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Last Updated :2024/10/18

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Master

Affiliation (Master)

  • Research Faculty of Agriculture　Fundamental AgriScience Research　Agrobiology and Bioresources

Affiliation (Master)

  • Research Faculty of Agriculture　Fundamental AgriScience Research　Agrobiology and Bioresources

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Profile and Settings

Degree

  • PhD
  • MA (Environmental Science)

Profile and Settings

  • Name (Japanese)

    Yoshizawa

  • Name (Kana)

    Kazunori

  • Name

    200901009650396974

Alternate Names

Achievement

Research Interests

  • 昆虫学 ・系統学・分類学   Evolution   Morphology   Taxonomy   Phylogenetics   Entomology   

Research Areas

  • Life sciences / Biodiversity and systematics

Research Experience

  • 2009/09 - Today Hokkaido University Graduate School of Agriculture Research Faculty of Agriculture
  • 2001/04 - Today Hokkaido University The Hokkaido University Museum
  • 2019/04 - 2021/03 京都大学大学院 理学研究科 客員准教授
  • 2007/04 - 2009/08 北海道大学大学院農学研究 助教
  • 2000/08 - 2007/03 北海道大学大学院農学研究科 助手
  • 2003/11 - 2004/10 University of Illinois, Illinois Natural History Survey Visiting Faculty
  • 1999/04 - 2000/08 Kyushu University
  • 1996/04 - 1999/03 日本学術振興会特別研究員 DC1（九州大学）

Education

  • 1996/04 - 1999/03  Kyushu University
  • 1994/04 - 1996/03  Kyushu University
  • 1990/04 - 1994/03  Kyushu University  School of Agriculture

Awards

  • 2017/09 Ig Nobel Prize in Biology
     Discovery of a female penis and male vagina in a cave insect 
    受賞者: YOSHIZAWA K;Ferreira R;Kamimura Y;Lienhard C受賞者：Yoshizawa, K., Ferreira, R.L., Kamimura, Y., Lienhard, C.
  • 2017/02 北海道大学 研究総長賞(奨励賞)
     
    受賞者: 吉澤 和徳
  • 2016/02 北海道大学 総長賞(奨励賞)
     
    受賞者: 吉澤 和徳
  • 2013/09 日本昆虫学会 学会賞
     
    受賞者: 吉澤和徳;我妻睦美

Published Papers

  • Tomoki Hirose, Kazunori Yoshizawa
    The Science of Nature 111 (5) 0028-1042 2024/09/23 [Refereed][Not invited]
  • Azuma Kawata, Naoki Ogawa, Kazunori Yoshizawa
    Journal of Morphology 285 (6) 0362-2525 2024/05/27 [Refereed][Not invited]
     
    Abstract Although the monophyly of Paraneoptera (=hemipteroid orders or Acercaria, composed of Psocodea, Thysanoptera and Hemiptera) has been widely accepted morphologically, the results from molecular phylogenetic and phylogenomic analyses contradict this hypothesis. In particular, phylogenomic analyses provide strong bootstrap support for the sister group relationship between Psocodea and Holometabola, that is, paraphyly of Paraneoptera. Here, we examined the pterothoracic musculature of Paraneoptera, as well as a wide range of other neopterous insect orders, and analysed its phylogenetic implication. By using the synchrotron microcomputed tomography (µCT) and parsimony‐based ancestral state reconstruction, several apomorphic conditions suggesting the monophyly of Paraneoptera, such as the absence of the II/IIItpm7, IIscm3, IIIspm2 and IIIscm3 muscles, were identified. In contrast, no characters supporting Psocodea + Holometabola were recovered from the thoracic muscles. These results provide additional support for the monophyly of Paraneoptera, together with the previously detected morphological apomorphies of the head, wing base, and abdomen.
  • Tatsuki MATOBA, Yoshitaka KAMIMURA, Kazunori YOSHIZAWA
    Entomological Science 27 (2) 1343-8786 2024/04/03 
    Abstract Female fritillary butterflies (Lepidoptera: Nymphalidae: Argynnini) are known to have diversified genital structures, especially at the entrance of the bursa copulatrix. However, the origin and function of such elaborated structures are poorly known to date. We examined the male and female genital structures of six Japanese species selected from five genera/subgenera [Argynnis (Argynnis), Argynnis (Damora), Argynnis (Argyronome), Fabriciana, and Speyeria] of the Argynnini tribe and found that traumatic copulation is widespread in Argynnini. Various male genital structures, such as the uncus, valva, and phallus, can cause wounds to the female genitalia. The eighth venter of the female, where the highest morphological diversity in female genitalia was detected, is hooked and held by the male uncus during copulation. It is very likely that the diversified female genital structure coevolved with the male uncus as a counterpart for traumatic coupling.
  • Shin-ichi Kudo, Tomohiro Harano, Jing-Fu Tsai, Kazunori Yoshizawa, Nobuyuki Kutsukake
    The American Naturalist 0003-0147 2024/02/27
  • MASAYASU SAWADA, KAZUNORI YOSHIZAWA, ISSEI OHSHIMA
    Zootaxa 5410 (2) 177 - 198 1175-5326 2024/02/13 
    Two new species of the genus Cameraria Chapman, 1902, Cameraria riparia sp. nov. and Cameraria corni sp. nov., are described from Japan. Larval setal maps, illustrations and/or photographs of wing patterns, wing venations and genitalia are provided for both species, and the systematic positions of the two species are discussed based on their morphologies. The larvae of C. riparia feed on Salix spp. and make blotch mines, whereas C. corni larvae feed on Cornus kousa subsp. kousa and make linear blotch mines. C. riparia was collected on Hokkaido and Honshu Islands, and considerable differences in the male genital characters were detected between the two populations. However, a molecular phylogenetic analysis based on the COI barcode region indicated that there was no clear genetic differentiation between the two populations (maximum divergence, 1.41%). C. corni was collected on Honshu and Kyushu Islands, and this is the first record of the trophic association of Lithocolletinae with Cornaceae.
  • Tatsuki Matoba, Yoshitaka Kamimura, Kazunori Yoshizawa
    Organisms Diversity & Evolution 23 (4) 871 - 879 1439-6092 2023/10/18
  • Zixin Cheng, Yoshitaka Kamimura, Rodrigo L. Ferreira, Charles Lienhard, Kazunori Yoshizawa
    The Science of Nature 110 (3) 0028-1042 2023/06/01
  • Zixin Cheng, Yoshitaka Kamimura, Rodrigo L. Ferreira, Charles Lienhard, Kazunori Yoshizawa
    Royal Society Open Science 10 (1) 2023/01/11 
    Brazilian sex-role reversed cave insects (genus Neotrogla ) have a striking structure called the gynosome (or female penis), which deeply penetrates male vagina-like genitalia during copulation to receive nutritious semen. However, the protruding and retracting mechanisms of the female penis, including their evolutionary origin, are poorly understood. By using micro-computed tomography (µCT), we compared the genital morphology and musculature between species with a gynosome and others lacking this structure. As a result, we discovered two groups of muscles related to the protrusion and retraction of gynosomes. These muscles were also observed in species with non-protrusible prepenis. This suggests that evolution of these muscles preceded the acquisition of the protruding function of the gynosome, originally having a putative stimulatory function to receive nutritious semen. This intermediate stage probably allowed for the reversal of genital functions.
  • An additional new species of Trichadenotecnum from Japan (Psocodea: Psocidae).
    Yoshizawa, K
    Insecta matsumurana, new series 78 21 - 25 2022/10 [Refereed]
  • Oriental Psyllipsocus (Psocodea: 'Psocoptera': Psyllipsocidae): checklist, new records and description of four new species from Southeast Asia .
    Lienhard, C, Yoshizawa, K, Idris, A
    Insecta matsumurana, new series 78 1 - 19 2022/10 [Refereed]
  • Azuma Kawata, Naoki Ogawa, Kazunori Yoshizawa
    Journal of Morphology 283 (8) 1106 - 1119 0362-2525 2022/07/18 
    Abstract The thoracic musculature of the insect order Psocodea has been examined in only a few species of a single suborder to date. In the present study, we examined the thoracic musculature of species selected from all three suborders of Psocodea to elucidate the ground plan of the order and to examine the phylogenetic utility of the character system. The sister‐group relationship between the suborders Troctomorpha and Psocomorpha received support from two novel nonhomoplasious synapomorphies, although the support from other morphological characters for this relationship is ambiguous. The sister‐group relationship between the infraorders Epipsocetae and Psocetae also received support from one nonhomoplasious synapomorphy, although no other morphological characters supporting this relationship have been identified to date. The present examination revealed the potential of thoracic muscle characters for estimating deep phylogeny, possibly including interordinal relationships.
  • Sheng Li, Kazunori Yoshizawa, Qiuzhu Wang, Dong Ren, Ming Bai, Yunzhi Yao
    Frontiers in Ecology and Evolution 10 2022/06/09 
    Two species of psocids discovered from the Mid-Cretaceous Burmese amber, Latempheria kachinensis Li, Yoshizawa, and Yao, gen. et sp. nov. and Burmempheria curvatavena Li, Yoshizawa, and Yao, sp. nov., are described and assigned to the Empheriidae (Trogiomorpha: Atropetae) family. A phylogenetic analysis of the infraorder Atropetae is conducted based on 38 morphological characters of three outgroups and fifteen ingroups, which supported the monophyly of Atropetae including fossil and extant taxa. In the phylogenetic result, all the genera of fossil families Empheriidae and Archaeatropidae form a monophyletic group, sister to the extant members of Atropetae. The two fossil families also share a lot of similarities in morphology, locality, and geological period. Recently discovered fossil species exhibited combined morphological characters of both families. Based on these observations and the results of the phylogenetic analysis, Archaeatropidae is treated here as a new junior synonym of Empheriidae.
  • Zixin Cheng, Kazunori Yoshizawa
    Arthropod Structure & Development 66 101141 - 101141 1467-8039 2022/01
  • Yoshitaka Kamimura, Kazunori Yoshizawa, Charles Lienhard, Rodrigo L. Ferreira, Jun Abe
    BMC Ecology and Evolution 21 (1) 2021/12 [Refereed]
     
