Research paper thumbnail of The nucleotide changes governing cuticular hydrocarbon variation and their evolution in <i>Drosophila melanogaster</i>

Proceedings of the National Academy of Sciences of the United States of America, Mar 20, 2001

The cuticular hydrocarbon (CH) pheromones in Drosophila melanogaster exhibit strong geographic va... more The cuticular hydrocarbon (CH) pheromones in Drosophila melanogaster exhibit strong geographic variation. African and Caribbean populations have a high ratio of 5,9 heptacosadiene͞ 7,11 heptacosadiene (the ''High'' CH type), whereas populations from all other areas have a low ratio (''Low'' CH type). Based on previous genetic mapping, DNA markers were developed that localized the genetic basis of this CH polymorphism to within a 13-kb region. We then carried out a hierarchical search for diagnostic nucleotide sites starting with four lines, and increasing to 24 and 43 lines from a worldwide collection. Within the 13-kb region, only one variable site shows a complete concordance with the CH phenotype. This is a 16-bp deletion in the 5 region of a desaturase gene (desat2) that was recently suggested to be responsible for the CH polymorphism on the basis of its expression [

Research paper thumbnail of A new species of Cixius from the United States (Homoptera: Fulgoroidea: Cixiidae)

Research paper thumbnail of The Cixiidae of Taiwan, Part VII: Tribe Pintaliini (Hemiptera: Fulgoroidea)

Zoological Studies, 2003

Shun-Chern Tsaur and Tung-Ching Hsu (2003) The Cixiidae of Taiwan, Part VII: tribe Pintaliini (He... more Shun-Chern Tsaur and Tung-Ching Hsu (2003) The Cixiidae of Taiwan, Part VII: tribe Pintaliini (Hemiptera: Fulgoroidea). Zoological Studies 42(3): 431-443. This study completes the first taxonomic revision of the planthopper family Cixiidae in Taiwan with emphasis on the morphology of the male and female genitalia. In total, 20 genera and 155 species are recognized. A key to the tribes of Bothriocerinae was given in a previous work. In this paper, keys to both the subfamilies and tribes of the Cixiidae are presented. The 5 tribes Pentastirini, Bothriocerini, Bennini, Stenophlepsini, and Cixiini were previously revised by the authors, and the remaining tribe, Pintaliini, is treated here with 6 species. Neocarpia maai represents a new genus and new species. Kirbyana lini, Eucarpia stellata, E. specialis, and E. truncata are described as new species. The male of K. pagana (Melichar) is described and illustrated for the first time.

Research paper thumbnail of <p><strong>Revision of the subgenus <em>Phortica</em> (sensu stricto) (Diptera, Drosophilidae) from East Asia, with assessment of species delimitation using DNA barcodes</strong></p&gt