    Abstract Background Many male animals donate nutritive materials during courtship or mating to their female mates. Donation of large-sized gifts, though costly to prepare, can result in increased sperm transfer during mating and delayed remating of the females, resulting in higher paternity. Nuptial gifting sometimes causes severe female-female competition for obtaining gifts (i.e., sex-role reversal in mate competition) and selection on females to increase their mating rate, changing the intensity of sperm competition and the resultant paternity gains. We built a theoretical model to simulate such coevolutionary feedbacks between nuptial gift size (male trait) and propensity for multiple mating (female trait). Donation of nuptial gifts sometimes causes development of female persistence trait for gift acquisition. We also analyzed the causes and consequences of this type of traits, taking double receptacles for nutritious seminal gifts, which are known to occur in an insect group with a “female penis” (Neotrogla spp.), as an illustrative example. Results Our individual-based simulations demonstrated that female-female competition for male-derived nutrients always occur when the environment is oligotrophic and mating costs are low for females. However, a positive correlation between donated gift size and the resultant paternity gain was a requisite for the co-occurrence of large gifts and females’ competitive multiple mating for the gifts. When gift donation satisfied female demands and thus resulted in monandry, exaggeration of nuptial gift size also occurred under the assumption that the last male monopolizes paternity. The evolution of double slots for gift acquisition and digestion (female persistence trait) always occurred when males could not satisfy the demands of females for gifts. However, through coevolutionary reduction in male gift size, fixation of this trait in a population drastically reduced the average female fitness. Conclusion Sperm usage patterns, which have rarely been examined for animals with nuptial gifts, can be a critical factor for determining the extent of exaggeration in nuptial gifting. Sex-role reversals in mate competition, as a result of donation of nuptial gifts from males to females, can involve the evolution of male-like, persistent traits in females that reduce population productivity, as is the case with persistence traits in males.
  • Mamoru Takahashi, Kazunori Yoshizawa
    Insect Systematics & Evolution 53 (2) 185 - 199 1399-560X 2021/11/02 
    Abstract The forewing base structure of the orthopteran suborder Ensifera, a group of insects well known for their sound-producing behavior, was examined by using light microscopy, confocal laser scanning microscopy, and µCT. We detected significant novel characters, such as functional changes in the tpm9 muscle from the flexor to the extensor of the forewing that are likely associated with the acquisition of sound-producing behavior using forewings. Phylogenetic analysis based on the characters selected from the forewing base structure showed that the character system contains a strong phylogenetic signal supporting the monophyly of Ensifera, Tettigonioidea and Grylloidea as well as the sister-group relationship between Grylloidea and Gryllotalpoidea, but no apomorphies to resolve the deep phylogeny among superfamilies could be determined from this character system.
  • The earliest fossil record of the suborder Psocomorpha (Insecta: Psocodea) from mid-Cretaceous Burmese amber, with description of a new genus and species.
    Yoshizawa, K, Yamamoto, S
    Insecta matsumurana, new series 77 1 - 15 2021/10 [Refereed]
  • Corentin Jouault, Kazunori Yoshizawa, Marina Hakim, Diying Huang, André Nel
    Cretaceous Research 126 104890 - 104890 0195-6671 2021/10 [Refereed]
  • Akiko Otsuki, Kazunori Yoshizawa, Shin-Ichi Akimoto
    Biological Journal of the Linnean Society 134 (2) 316 - 330 0024-4066 2021/09/10 [Refereed]
     
    Abstract To elucidate the effect of Pleistocene climatic fluctuations on the historical distribution and geographical genetic structure of temperate Japanese species, we performed phylogeographical and demographic analyses using mitochondrial gene sequences obtained from the stonefly species Kamimuria tibialis, sampled from four main islands of the Japanese Archipelago (i.e. Hokkaido, Honshu, Shikoku and Kyushu) and Tsushima Island. We detected three main clades with distinct geographical distributions, including the Tsushima, Kyushu and Hokkaido–Honshu–Shikoku phylogroups. These groups were estimated to have diverged from one another 0.54–2.02 Mya, suggesting they have undergone several glacial cycles in different refugia. Our results showed that during the glacial epochs and with a fall in sea-level, gene flow was limited among Tsushima and Kyushu, and among Hokkaido and Honshu, probably because the straits between these islands are deep. The population in Kyushu and Shikoku, the southernmost islands, exhibited high genetic diversity, with two distinct haplotype lineages occurring sympatrically. These results suggest that the population division into multiple refugia and the existence of stable southern refugia have contributed to the high genetic diversity of the species in this region.
  • Tatsushi Takayanagi, Kazunori Yoshizawa
    Biodiversity Data Journal 9 1314-2836 2021/07/08 [Refereed]
     
    Caenis rivulorum Eaton, 1884 is widely distributed and has been reported from a wide range in the Palearctic Region. We report this species from Japan for the first time, from five localities of Hokkaido, based on morphology and molecular data.
  • Oscar Fernando Saenz Manchola, Stephany Virrueta Herrera, Lorenzo Mario D'Alessio, Kazunori Yoshizawa, Alfonso Neri García Aldrete, Kevin P. Johnson
    Systematic Entomology 46 (4) 938 - 951 0307-6970 2021/07 [Refereed]
     
    ABSTRACT Psocodea (booklice and parasitic lice) is an order of insects containing species with extensive mitochondrial genome rearrangements, particularly within the suborder Troctomorpha, in which some species possess an extremely fragmented mitochondrial genome with several small minichromosomes. In the remaining suborders of Psocodea, there are groups with the ancestral pancrustacean arrangement, quite extensive rearrangements (e.g. Trogiomorpha), or in which the small number of species analysed to date have rearrangements of only a few protein‐coding genes and/or tRNAs (e.g. Psocomorpha). Despite the apparent high rate of rearrangements in the order as a whole, a small number of complete mitochondrial genomes are available, especially for suborder Psocomorpha, the largest free‐living suborder. To understand the evolution of the gene arrangement of the mitochondrial genome within Psocomorpha and its phylogenetic implications, we assembled and analysed the mitochondrial genomes of 33 species of bark lice belonging to nine families in two infraorders. Within the infraorder Homilopsocidea, four families were analysed, mainly from Lachesillidae (which included 22 species of this family). Within the infraorder Caeciliusetae, seven species representing five families were analysed. Mitochondrial gene rearrangements were identified in seven of the nine families. Some of these rearrangements were unique to a single species, while some contained phylogenetic signal, being shared by related species. These rearrangements typically corresponded to transpositions and inversions of tRNAs, possibly caused by tandem duplication–random loss (TDRL) and/or recombination events. Phylogenetic analyses of mitochondrial gene sequences provided phylogenetic resolution for several branches of the tree, including monophyly of Lachesillinae. The genus Hemicaecilius Enderlein was found to be embedded within the genus Lachesilla Westwood, rending the latter paraphyletic. Monophyly was also never recovered for Lachesillidae and Elipsocidae as currently defined. However, instability was observed for some higher level relationships within Psocomorpha, including the relationships among the major clades of Lachesillidae.
  • Robert S de Moya, Kazunori Yoshizawa, Kimberly K O Walden, Andrew D Sweet, Christopher H Dietrich, Johnson Kevin P
    Systematic Biology 70 (4) 719 - 738 1063-5157 2021/06/16 [Refereed]
     
    Abstract The insect order Psocodea is a diverse lineage comprising both parasitic (Phthiraptera) and nonparasitic members (Psocoptera). The extreme age and ecological diversity of the group may be associated with major genomic changes, such as base compositional biases expected to affect phylogenetic inference. Divergent morphology between parasitic and nonparasitic members has also obscured the origins of parasitism within the order. We conducted a phylogenomic analysis on the order Psocodea utilizing both transcriptome and genome sequencing to obtain a data set of 2370 orthologous genes. All phylogenomic analyses, including both concatenated and coalescent methods suggest a single origin of parasitism within the order Psocodea, resolving conflicting results from previous studies. This phylogeny allows us to propose a stable ordinal level classification scheme that retains significant taxonomic names present in historical scientific literature and reflects the evolution of the group as a whole. A dating analysis, with internal nodes calibrated by fossil evidence, suggests an origin of parasitism that predates the K-Pg boundary. Nucleotide compositional biases are detected in third and first codon positions and result in the anomalous placement of the Amphientometae as sister to Psocomorpha when all nucleotide sites are analyzed. Likelihood-mapping and quartet sampling methods demonstrate that base compositional biases can also have an effect on quartet-based methods.[Illumina; Phthiraptera; Psocoptera; quartet sampling; recoding methods.]
  • Kazunori Yoshizawa, Yuri M Marusik, Izumi Yao, Charles Lienhard
    Insect Systematics and Diversity 4 (6) 2020/11/01 [Refereed]
     
    Abstract The systematic placement of an enigmatic psocid family restricted to Africa, Lesneiidae, was estimated by using a multiple gene data set. The candidates for its close relatives are now classified under two different infraorders, the family Archipsocidae of the infraorder Archipsocetae or the families Elipsocidae/Mesopsocidae of the infraorder Homilopsocidea. The maximum likelihood and Bayesian analyses of the molecular data set strongly suggested that the Lesneiidae belongs to Homilopsocidea and forms a clade with Elipsocidae/Mesopsocidae/Eolachesillinae (Lachesillidae). However, the relationships among these (sub)families and Lesneiidae, including the monophyly of Elipsocidae and Mesopsocidae, were ambiguous or questionable, showing the necessity of further investigations for elucidating their relationships and validating the status of these families. Two species, L. johnsoni Yoshizawa & Lienhard, n. sp. and L. testudinata Yoshizawa & Lienhard, n. sp. (Psocodea: Lesneiidae), were described from South Africa. There appears to be a tight association between the reproductive biology and morphological specialization of this group.
  • Yoshizawa Kazunori, Charles Lienhard
    Dugesiana 27 (2) 173 - 181 1405-4094 2020/07/01 [Refereed]
     