Zootaxa, Sep 30, 2019

A total of 50 (43 known and seven new) species in the subgenus Phortica (sensu stricto) were surv... more A total of 50 (43 known and seven new) species in the subgenus Phortica (sensu stricto) were surveyed and (re)described from China: P. bicornuta (Chen &amp; Toda, 1997); P. bipartita (Toda &amp; Peng, 1992); P. biprotrusa (Chen &amp; Toda, 1998); P. cardua (Okada, 1977); P. chi (Toda &amp; Sidorenko, 1996); P. conifera (Okada, 1977); P. eparmata (Okada, 1977); P. eugamma (Toda &amp; Peng, 1990); P. excrescentiosa (Toda &amp; Peng, 1990); P. fangae (Máca, 1993); P. flexuosa (Zhang &amp; Gan, 1986); P. foliata (Chen &amp; Toda, 1997); P. gamma (Toda &amp; Peng, 1990); P. gigas (Okada, 1977); P. glabtabula Chen &amp; Gao, 2005; P. hainanensis (Chen &amp; Toda, 1998); P. hongae (Máca, 1993); P. huazhii Cheng &amp; Chen, 2008; P. iota (Toda &amp; Sidorenko, 1996); P. jadete Zhu, Cao &amp; Chen, 2018; P. kappa (Máca, 1977); P. lambda (Toda &amp; Peng, 1990); P. latifoliacea Chen &amp; Watabe, 2008; P. magna (Okada, 1960); P. okadai (Máca, 1977); P. omega (Okada, 1977); P. orientalis (Hendel, 1914); P. pangi Chen &amp; Wen, 2005; P. paramagna (Okada, 1971); P. perforcipata (Máca &amp; Lin, 1993); P. pi (Toda &amp; Peng, 1990); P. protrusa (Zhang &amp; Shi, 1997); P. pseudopi (Toda &amp; Peng, 1990); P. pseudotau (Toda &amp; Peng, 1990); P. psi (Zhang &amp; Gan, 1986); P. rhagolobos Chen &amp; Gao, 2008; P. saeta (Zhang &amp; Gan, 1986); P. setitabula Chen &amp; Gao, 2005; P. subradiata (Okada, 1977); P. tau (Toda &amp; Peng, 1990); P. uncinata Chen &amp; Gao, 2005; P. unipetala Chen &amp; Wen, 2005; P. allomega Gong &amp; Chen, sp. nov.; P. archikappa Gong &amp; Chen, sp. nov.; P. dianzangensis Gong &amp; Chen, sp. nov.; P. imbacilia Gong &amp; Chen, sp. nov.; P. liukuni Gong &amp; Chen, sp. nov.; P. tibeta Gong &amp; Chen, sp. nov.; and P. xianfui Gong &amp; Chen, sp. nov. In addition, seven new synonyms were recognized: P. acongruens (Zhang &amp; Shi, 1997), syn. nov.; P. antillaria (Chen &amp; Toda, 1997), syn. nov.; P. kukuanensis Máca, 2003, syn. nov.; P. linae (Máca &amp; Chen, 1993), syn. nov.; P. shillongensis (Singh &amp; Gupta, 1979), syn. nov.; P. takadai (Okada, 1977), syn. nov.; and P. watanabei (Máca &amp; Lin, 1993), syn. nov. A key to all Asian species (except for the eparmata species complex) of this subgenus was provided. All currently available DNA barcode (partial mitochondrial cytochrome c oxidase subunit I (COI) gene) sequences of this subgenus (217 sequences of 54 species) are employed in a molecular analysis using different species delimitation methods. The results indicate that approximately 68.5% (37 of 54 spp.) of Phortica (s. str.) species could be clearly distinguished from closely related morphospecies or cryptic species.

Research paper thumbnail of Survey of the genus Stegana Meigen (Diptera, Drosophilidae) from Taiwan, with DNA barcodes and descriptions of three new species

Zoological Studies, Jan 15, 2014

Background: Twelve Stegana species have been reported from Taiwan, yet only four were also record... more Background: Twelve Stegana species have been reported from Taiwan, yet only four were also recorded from Mainland China. This may not reflect the actual fauna between both sides of the strait. This report mainly deals with a fly collection of the genus Stegana during a short visit to Taiwan in 2012. It represents the most recent drosophilid faunal survey of Taiwan associating with bleeding trees. Results: In this study, 17 species were recognized including three new ones and eight new records. They are Stegana (Oxyphortica) convergens (de Meijere, 1911); Stegana (Oxyphortica) nigripennis (Hendel, 1914); Stegana

Research paper thumbnail of Sex in Drosophila mauritiana: A Very High Level of Amino Acid Polymorphism in a Male Reproductive Protein Gene, Acp26Aa

Molecular Biology and Evolution, 2001

Many genes pertaining to male reproductive functions have been shown to evolve rapidly between sp... more Many genes pertaining to male reproductive functions have been shown to evolve rapidly between species, and evidence increasingly suggest the influence of positive Darwinian selection. The accessory gland protein gene (Acp26Aa) of Drosophila is one such example. In order to understand the mechanism of selection, it is often helpful to examine the pattern of polymorphism. We report here that the level of amino acid polymorphism in the Nterminal quarter of Acp26Aa is high in Drosophila melanogaster and is unprecedented in its sibling species Drosophila mauritiana. We postulate that (1) this N-terminal segment may play a role in sperm competition, and (2) D. mauritiana may have been under much more intense sexual selection than other species. Both postulates have important ramifications and deserve to be tested rigorously.

Research paper thumbnail of Gene duplication and speciation in <i>Drosophila</i> : Evidence from the <i>Odysseus</i> locus

Proceedings of the National Academy of Sciences of the United States of America, Aug 10, 2004

The importance of gene duplication in evolution has long been recognized. Because duplicated gene... more The importance of gene duplication in evolution has long been recognized. Because duplicated genes are prone to diverge in function, gene duplication could plausibly play a role in species differentiation. However, experimental evidence linking gene duplication with speciation is scarce. Here, we show that a hybridmale sterility gene, Odysseus (OdsH), arose by gene duplication in the Drosophila genome. OdsH has evolved at a very high rate, whereas its most immediate paralog, unc-4, is nearly identical among species in the Drosophila melanogaster subgroup. The disparity in their sequence evolution is echoed by the divergence in their expression patterns in both soma and reproductive tissues. We suggest that duplicated genes that have yet to evolve a stable function at the time of speciation may be candidates for ''speciation genes,'' which is broadly defined as genes that contribute to differential adaptation between species.