    Five new species of the corniculum species group of the psocid genus Trichadenotecnum Enderlein, T. alfonsoi, T. lingens, T. palmula, T. affinifelix and T. separatum were described from Thailand. These species share a couple of male genital apomorphies with the other Oriental species of this species group recorded from China, Hong Kong, Malaysia, Singapore and Indonesia. The Japanese species of the corniculum group are considered to form a different clade placed to the sister of the Oriental species.
  • Yoshitaka Kamimura, Kazunori Yoshizawa, Charles Lienhard, Rodrigo L. Ferreira, Jun Abe
    bioRxiv 2020/06/23 [Not refereed]
     
    ABSTRACTMany male animals donate nutritive materials during courtship or mating to their female mates. Donation of large-sized gifts, though costly to prepare, can result in increased sperm transfer during mating and delayed remating of the females, resulting in a higher paternity Nuptial gifting sometimes causes severe female-female competition for obtaining gifts (i.e., sex-role reversal in mate competition) and female polyandry, changing the intensity of sperm competition and the resultant paternity gains. We built a theoretical model to analyze such coevolutionary feedbacks between nuptial gift size (male trait) and propensity for multiple mating (female trait). Our genetically explicit, individual-based computer simulations demonstrate that a positive correlation between donated gift size and the resultant paternity gain is a requisite for the co-occurrence of large-sized gifts and females’ competitive multiple mating for the gifts. When donation of gifts imposes monandry, exaggeration in nuptial gift size also occurs under the assumption that the last male monopolizes paternity. We also analyzed the causes and consequences of the evolution of a female persistence trait in trading of nuptial gifts, that is, double receptacles for nuptial gifts known to occur in an insect group with a “female penis” (Neotrogla spp.).
  • Kazunori Yoshizawa, Charles Lienhard
    Entomological Science 23 (2) 208 - 215 1343-8786 2020/06 [Refereed]
  • MEGUMI SHIMADA, KAZUNORI YOSHIZAWA
    Zootaxa 4779 (4) 501 - 521 1175-5326 2020/05/21 [Refereed]
     
    The Japanese species of the genus Strigiphilus Mjöberg, 1910 (Insecta: Phthiraptera: Philopteridae) are revised. Six species are recorded, including a new species belonging to the cursitans species-group: Strigiphilus stenocephalus new species, described from the type host Otus bakkamoena semitorques and based on specimens originally identified and reported by Uchida (1949) as Strigiphilus rostratus (Burmeister, 1838). A lectotype for Strigiphilus laticephalus (Uchida, 1949) (type host: Strix aluco yamadae) is designated and redescribed, and this louse species is synonymized under Strigiphilus cursor (Burmeister, 1838). Strigiphilus ceblebrachys (Denny, 1842), S. heterogenitalis Emerson & Elbel, 1957 and S. tuleskovi Balát, 1958 are recorded for the first time in Japan. Also, Strix uralensis and Otus sunia japonicus are recorded as new hosts for Strigiphilus heterogenitalis and S. tuleskovi respectively.
  • Yoko Matsumura, Rolf G. Beutel, José A. Rafael, Izumi Yao, Josenir T. Câmara, Sheila P. Lima, Kazunori Yoshizawa
    Systematic Entomology 45 (2) 349 - 364 0307-6970 2020/04 [Refereed][Not invited]
     
    Zoraptera is one of the most enigmatic and least understood orders in insects. Based on a wide taxon sampling from all continents where the group is known, we applied a phylogenetic approach using multiple DNA sequences to elucidate species-level relationships. The resulting phylogeny shows that Zoraptera is divided into three major clades, and that two comprise species distributed on different continents. The monophyly of these clades is at least partly supported by shared derived morphological features. The divergence age estimation and ancestral distribution area reconstruction suggest an ancient origin and early radiation initiated in the Permian. Plate tectonics theory suggests that the present distribution of Zoraptera was mainly established by vicariance, rather than dispersal. The three major clades probably originated on the Pangaea supercontinent, or alternatively on the linked Gondwana and Laurasia supercontinents. Their ancient origin explains previously found conspicuous interspecific variation of the genital apparatus, sperm structure and mating behaviour, in striking contrast to a highly conserved general body morphology. We compiled data of available reproductive features and reconstructed the character evolution. Our analyses revealed repeated acquisitions and/or losses of a hyperelongated intromittent organ, mating hooks and tergal protuberances.
  • Kazunori Yoshizawa, Charles Lienhard, Izumi Yao, Rodrigo L. Ferreira
    Entomological Science 22 (3) 334 - 338 1343-8786 2019/09 [Refereed][Not invited]
     
    The divergence date and ancestral distributional area of the psocid subfamily Speleketorinae, which includes taxa with reversed genitalia (female penis and male vagina of Afrotrogla and Neotrogla, tribe Sensitibillini), were estimated. The most basal divergence of the subfamily (between the North American Speleketor and the tribe Sensitibillini) was estimated to have occurred according to the separation between the North American continent and Gondwana, ca. 175 Ma. The most basal divergence of Sensitibillini (between African Afrotrogla + Sensitibilla and Brazilian Neotrogla) was estimated to have occurred according to the split of West Gondwana (separation between the African and South American continents), ca. 127 Ma. The biome of the ancestral distributional area of Sensitibillini (inland of West Gondwana) is believed to be arid to semi-arid, which might strengthen the reversed sexual selection and then facilitate the origin of preadaptive features related to the evolution of a female penis. All extant Sensitibillini species inhabit carbonatic caves, but geological evidence suggested independent shifts of these genera to the carbonatic cave habitat in the Tertiary/Quaternary.
  • KAMIMURA Yoshitaka, ABE Jun, FERREIRA Rodrigo L, YOSHIZAWA Kazunori
    Entomological Science 22 (1) 48 - 55 1343-8786 2019 [Refereed][Not invited]
  • Shimada, M, Yoshizawa, K
    Zootaxa 4612 (1) 133 - 137 1175-5326 2019 [Refereed][Not invited]
     
    Three species of chewing lice—Eucolpocephalum femorale (Piaget, 1880) (Amblycera: Menoponidae), Ibidoecus plataleae (Denny, 1842) (Ischnocera: Philopteridae) and Ardeicola plataleae (Linnaeus, 1758) (Ischnocera: Philopteridae)—are reported from the black-faced spoonbill (Platalea minor Temminck & Schlegel, 1849) in Japan. These three louse species are new records for this host. The black-faced spoonbill is classified as a “Critically Endangered” species in the IUCN Red List; although its species of lice are known to infest other species of spoonbills, these lice are endangered at the population level.
  • Cheng Z, Yoshizawa K
    Journal of Morphology 280 555 - 567 2019 [Refereed][Not invited]
  • Why did a female, penis evolve in, a, small, group of, cave insect
    BioEssays 41 (6) 1900005  1521-1878 2019 [Refereed][Invited]
     
    The evolution of a female penis is an extremely rare event and is only known to have occurred in a tribe of small cave insects, Sensitibillini (Psocodea: Trogiomorpha: Prionoglarididae). The female penis, which is protrudable and inserted into the male vagina-like cavity during copulation to receive semen, is thought to have evolved independently twice in this tribe, in the Brazilian Neotrogla and the African Afrotrogla. These findings strongly suggest that there are some factors unique to Sensitibillini that have facilitated female penis evolution. Here, several hypothetical factors are presented that may have enabled the evolution of the female penis in Sensitibillini. The female–female competition for nutritious semen, the oligotrophic environment, and the twin insemination slots with switching valve are considered to be the driving factors for female penis evolution. Additionally, the following factors are considered responsible for relaxing the constraint against female penis evolution: preexistence of the female-above mating position, the elongated duct connecting the female pre-penis with the sperm storage organ, and the small male genital cavity accepting the female genital tubercle bearing the opening of this duct. Understanding the factors enabling female penis evolution may also shed light on the evolution of the male penis.
  • YOSHIZAWA Kazunori
    Insecta matsumurana new series 75 19 - 28 2019 [Refereed][Not invited]
  • Yoshizawa, K, Ferreira, R, Yao, I, Lienhard, C, Kamimura, Y
    Biology Letters 2018/11 [Refereed][Not invited]
  • Kevin P. Johnson, Christopher H. Dietrich, Frank Friedrich, Rolf Beutel, Benjamin Wipfler, Ralph S. Peters, Julie M. Allen, Malte Petersen, Alexander Donath, Kimberly K. O. Walden, Alexey Kozlov, Lars Podsiadlowski, Christoph Mayer, Karen Meusemann, Alexandros Vasilikopoulos, Robert M. Waterhouse, Stephen Cameron, Christiane Weirauch, Daniel R. Swanson, Diana Percy, Nathaniel Hardy, Irene Terry, Shanlin Liu, Xin Zhou, Bernhard Misof, Hugh M. Robertson, Kazunori Yoshizawa
    PNAS 115 12775 - 12780 2018/11 [Refereed][Not invited]
  • YOSHIZAWA, K, KAMIMURA, Y, LIENHARD, C, FERREIRA, R.L, BLANKE, A
    eLife 2018/10 [Refereed][Not invited]
  • Naoki Ogawa, Kazunori Yoshizawa
    Journal of Morphology 279 (4) 517 - 530 1097-4687 2018/04/01 [Refereed][Not invited]
     
    Many four-winged insects have mechanisms that unite the forewings and hindwings in a single plane. Such an in-flight wing coupling apparatus may improve flight performance in four-winged insects, but its structure is variable among different insect groups. The wings of bark lice (Insecta: Psocodea: “Psocoptera”) also have an in-flight wing coupling apparatus, but to date, its morphology has not been studied in detail. In this study, we examined the wing-coupling structure in representative species of the three suborders of bark lice (Trogiomorpha, Troctomorpha, and Psocomorpha) and inferred its origin and transformation. We conclude that the main component of the psocodean wing coupling apparatus evolved once in the common ancestor via modification of cuticular structures at the apex of the forewing CuP vein. Morphological differences in components of the coupling structures are phylogenetically informative at the intraorder level and include an autapomorphy that characterizes Troctomorpha and a synapomorphy that supports a sister relationship between Troctomorpha and Psocomorpha.
  • Kazunori Yoshizawa, Kevin P. Johnson, Andrew D. Sweet, Izumi Yao, Rodrigo L. Ferreira, Stephen L. Cameron
    Molecular Phylogenetics and Evolution 119 118 - 127 1095-9513 2018/02/01 [Refereed][Not invited]
     
    The mitochondrial genome arrangement in the insect order Psocodea (booklice, barklice, and parasitic lice) is extremely variable. Genome organization ranges from the rearrangement of a few tRNAs and protein coding genes, through extensive tRNA and protein coding gene rearrangements, to subdivision into multiple mini-chromosomes. Evolution of the extremely modified mitochondrial genome in parasitic lice (Phthiraptera) has been the subject of several studies, but limited information is available regarding the mitochondrial genome organization of the more plesiomorphic, free-living Psocodea (formerly known as the “Psocoptera”). In particular, the ancestral state of the psocodean mitochondrial genome arrangement and the evolutionary pathway to the rearranged conditions are still unknown. In this study, we addressed mitochondrial evolutionary questions within the Psocodea by using mitochondrial genome sequences obtained from a wide range of Psocoptera, covering all three suborders. We identified seven types of mitochondrial genome arrangements in Psocoptera, including the first example in Psocodea of retention of the ancestral pancrustacean condition in Prionoglaris (Prionoglarididae). Two methods (condition-based parsimony reconstruction and common-interval genome distances) were applied to estimate the ancestral mitochondrial arrangement in Psocodea, and both provided concordant results. Specifically, the common ancestor of Psocodea retained the ancestral pancrustacean condition, and most of the gene arrangement types have originated independently from this ancestral condition. We also utilized the genomic data for phylogenetic estimation. The tree estimated from the mitochondrial genomic data was well resolved, strongly supported, and in agreement with previously estimated phylogenies. It also provided the first robust support for the family Prionoglarididae, as its monophyly was uncertain in previous morphological and molecular studies.
  • Naoki Ogawa, Kazunori Yoshizawa
    Arthropod Structure and Development 47 416 - 422 1873-5495 2018 [Refereed][Not invited]
     
    The gain of foldable wings is regarded as one of the key innovations enabling the present-day diversity of neopteran insects. Wing folding allows compact housing of the wings and shields the insect body from damage. Wing-fixing systems have evolved in some insects, probably to increase the durability of the shielding function by the wings. Bark lice (Psocodea) are known to possess a unique wing-to-wing repose coupling system, but a detailed morphological and evolutionary study of this system is lacking. In this study, we examined this repose coupling structure by SEM in 32 species including representatives of all three suborders of bark lice (Trogiomorpha, Troctomorpha and Psocomorpha). We concluded that the repose wing-coupling apparatus independently evolved twice within Psocodea. In Trogiomorpha, the apparatus is located on the subcostal vein of the forewing and is composed of elongated rib-like structures. In Troctomorpha and Psocomorpha, in contrast, the repose coupling structure is located on the radius vein of the forewing and is formed by a swollen vein. These morphological and developmental differences in the repose coupling structures also provide phylogenetic information at different systematic levels.
  • SUGIMOTO Miwa, OGAWA Naomi, YOSHIZAWA Kazunori
    Arthropod Structure & Development 2018 [Refereed][Not invited]
  • Alberto Sendra, Kazunori Yoshizawa, Rodrigo Lopes Ferreira
    Subterranean Biology 27 53 - 73 2018 [Refereed][Not invited]
  • Satoshi D. Ohdachi, Kazunori Yoshizawa, Yasushi Takada, Masaharu Motokaw, Masahiro A. Iwasa, Satoru Arai, Junji Moribe, Yasushi Uematsu, Eiichi Sakai, Takashi Tateishi, Hong-Shik Oh, Gohta Kinoshita
    Mammal Study 43 2018 [Refereed][Not invited]
  • Naoki Ogawa, Kazunori Yoshizawa
    ORGANISMS DIVERSITY & EVOLUTION 17 (3) 521 - 530 1439-6092 2017/09 [Refereed][Not invited]
     
    Morphological and behavioral characters are frequently examined for comparative studies. Unlike morphology, a single behavioral trait is difficult to subdivide as multiple characters, even when achieved by many evolutionary changes. Therefore, when similar behavioral traits evolved independently among closely related taxa, their distinction is difficult. Almost all members of the suborder Auchenorrhyncha (Insecta: Hemiptera) possess a jumping ability that uses metathoracic muscles, and this behavioral trait has been regarded as a synapomorphy. In this study, the anatomical observations of metathoracic muscles revealed that highly elaborated jumping ability was gained independently within the suborder, although the evolution of jumping ability might have been initiated at their common ancestor. Our results provide an example of identifying a true evolutionary pathway by dissecting a behavioral character into mechanical elements.
  • Kazunori Yoshizawa, Naoki Ogawa, Christopher H. Dietrich
    JOURNAL OF ZOOLOGICAL SYSTEMATICS AND EVOLUTIONARY RESEARCH 55 (3) 199 - 207 0947-5745 2017/08 [Refereed][Not invited]
     
    The phylogenetic placement of the moss bugs (Insecta: Hemiptera: Coleorrhyncha) has been highly controversial. Many apparent morphological apomorphies support the close relationship between Coleorrhyncha and Heteroptera (=true bugs). However, a recent phylogenomic study strongly supported a sister group relationship between Coleorrhyncha and Auchenorrhyncha (planthoppers, leafhoppers, treehoppers, spittlebugs, and cicadas). To test these two alternative hypotheses, we examined the fore- and hindwing base structure of the only known extant macropterous species of Coleorrhyncha using binocular and confocal laser scanning microscopes and analyzed the data selected from the wing base phylogenetically. When full morphological data including the wing base characters were analyzed, the sister group relationship between Coleorrhyncha+Heteroptera was supported, agreeing with previous consensus based on morphology. In contrast, when only wing base characters were analyzed separately, the clade Coleorrhyncha+Auchenorrhyncha was recovered, in agreement with the result from the phylogenomic study. The membranous condition of the proximal median plate in the forewing was identified as a potential synapomorphy of the latter grouping, and the absence of the tegula was excluded as a potential synapomorphy of Coleorrhyncha and Heteroptera.
  • Kazunori Yoshizawa, Kevin P. Johnson, Izumi Yao, Jose Arturo Casasola Gonzalez, Emilie Bess, Alfonso Neri Garcia Aldrete
    BIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY 121 (3) 501 - 513 0024-4066 2017/07 [Refereed][Not invited]
     
    The causes underlying disjunct distributions are of major importance in biogeography. Arcto-Tertiary relict biotas in the temperate northern hemisphere, which typically show disjunct distributions between Asia and the Nearctic region, are widely known, but often poorly understood examples of disjunct distributions. The distributional pattern of the barklouse genus Trichadenotecnum is an example of an Arcto-Tertiary relict, with centres of species diversity in Asia and Central America. We evaluated the potential causes of this disjunct distribution in Trichadenotecnum using a molecular phylogeny, divergence dating and ancestral area reconstruction. Phylogenetic analysis identified three separate clades of New World Trichadenotecnum, whereas all other groups were distributed in the Old World. Ancestral area and dating analyses recovered three independent events of trans-Beringian dispersal in the Oligocene to Miocene (27-15 Mya). The formation of two disjunct centres of diversity can be explained by the restriction of distributional areas to temperate refugia during the Quaternary glaciations (2.5-0.02 Mya). The South American Trichadenotecnum appeared to have arisen from two independent dispersal events in the Miocene (19-9 Mya). These estimated dispersal dates are much older than the generally assumed date (Pleistocene: c. 3 Mya) for the formation of the Isthmus of Panama.
  • K. Yoshizawa, M. Souza-Silva, R. L. Ferreira
    NEOTROPICAL ENTOMOLOGY 46 (2) 169 - 174 1519-566X 2017/04 [Refereed][Not invited]
     
    A new barklouse species belonging to the Trichadenotecnum s.str. clade (Psocodea: "Psocoptera": Psocidae) was described from southeastern Brazil (Minas Gerais state). This locality is highly isolated (about 3400 km) from the closest known distributional range of this clade. It also represents the southernmost distributional record of Trichadenotecnum s.str. Phylogenetic placement of the species and biogeography of Neotropical Trichadenotecnum were briefly discussed. Trichadenotecnum punctipenne New, 1972 described from Brazil was synonymized with Trichadenotecnum pardus Badonnel, 1955.
  • Kazunori Yoshizawa, Charles Lienhard
    ZOOTAXA 4079 (2) 229 - 245 1175-5326 2016/02 [Refereed][Not invited]
     
    The diversity of feeding apparatuses in insects far exceeds that observed in any other animal group. Consequently, tracking mouthpart innovation in insects is one of the keys toward understanding their diversification. In hemipteroid insects (clade Paraneoptera or Acercaria: lice, thrips, aphids, cicadas, bugs, etc.), the transition from chewing to piercing-and-sucking mouthparts is widely regarded as the turning point that enabled hyperdiversification of the Hemiptera, the fifth largest insect order. However, the transitional process from chewing to piercing-and-sucking in the Paraneoptera was hitherto completely unknown. In this paper, we report a well preserved mid Cretaceous amber fossil of the paraneopteran insect family Archipsyllidae and describe it as Mydiognathus eviohlhoffae gen. et sp. n. This species has elongate mandibles and styliform laciniae similar to Hemiptera but retains functional chewing mouthparts. A number of morphological characters place the Archipsyllidae as the sister group of the thrips plus hemipterans, which strongly suggests that the mouthparts of M. eviohlhoffae represent a transitional condition from primitive chewing to derived piercing-and-sucking mouthparts. The clade composed of Archipsyllidae, thrips, and hemipterans is here named Pancondylognatha, a new supra-ordinal taxon. Based on newly obtained information, we also assess the monophyly of the Paraneoptera, which was called into question by recent phylogenomic analyses. A phylogenetic analysis that includes Mydiognathus strongly supports the monophyly of the Paraneoptera.
  • YOSHIZAWA Kazunori
    Insecta matsumurana, new series The Research Faculty of Agriculture, Hokkaido University 72 95 - 100 0020-1804 2016 [Refereed][Not invited]
     