Research paper thumbnail of The phylogeny of closely related species as revealed by the genealogy of a speciation gene, <i>Odysseus</i>

Proceedings of the National Academy of Sciences of the United States of America, Apr 25, 2000

Molecular differentiation between races or closely related species is often incongruent with the ... more Molecular differentiation between races or closely related species is often incongruent with the reproductive divergence of the taxa of interest. Shared ancient polymorphism and͞or introgression during secondary contact may be responsible for the incongruence. At loci contributing to speciation, these two complications should be minimized (1, 2); hence, their variation may more faithfully reflect the history of the species' reproductive differentiation. In this study, we analyzed DNA polymorphism at the Odysseus (OdsH) locus of hybrid sterility between Drosophila mauritiana and Drosophila simulans and were able to verify such a prediction. Interestingly, DNA variation only a short distance away (1.8 kb) appears not to be influenced by the forces that shape the recent evolution of the OdsH coding region. This locus thus may represent a test case of inferring phylogeny of very closely related species. S pecies are delineated by shared reproductive physiology, development, sexual behavior, and morphology (3, 4). Divergence in these systems is manifested as hybrid sterility, hybrid inviability, premating isolation, and morphological differences, respectively. Races are less well defined but members often may cluster by morphological traits. One of the paradoxes concerning race or species differentiation is the common occurrences of ambiguity in distinguishing taxa by molecular means, even when grouping by reproductive or morphological traits is straightforward and clearcut. Human racial differentiation may be a most obvious example in which many morphological characters cluster by geographical origin, whereas almost all molecular polymorphisms are extensively shared among races (5). Morphological distinction among dog breeds is another example (6). In Drosophila, sexual isolation between the Zimbabwe and non-African races of Drosophila melanogaster is clearly determined by many genes spread over the autosomal genome (7), and yet, recent molecular data have failed to show much differentiation at autosomal loci (8, 9). An explanation for the discordance between the ''reproductive'' and ''molecular'' phylogeny is that genomes may be mosaics with respect to molecular genealogy, as illustrated in Fig. 1. Most loci, chosen without regard to their roles in reproductive differentiation, may not reflect the biological divergence in their sequence polymorphism because of either shared ancient polymorphism or gene introgression through secondary contact (Fig. 1b). Ancient polymorphism may persist until present day in species with large population sizes (10, 11), and gene introgression, even at a very low level, may be sufficient to obliterate differentiation (12). In this context, we shall consider separately ''speciation genes,'' defined as loci that contribute directly to some aspects of biological divergence between closely related species (such as gametogenesis, behavior, or morphology). A hypothesis, proposed in various forms (1, 2, 13, 14), is that ''speciation genes'' may record a phylogenetic history more consistent with species' reproductive biology. This is because polymorphism and divergence at these loci should be relatively unaffected by shared polymorphisms or introgressions (see the legend of Fig. 1a). The cloning of the Odysseus (OdsH, H for homeodomain) locus of hybrid male sterility in the Drosophila simulans clade (15) therefore provides an opportunity to test this hypothesis. The sibling species of D. simulans, Drosophila mau-This paper was submitted directly (Track II) to the PNAS office. Data deposition: The sequences reported in this paper have been deposited in the GenBank database (accession nos. AF254805 for D. simulans, AF254806 for D. mauritiana, and AF254807 for D. sechellia).

Research paper thumbnail of Two New Species of Oliarus from Taiwan (Homoptera : Fulgoroidea : Cixiidae), with Proposition and Discussion on 0. Horjshanus Group 1

~.. . . / . Shun-Chern Tsaur (1990) Two new species of Oliarus from Taiwan (Homo­ ptera : Fulgoro... more ~.. . . / . Shun-Chern Tsaur (1990) Two new species of Oliarus from Taiwan (Homo­ ptera : Fulgoroidea : Cixiidae), with proposition and discussion on 0. horishanus group. Bull. Inst. Zoo!., Academia Sinica 29(3): 135-139. Two new species of Oliarus ( 0. hsui and 0. shiaoi) from Taiwan have been added. The 0 . horishanus group proposed here includes five species; 0. horishanus, 0. hsui, 0. nigroner\latus, 0. shiaoi and 0 . yangi.