    The male of Manicapsocus alettae Smithers was recorded for the first time and its genital morphology was described and illustrated. This also represented the first report of the male of the genus. The highly specialized phallosome of M. alettae supported its close relationship with Epitroctes Mockford and Nothoentomum Badonnel, as previously suggested from general and female genital characters.
  • Kazunori Yoshizawa, Izumi Yao, Charles Lienhard
    MOLECULAR PHYLOGENETICS AND EVOLUTION 94 358 - 364 1055-7903 2016/01 [Refereed][Not invited]
     
    Trichadenotecnum is one of the most diverse genera among the non-parasitic members of Psocodea (Insecta: "Psocoptera"). The genus shows a world-wide distribution (excluding the Australian Region, where only one introduced species is known) with its center of diversity in southern to eastern Asia. Several species groups had been proposed for this large genus based on morphology, but their validity and phylogenetic relationships are still unclear because of great morphological diversity in the genitalia, systematically the most relevant character. In this study, we estimated the molecular phylogeny of the Old World species of Trichadenotecnum based on extensive taxon sampling. As a result, the monophyly of morphology-based species groups was very strongly supported in most cases. However, two groups were recovered as non-monophyletic, which had been inadequately defined on the basis of plesiomorphies or convergences of genital characters. First, the monophyly of the sexpunctatum group was not supported because the medium group was found to be embedded within this group. The simpler genitalia observed in the medium group were considered to be derived from the more complicated genitalia present in the sexpunctatum group. Second, the monophyly of the majus group was not supported for two reasons: (1) It was divided into two distant clades which initially had been united on the basis of convergent similarities of the male genitalia. (2) Two species groups were revealed to be embedded within the main clade of the majus group; the initial separation of these groups had been based on reversals to the ancestral genital condition. (C) 2015 Elsevier Inc. All rights reserved.
  • Jing-Fu Tsai, Shin-ichi Kudo, Kazunori Yoshizawa
    BMC EVOLUTIONARY BIOLOGY 15 258  1471-2148 2015/11 [Refereed][Not invited]
     
    Background: Maternal care (egg-nymph guarding behavior) has been recorded in some genera of Acanthosomatidae. However, the origin of the maternal care in the family has remained unclear due to the lack of phylogenetic hypotheses. Another reproductive mode is found in non-caring species whose females smear their eggs before leaving them. They possess pairs of complex organs on the abdominal venter called Pendergrast's organ (PO) and spread the secretion of this organ onto each egg with their hind legs, which is supposed to provide a protective function against enemies. Some authors claim that the absence of PO may be associated with the presence of maternal care. No study, however, has tested this hypothesis of a correlated evolution between the two traits. Results: We reconstructed the molecular phylogeny of the subfamily Acanthosomatinae using five genetic markers sequenced from 44 species and one subspecies with and without maternal care. Eight additional species from the other two acanthosomatid subfamilies were included as outgroups. Our results indicated that maternal care has evolved independently at least three times within Acanthosomatinae and once in the outgroup species. Statistical tests for correlated evolution showed that the presence of maternal care is significantly correlated with the secondary loss or reduction of PO. Ancestral state reconstruction for the node of Acanthosoma denticaudum (a non-caring species in which egg smearing with developed POs occurs) and A. firmatum (a caring species with reduced POs) suggested egg smearing was still present in their most recent common ancestor and that maternal care in A. firmatum has evolved relatively recently. Conclusions: We showed that maternal care is an apomorphic trait that has arisen multiple times from the presence of PO within the subfamily Acanthosomatinae. The acquisition of maternal care is correlated with the reduction or loss of PO, which suggests an evolutionary trade-off between the two traits resulting from physiological costs. This prediction also implies that presence of maternal care can be highly expected for those groups lacking behavioral data, which invariably also lack the organ. No secondary loss of maternal care was detected in the present tree. We suggest that the loss of maternal care may be suppressed due to the vulnerability of the PO-free condition, which thus maintains maternal care.
  • Kazunori Yoshizawa, Charles Lienhard
    ZOOTAXA 3957 (4) 480 - 488 1175-5326 2015/05 [Refereed][Not invited]
     
    The genus Cryptopsocus Li, 2002 is synonymized with Trichadenotecnum Enderlein, 1909. The type species of Cryptopsocus, T. cynostigmus (Li, 2002) n. comb., is considered to be a close relative of T. marginatum New & Thornton, 1976. These species cannot be assigned to any species group previously established in Trichadenotecnum so that the marginatum species group is here proposed for them. Three new species belonging to this species group are described: T. tigrinum and T. sharkeyi from Thailand and T. sabahense from Sabah, Malaysia. The phylogenetic position of the marginatum group is discussed using morphological data.
  • Naoki Ogawa, Munetoshi Maruyama, Kazunori Yoshizawa
    ENTOMOLOGICAL SCIENCE 18 (2) 262 - 265 1343-8786 2015/04 [Refereed][Not invited]
     
    The Aetalionidae is a small family belonging to the treehopper superfamily Membracoidea (Hemiptera: Cicadomorpha). Although the wing-base morphology of Cicadomorpha was examined in detail recently, the wing base of this family has not been investigated to date. We examined morphology of the wing-base structure of Aetalionidae. Using the characters selected from the wing base, we inferred the phylogenetic placement of this family and confirmed that it belongs to the superfamily Membracoidea and is likely a sister group of the Membracidae.
  • Yoshizawa, K, Lienhard, C
    Insecta matsumurana, new series The Research Faculty of Agriculture, Hokkaido University 71 179 - 188 0020-1804 2015 [Refereed][Not invited]
     
    The alinguum species group is newly proposed in the genus Trichadenotecnum (Psocidae: Ptyctini), and diagnosis of the species group is provided. Four named species, T. alinguum Endang, Thornton & New, T. laticornutum Endang, Thornton & New, T. sibolangitense Endang & New, and T. waykananense Endang & New are assigned to this species group. Two new species of the alinguum group, T. siamense and T. triceratum, are described and T. alinguum is newly recorded from Thailand.
  • Bernhard Misof, Shanlin Liu, Karen Meusemann, Ralph S. Peters, Alexander Donath, Christoph Mayer, Paul B. Frandsen, Jessica Ware, Tomas Flouri, Rolf G. Beutel, Oliver Niehuis, Malte Petersen, Fernando Izquierdo-Carrasco, Torsten Wappler, Jes Rust, Andre J. Aberer, Ulrike Aspoeck, Horst Aspoeck, Daniela Bartel, Alexander Blanke, Simon Berger, Alexander Boehm, Thomas R. Buckley, Brett Calcott, Junqing Chen, Frank Friedrich, Makiko Fukui, Mari Fujita, Carola Greve, Peter Grobe, Shengchang Gu, Ying Huang, Lars S. Jermiin, Akito Y. Kawahara, Lars Krogmann, Martin Kubiak, Robert Lanfear, Harald Letsch, Yiyuan Li, Zhenyu Li, Jiguang Li, Haorong Lu, Ryuichiro Machida, Yuta Mashimo, Pashalia Kapli, Duane D. McKenna, Guanliang Meng, Yasutaka Nakagaki, Jose Luis Navarrete-Heredia, Michael Ott, Yanxiang Ou, Guenther Pass, Lars Podsiadlowski, Hans Pohl, Bjoern M. von Reumont, Kai Schuette, Kaoru Sekiya, Shota Shimizu, Adam Slipinski, Alexandros Stamatakis, Wenhui Song, Xu Su, Nikolaus U. Szucsich, Meihua Tan, Xuemei Tan, Min Tang, Jingbo Tang, Gerald Timelthaler, Shigekazu Tomizuka, Michelle Trautwein, Xiaoli Tong, Toshiki Uchifune, Manfred G. Walzl, Brian M. Wiegmann, Jeanne Wilbrandt, Benjamin Wipfler, Thomas K. F. Wong, Qiong Wu, Gengxiong Wu, Yinlong Xie, Shenzhou Yang, Qing Yang, David K. Yeates, Kazunori Yoshizawa, Qing Zhang, Rui Zhang, Wenwei Zhang, Yunhui Zhang, Jing Zhao, Chengran Zhou, Lili Zhou, Tanja Ziesmann, Shijie Zou, Yingrui Li, Xun Xu, Yong Zhang, Huanming Yang, Jian Wang, Jun Wang, Karl M. Kjer, Xin Zhou
    SCIENCE 346 (6210) 763 - 767 0036-8075 2014/11 [Refereed][Not invited]
     
    Insects are the most speciose group of animals, but the phylogenetic relationships of many major lineages remain unresolved. We inferred the phylogeny of insects from 1478 protein-coding genes. Phylogenomic analyses of nucleotide and amino acid sequences, with site-specific nucleotide or domain-specific amino acid substitution models, produced statistically robust and congruent results resolving previously controversial phylogenetic relationships. We dated the origin of insects to the Early Ordovician [similar to 479 million years ago (Ma)], of insect flight to the Early Devonian (similar to 406 Ma), of major extant lineages to the Mississippian (similar to 345 Ma), and the major diversification of holometabolous insects to the Early Cretaceous. Our phylogenomic study provides a comprehensive reliable scaffold for future comparative analyses of evolutionary innovations among insects.
  • Kazunori Yoshizawa, Kevin P. Johnson
    ZOOLOGICAL JOURNAL OF THE LINNEAN SOCIETY 171 (4) 716 - 731 0024-4082 2014/08 [Refereed][Not invited]
     
    The largest suborder of bark lice (Insecta: Psocodea: 'Psocoptera') is Psocomorpha, which includes over 3600 described species. We estimated the phylogeny of this major group with family-level taxon sampling using multiple gene markers, including both nuclear and mitochondrial ribosomal RNA and protein-coding genes. Monophyly of the suborder was strongly supported, and monophyly of three of four previously recognized infraorders (Caeciliusetae, Epipsocetae, and Psocetae) was also strongly supported. In contrast, monophyly of the infraorder Homilopsocidea was not supported. Based on the phylogeny, we divided Homilopsocidea into three independent infraorders: Archipsocetae, Philotarsetae, and Homilopsocidea. Except for a few cases, previously recognized families were recovered as monophyletic. To establish a classification more congruent with the phylogeny, we synonymized the families Bryopsocidae (with Zelandopsocinae of Pseudocaeciliidae), Calopsocidae (with Pseudocaeciliidae), and Neurostigmatidae (with Epipsocidae). Monophyly of Elipsocidae, Lachesillidae, and Mesopsocidae was not supported, but the monophyly of these families could not be rejected statistically, so they are tentatively maintained as valid families. The molecular tree was compared with a morphological phylogeny estimated previously. Sources of congruence and incongruence exist and the utility of the morphological data for phylogenetic estimation is evaluated. (C) 2014 The Linnean Society of London
  • Kazunori Yoshizawa, Charles Lienhard, Idris Abd Ghani
    ZOOTAXA 3835 (4) 469 - 500 1175-5326 2014/07 [Refereed][Not invited]
     
    Species of the bark louse genus Trichadenotecnum Enderlein (Insecta: Psocodea) from Peninsular Malaysia and Singapore are revised with illustrations and identification keys. Twenty species are here recognised, with four new species and ten recorded for the first time from this region, together with an unnamed species represented by a single female. The previously described species T. marginatum New & Thornton is not included because its generic assignment is questionable. Females of T. cinnamonum Endang & New, T. imrum New & Thornton and T. sibolangitense Endang, Thornton & New, and the male of T. kerinciense Endang & New are described for the first time. A new species group is defined for T. krucilense Endang, Thornton & New.
  • Yoko Matsumura, Izumi Yao, Rolf G. Beutel, Kazunori Yoshizawa
    ARTHROPOD SYSTEMATICS & PHYLOGENY 72 (2) 95 - 110 1863-7221 2014/07 [Refereed][Not invited]
     
    Phylogenetic relationships among major groups of Criocerinae were reconstructed using molecular data (mitochondrial cytochrome oxidase I and 12S rDNA, and nuclear histone 3). The monophyly of Criocerinae was consistently and robustly supported. The Lema group including Lema, Oulema and Neolema was recovered as a clade, with the latter two genera imbedded within Lema. The Lilioceris group was placed as the sister taxon of the Lema group, and the genus Crioceris was identified as the sister taxon of the Lilioceris + Lema groups. The monophyly and/or validity of Mecoprosopus Chujo, 1951 and the subgenera Lema, Petauristes Latreille, 1829, Quasilema Monros, 1960, Microlema Pic, 1932, and Bradyceris Chujo, 1951 were not confirmed. The monophyly of the subgenus Lema except for the type species L. cyanea was supported by molecular and morphological data, and we termed it the cyanella clade. The present molecular phylogeny was compared with previous concepts with respect to the validity of each genus/subgenus. A revision of several genera is necessary. Based on the phylogenic result, the character evolution of the reproductive organs was analyzed. The ancestral states of this character system were parsimoniously reconstructed. Various shapes of the spermatheca were observed in the subfamily. A convoluted spermatheca evolved once, and reversals to the ancestral state took place several times independently. An elongation of a part of the intromittent organ also occurred several times independently. The length of the male and female reproductive ducts, which are in physical contact during copulation, showed a tight positive correlation even after removing phylogenetic effects. This strongly suggests coevolution between the male and female genital length.
  • Ayaka Yamagishi, Izumi Yao, Kevin P. Johnson, Kazunori Yoshizawa
    ZOOLOGICAL SCIENCE 31 (6) 383 - 389 0289-0003 2014/06 [Refereed][Not invited]
     
    Data from gene sequences and morphological structures were collected for the gull feather lice, Saemundssonia lari, Quadraceps punctatus, and Q. ornatus, parasitizing Larus crassirostris and L. schistisagus. Saemundssonia lari was collected from both gull species, and no detectable morphological and genetic differences were found between lice collected from the two different hosts. In contrast, Q. punctatus was only collected from L. crassirostris, whereas Q. ornatus was only collected from L. schistisagus. The two Quadraceps species were genetically highly divergent, and body-size differences corresponding to the gull's body size (Harrison's rule) were also detected between them. Both Quadraceps species were collected from the interbarb of the remex or rectrix, and a match in body size between the louse and the interbarb space may be important in escape from host preening defenses. In contrast, Saemundssonia is a head louse, inhabiting the finer feathers of the head and neck, which the bird cannot preen. A close match to host body size may be less important for lice in the head microhabitat. The differences in the pattern of host-specificity between Saemundssonia and Quadraceps on the two focal host species of this study were probably due to their different microhabitat preferences. More broadly, comparisons of the gene sequences of S. lari and Q. punctatus to those from other gull hosts showed that genetically almost undifferentiated populations of both species were distributed on wide range of gull species. Frequent interspecific hybridization of gulls is one possible factor that may allow these lice to maintain gene flow across multiple host species.
  • Kazunori Yoshizawa, Rodrigo L. Ferreira, Yoshitaka Kamimura, Charles Lienhard
    CURRENT BIOLOGY 24 (9) 1006 - 1010 0960-9822 2014/05 [Refereed][Not invited]
     
    Sex-specific elaborations are common in animals and have attracted the attention of many biologists, including Darwin [1]. It is accepted that sexual selection promotes the evolution of sex-specific elaborations. Due to the faster replenishment rate of gametes, males generally have higher potential reproductive and optimal mating rates than females. Therefore, sexual selection acts strongly on males [2], leading to the rapid evolution and diversification of male genitalia [3]. Male genitalia are sometimes used as devices for coercive holding of females as a result of sexual conflict over mating [4, 5]. In contrast, female genitalia are usually simple. Here we report the reversal of intromittent organs in the insect genus Neotrogla (Psocodea: Prionoglarididae) from Brazilian caves. Females have a highly elaborate, penis-like structure, the gynosome, while males lack an intromittent organ. The gynosome has species-specific elaborations, such as numerous spines that fit species-specific pouches in the simple male genital chamber. During prolonged copulation (similar to 40-70 hr), a large and potentially nutritious ejaculate is transferred from the male via the gynosome. The correlated genital evolution in Neotrogla is probably driven by reversed sexual selection with females competing for seminal gifts. Nothing similar is known among sex-role reversed animals.
  • Yoko Matsumura, Kazunori Yoshizawa, Ryuichiro Machida, Yuta Mashimo, Romano Dallai, Marco Gottardo, Thomas Kleinteich, Jan Michels, Stanislav N. Gorb, Rolf G. Beutel
    BIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY 112 (1) 40 - 54 0024-4066 2014/05 [Refereed][Not invited]
     
    Very unusual genitalia of the species Zorotypus caudelli are described. It contains the unique configuration of two different intromittent organs, one of them strongly elongated. Hyper elongated genitalia are known in different groups of insects. Males have to accommodate these unwieldy structures in the limited spaces of the abdomen and manipulate them acutely during copulation. A crucial question is how do species with elongated genitalia cope with these requirements? To investigate this, we studied key features enabling storage, insertion, and withdrawal of the elongated genitalia. The co-existence of an elongated narrow tube and a bulky spermatophore is a highly unusual and apparently paradoxical condition. However, we demonstrate that the tube is not involved in sperm transmission, whereas the large spermatophore is transferred to females by a membranous fold of the genitalia. The movement of the spermatophore is caused by haemolymph pressure, which likely also promotes the insertion of both intromittent organs. A comparison with the genital anatomy and reproductive mode in related groups suggests that the elongated tube and its accommodating pouch is a de novo structure, and that the ancestral sperm transport via spermatophore is a preadaptive condition for the acquisition of this unusual structure. (c) 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112, 40-54.
  • Kazunori Yoshizawa, Edward L. Mockford, Kevin P. Johnson
    SYSTEMATIC ENTOMOLOGY 39 (2) 279 - 285 0307-6970 2014/04 [Refereed][Not invited]
     
    The phylogenetic relationships of bark lice and parasitic lice (Insecta: Psocodea) have been studied in a number of recent molecular phylogenetic analyses based on DNA sequences. Many of these studies have focused on the position of parasitic lice within the free-living bark lice. However, fewer such studies have examined the relationships among major groups of free-living bark lice and their implications for classification. In this study we focus on the infraorder Caeciliusetae, a large group of bark lice (similar to 1000 species) within the suborder Psocomorpha. Using sequences of two mitochondrial and two nuclear genes, we estimated the phylogeny for relationships among the five recognized families within the infraorder Caeciliusetae. Based on the results, the sister-group relationship and respective monophyly of Stenopsocidae and Dasydemellidae is strongly supported. Monophyly of the larger families Amphipsocidae and Caeciliusidae was not supported, although the causes of this were the placement of two distinct subfamilies (Paracaeciliinae and Calocaeciliinae). The monophyly of Asiopsocidae could not be tested because it was sampled only by one species. Based on these results and consideration of morphological characters, we propose a new classification for Caeciliusetae, recognizing six families: Amphipsocidae, Stenopsocidae, Dasydemellidae, Asiopsocidae, Paracaeciliidae and Caeciliusidae. We expect that this new classification will stabilize the higher-level taxonomy of this group and help to identify groups in need of further work among these insects.
  • Katrin Friedemann, Rico Spangenberg, Kazunori Yoshizawa, Rolf G. Beutel
    CLADISTICS 30 (2) 170 - 201 0748-3007 2014/04 [Refereed][Not invited]
     
    Acercaria display an unusually broad array of adhesive devices occurring on different parts of the legs. Attachment structures of all major subgroups are described and illustrated. Nineteen characters of the distal leg region were combined with a data matrix containing 99 additional morphological characters of different body parts. The results of the cladistic analysis are largely congruent with current hypotheses. Zoraptera are not retrieved as close relatives of Acercaria. The monophyly of the entire lineage and of the major subgroups Psocodea, Phthiraptera, and Hemiptera is confirmed. Our data also support the monophyly of Auchenorrhycha and a sister-group relationship between Thysanoptera and Hemiptera (Condylognatha). In contrast to other lineages of insects, the hairy type of adhesive device is present only in one group within the Acercaria (Heteroptera, Cimicomorpha). The arolium is present in the groundplan but missing in several groups (e.g. Psocodea, Cicadoidea, Aphidoidea). Pretarsal pulvilli evolved several times independently. Tarsal euplantulae and different specialized clasping devices have evolved within Phthiraptera, whereas pretarsal attachment devices are missing in this ectoparasitic group. The potential to modify pretarsal attachment devices in their structural details has probably contributed to the very successful diversification of the predominantly phytophagous Hemiptera.
  • Yoshizawa K, Ferreira R.L, Kamimura Y, Lienhard C
    The Winnower 2014 [Refereed][Invited]
  • Y. Mashimo, Y. Matsumura, R. Machida, R. Dallai, M. Gottardo, K. Yoshizawa, F. Friedrich, B. Wipfler, R. G. Beutel
    INSECT SYSTEMATICS & EVOLUTION 45 (4) 371 - 393 1399-560X 2014 [Refereed][Not invited]
     
    Zoraptera are a cryptic and enigmatic group of insects. The species diversity is lower than in almost all other groups of Hexapoda, but may be distinctly higher than presently known. Several new species were described from different regions recently. The systematic placement was discussed controversially since the group was discovered 100 years ago. Affinities with Isoptera and Psocoptera were discussed in earlier studies. A sister group relationship with Acercaria (Psocodea, Thysanoptera, Hemiptera) was proposed by W. Hennig, for the first time based on a strictly phylogenetic argumentation. More recent studies consistently suggest a placement among the "lower neopteran orders" (Polyneoptera). Close affinities to Dictyoptera were proposed and alternatively a sister group relationship with Embioptera or with Embioptera + Phasmatodea (Eukinolabia), respectively. The precise placement is still controversial and the intraordinal relationships are largely unclear. Recent transcriptome analyses tentatively suggest a clade Zoraptera + Dermaptera as sister group of all other polyneopteran orders. The oldest fossils are from Cretaceous amber. An extinct genus from this era may be the sister group of all the remaining zorapterans. The knowledge of the morphology, development and features related to the reproductive system greatly increased in recent years. The general body morphology is very uniform, whereas the genitalia differ strongly between species. This is likely due to different kinds of selection, i.e. sexual selection in the case of the genital organs. The mating pattern also differs profoundly within the order. A unique external sperm transfer occurs in Zorotypus impolitus. A species-level phylogeny and more investigations of the reproductive system should have high priority.
  • Kazunori Yoshizawa, Kevin P. Johnson
    GENETICA 141 (10-12) 491 - 499 0016-6707 2013/12 [Refereed][Not invited]
     
    While it is well known that changes in the general processes of molecular evolution have occurred on a variety of timescales, the mechanisms underlying these changes are less well understood. Parasitic lice ("Phthiraptera") and their close relatives (infraorder Nanopsocetae of the insect order Psocodea) are a group of insects well known for their unusual features of molecular evolution. We examined changes in base composition across parasitic lice and bark lice. We identified substantial differences in percent GC content between the clade comprising parasitic lice plus closely related bark lice (=Nanopsocetae) versus all other bark lice. These changes occurred for both nuclear and mitochondrial protein coding and ribosomal RNA genes, often in the same direction. To evaluate whether correlations in base composition change also occurred within lineages, we used phylogenetically controlled comparisons, and in this case few significant correlations were identified. Examining more constrained sites (first/second codon positions and rRNA) revealed that, in comparison to the other bark lice, the GC content of parasitic lice and close relatives tended towards 50 % either up from less than 50 % GC or down from greater than 50 % GC. In contrast, less constrained sites (third codon positions) in both nuclear and mitochondrial genes showed less of a consistent change of base composition in parasitic lice and very close relatives. We conclude that relaxed selection on this group of insects is a potential explanation of the change in base composition for both mitochondrial and nuclear genes, which could lead to nucleotide frequencies closer to random expectation (i.e., 50 % GC) in the absence of any mutation bias. Evidence suggests this relaxed selection arose once in the non-parasitic common ancestor of Phthiraptera + Nanopsocetae and is not directly related to the evolution of the parasitism in lice.
  • Yuta Mashimo, Kazunori Yoshizawa, Michael S. Engel, Idris Abd Ghani, Romano Dallai, Rolf G. Beutel, Ryuichiro Machida
    ZOOTAXA 3717 (4) 498 - 514 1175-5326 2013/10 [Refereed][Not invited]
     
    Three new species of the uncommonly encountered insect order Zoraptera are described and figured from Peninsular Malaysia-Zorotypus magnicaudelli sp. n., Zorotypus cervicornis sp. n., and Zorotypus impolitus sp. n. Another species from the region, identified as Zorotypus caudelli Karny, 1927, was also collected and is reevaluated herein based on new material. A brief discussion of characters used in zorapteran systematics is provided, and a key to the species of Peninsular Malaysia provided. This is the first report for the order Zoraptera from Peninsular Malaysia.
  • Luxi Liu, Kazunori Yoshizawa, Fasheng Li, Zhiqi Liu
    ZOOTAXA 3701 (4) 460 - 466 1175-5326 2013/08 [Refereed][Not invited]
     
    Atrichadenotecnum multispinosus sp. n. is described from southwestern China. Psocomesites and Clematostigma from China are discussed, with five species transferred to Atrichadenotecnum and four species placed as new synonyms. Keys to adult males and females of Atrichadenotecnum species are presented.
  • Yoko Matsumura, Ryuichiro Machida, Benjamin Wipfler, Rolf G. Beutel, Kazunori Yoshizawa
    EVOLUTION & DEVELOPMENT 15 (4) 305 - 315 1520-541X 2013/07 [Refereed][Not invited]
     
    Extreme elongation of a part of the intromittent organ, the flagellum, has occurred several times in Criocerinae (Chrysomelidae). These leaf beetles have acquired a specialized pocket to store the flagellum in the abdominal cavity, at the same time allowing a quick control of movements of this structure during copulation. We investigated the morphogenesis of the intromittent organs of species with and without a flagellum to discuss the evolutionary background of parallel evolution of novel structures. We found that the specialized pocket is formed by the invagination of an epidermal layer and a resultant rotation of the primary gonopore. Invagination itself is a well-known phenomenon in morphogenetic processes, which leads us to hypothesize that the novelty is formed by co-opting a previously acquired genetic system. A large open-space is present within the intromittent organ during the entire morphogenesis in species without a flagellum, and the invagination in the species with a flagellum grows in the corresponding area. This means that there are no physical impediments for the growth of a large pocket. In addition the sites of muscular attachments in the species with a flagellum are also different from those without it. The differentiation of muscles is completed immediately before adult emergence, which means the muscles are adjustable during the entire morphogenesis in this group. Simple modifications probably based on a co-option of previously acquired genetic systems, the potential space for adding a new element, and an adjustable factor in morphogenesis of the intromittent organ facilitate the parallel evolution of the extreme elongation.
  • Satoshi D. Ohdachi, Kazunori Yoshizawa, Ilkka Hanski, Kuniko Kawai, Nikolai E. Dokuchaev, Boris I. Sheftel, Alexei V. Abramov, Igor Moroldoevs, Atsushi Kawahara
    MAMMAL STUDY 37 (4) 281 - 297 1343-4152 2012/12 [Not refereed][Not invited]
     
    Phylogenetic analysis was conducted for various populations of the Sorex minutissimus-S. yukonicus complex based on mitochondrial gene (cytochrome b and/or the control region) sequences. Sorex minutissimus was divided into some monophyletic groups in Eurasia; it was divided into 2 main groups, eastern and western Eurasian clades, based on combined data of the cytochrome b and the control region. Monophyly of shrews from Hokkaido-Sakhalin, Primorye, Mongolia-Transbaikalia, southeastern Finland was strongly supported respectively in most analyses. Sorex yukonicus was phylogenetically close to S. minutissimus in eastern Siberia. Some shrews from western and central Siberia were included in the clade of southeastern Finland Also, most shrews from central-northern Finland and Norway made a clade close to but different from the southeastern Finland clade. This finding suggests that Fennoscandian shrews might consist of individuals which were recolonised from various refugia after the Last Glacial Maximum. Nucleotide diversity of shrews from Hokkaido and Alaska was low. Three regional groups in Kamchatka-Sakha, Sakhalin, and Mongolia-Transbaikalia tended to have medium nucleotide diversity. In contrast, shrews from Cisbaikalia-western Siberia and Fennoscandia had high nucleotide diversity. The S. minutissimus-S. hosonoi group appears to have experienceed a quit different biogeographic history from two shrews with similar ranges, the S. caecutiens-S. hosonoi group and S. tundrensis.
  • 吉澤 和徳
    化学と生物 日本農芸化学会 ; 1962- 50 (11) 840 - 843 0453-073X 2012/11/01 [Not refereed][Not invited]
  • YOSHIZAWA KAZUNORI, MOCKFORD EDWARD L
    Insecta Matsumurana New Ser Research faculty of agriculture, Hokkaido University 68 (68) 133 - 141 0020-1804 2012/10 [Not refereed][Not invited]
     
    Symbiopsocus hastatus was described based only on two males from KunashiriIsland. Here we redescribe the species based on the specimens collected throughout the Japanese Archipelago, with first description of females. Morphological examinations showed that the species is closely related to Mecampsis multimacularis Li from China. The type species of Mecampsis is endemic to Chile and, except for 10 Chinese species described recently, distributional ranges of all the species are restricted to South America and the Greater Antilles. These distributional areas of Mecampsis are widely disjunct and thus assignments of the Chinese species to this genus are highly questionable. Rather, judging from the phallosomal and hypandrial apomorphies, closer relationship between the Chinese Mecampsis with Symbiopsocus is likely. Therefore, all the Chinese species of Mecampsis are here transferred to Symbiopsocus. Symbiopsocus sturmi (Badonnel) is transferred back to its original genus, Psocomesites.
  • Kazunori Yoshizawa, Mutsumi Wagatsuma
    ENTOMOLOGICAL SCIENCE 15 (4) 408 - 421 1343-8786 2012/10 [Not refereed][Not invited]
     
    The infraorder Cicadomorpha is a monophyletic group of the order Hemiptera, suborder Auchenorrhyncha, and is composed of three superfamilies: Cercopoidea (spittle bugs), Cicadoidea (cicadas) and Membracoidea (leafhoppers and treehoppers). Phylogenetic relationships among the superfamilies have been highly controversial morphologically and molecularly, but recent molecular phylogenetic analyses provided support for Cercopoidea + Cicadoidea. In this study, we examined morphology of the wing base structure in Cicadomorpha and tested the previous phylogenetic hypotheses using the characters selected from the wing base. As a result, a sister-group relationship between Cicadoidea and Cercopoidea was supported by three synapomorphies (presence of a projection posterior to the anterior notal wing process, presence of a novel notal process anterior to the posterior notal wing process, presence of a novel sclerite between the distal median plate and the base of anal vein). The present study provides the first unambiguous and prominent morphological support for Cicadoidea + Cercopoidea.
  • Atsushi Mizukoshi, Kevin P. Johnson, Kazunori Yoshizawa
    PARASITOLOGY 139 (12) 1614 - 1629 0031-1820 2012/10 [Refereed][Not invited]
     
    Lice are obligate parasites of mammals and birds and have become an important model for studies of host-parasite co-evolution and co-phylogenetics. Population genetic and phylogeographic studies represent an important bridge between microevolution and co-phylogenetic patterns. We examine co-phylogeographic patterns in sika deer and their parasitic lice. Co-phylogeographic patterns in deer and lice were evaluated using homologous regions of mitochondrial COI sequences. The phylogeographic breaks recovered for deer populations matched those of previous studies. Comparisons of the phylogeographic tree topology for deer lice with that of their hosts revealed a significant level of congruence. However, comparisons of genetic distances between deer and lice suggested that one of the estimated co-divergence events is more likely a recent host switch. Taking into account genetic divergence, there is not strong evidence for complete phylogeographic co-divergence between deer and their parasitic lice. However, mitochondrial phylogenies only track genetic structure of female lineages, and the incongruence between deer and louse phylogeography may be explained by louse migration mediated by male deer. Morphological analysis of head shape variation based on an elliptic Fourier descriptor showed that overall morphological variation contained phylogenetic signal, suggesting that in general morphology of these lice evolves congruent to population history.
  • Yoko Matsumura, Kazunori Yoshizawa
    JOURNAL OF MORPHOLOGY 273 (5) 507 - 518 0362-2525 2012/05 [Refereed][Not invited]
     
    Extremely elongated intromittent organs are found in a wide range of taxa, especially among insects. This phenomenon is generally thought to result from sexual selection, but it is predicted that limited storage space in the body cavity and the difficulty of using the elongated organs should have constrained the evolution of extreme elongation, neutralizing any selective advantage. Therefore, in groups with long intromittent organs, features that overcome these constraints may have evolved or coevolved together with intromittent organ elongation. Using a comparative morphological approach and outgroup comparisons, we identified potential constraints and key novelties that may neutralize such constraints in the leaf beetle subfamily Criocerinae. Observations of the internal sac structure throughout Criocerinae were performed. Comparing the results with preceding studies from outgroups, a ground plan of the criocerine internal sac was constructed. Our analysis also identified specific features that are always correlated with extreme elongation: the rotation of whole internal-sac sclerites and the possession of a pocket in which to store the elongated flagellum. The pocket is thought to be formed by the rotation of the sclerites, markedly altering internal sac shape from the criocerine ground plan. Onlythe clades that have acquired this derived state contain species with an elongated flagellum that distinctly exceeds the median lobe length. It is presumed that these character correlations evolved independently three times. The detected character correlations corroborate the hypothesis that there are latent adaptive constraints for the evolution of extremely elongated intromittent organs. The constraints may have been neutralized by the alteration from the criocerine ground plan resulting in the formation of a storage pocket. In conclusion, deviation from the criocerine ground plan isconsidered to be the evolutionary innovation that neutralized the latent adaptive constraints of flagellum elongation in the subfamily Criocerinae. J. Morphol., 2012. (c) 2011 Wiley Periodicals, Inc.
  • Sebastian Buesse, Philipp von Grumbkow, Susanne Hummel, Deep Narayan Shah, Ram Devi Tachamo Shah, Jingke Li, Xueping Zhang, Kazunori Yoshizawa, Sonja Wedmann, Thomas Hoernschemeyer
    PLOS ONE 7 (5) e38132  1932-6203 2012/05 [Refereed][Not invited]
     
    Unusual biogeographic patterns of closely related groups reflect events in the past, and molecular analyses can help to elucidate these events. While ample research on the origin of disjunct distributions of different organism groups in the Western Paleartic has been conducted, such studies are rare for Eastern Palearctic organisms. In this paper we present a phylogeographic analysis of the disjunct distribution pattern of the extant species of the strongly cool-adapted Epiophlebia dragonflies from Asia. We investigated sequences of the usually more conserved 18 S rDNA and 28 S rDNA genes and the more variable sequences of ITS1, ITS2 and CO2 of all three currently recognised Epiophlebia species and of a sample of other odonatan species. In all genes investigated the degrees of similarity between species of Epiophlebia are very high and resemble those otherwise found between different populations of the same species in Odonata. This indicates that substantial gene transfer between these populations occurred in the comparatively recent past. Our analyses imply a wide distribution of the ancestor of extant Epiophlebia in Southeast Asia during the last ice age, when suitable habitats were more common. During the following warming phase, its range contracted, resulting in the current disjunct distribution. Given the strong sensitivity of these species to climatic parameters, the current trend to increasing global temperatures will further reduce acceptable habitats and seriously threaten the existences of these last representatives of an ancient group of Odonata.
  • Kazunori Yoshizawa
    SYSTEMATIC ENTOMOLOGY 37 (1) 2 - 6 0307-6970 2012/01 [Not refereed][Not invited]
  • Lu-Xi Liu, Kazunori Yoshizawa, Fa-Sheng Li, Zhi-Qi Liu
    ZOOKEYS 203 (203) 27 - 46 1313-2989 2012 [Refereed][Not invited]
     
    A review of species of the genus Neopsocopsis Badonnel, 1936 is presented. Four species are redescribed, viz. N. hirticornis (Reuter, 1893), N. quinquedentata (Li & Yang, 1988), N. profunda (Li, 1995), and N. flavida (Li, 1989), as well as the description of one new species, N. convexa sp. n. Seven new synonymies are proposed as follows: Pentablaste obconica Li syn. n. and P. clavata Li syn. n. of N. hirticornis, P. tetraedrica Li syn. n. of N. longiptera, Neoblaste schizopetala Li syn. n. and Nb. flavae Li syn. n. of N. profunda, Blastopsocidus pini Li syn. n. and P. lanceolata Li syn. n. of N. flavida. N. hirticornis (Reuter, 1893) is recorded from Japan and China for the first time, and N. longiptera Vishnyakova, 1986 is newly recorded from China. Illustrated keys to adult males and females are presented.
  • SEKIMOTO SHIGEYUKI, YOSHIZAWA KAZUNORI
    Insecta Matsumurana New Ser Research Faculty of Agriculture, Hokkaido University 67 (67) 1 - 22 0020-1804 2011/10 [Not refereed][Not invited]
     
    Japanese species of Osmylus were recognized: O. (O.) pryeri McLachlan, 1875, O. (O.) hyalinatus McLachlan, 1875 and O. (O.) decoratus Nakahara, 1914. One species of the subgenus Plesiosmylus Makarkin, O. (P.) tessellatus McLachlan, 1875 was recognized. All Japanese species of Osmylus were redescribed and illustrated but O. kisoensis Iwara, 1928, which is known only from a larva to date, was not treated here. A key to the Japanese species of the genus is provided. In Appendix, two new combinations were proposed for two Chinese species, O. atomatus (Yang, 1988) and O. zheanus (Yang & Liu, 2001)(both originally assigned in the genus Plethosmylus Kruger).
  • Vincent S. Smith, Tom Ford, Kevin P. Johnson, Paul C. D. Johnson, Kazunori Yoshizawa, Jessica E. Light
    BIOLOGY LETTERS 7 (5) 782 - 785 1744-9561 2011/10 [Refereed][Not invited]
     
    For modern lineages of birds and mammals, few fossils have been found that predate the Cretaceous-Palaeogene (K-Pg) boundary. However, molecular studies using fossil calibrations have shown that many of these lineages existed at that time. Both birds and mammals are parasitized by obligate ectoparasitic lice (Insecta: Phthiraptera), which have shared a long coevolutionary history with their hosts. Evaluating whether many lineages of lice passed through the K-Pg boundary would provide insight into the radiation of their hosts. Using molecular dating techniques, we demonstrate that the major louse suborders began to radiate before the K-Pg boundary. These data lend support to a Cretaceous diversification of many modern bird and mammal lineages.
  • Stephen L. Cameron, Kazunori Yoshizawa, Atsushi Mizukoshi, Michael F. Whiting, Kevin P. Johnson
    BMC GENOMICS 12 394  1471-2164 2011/08 [Refereed][Not invited]
     
    Background: The gene composition, gene order and structure of the mitochondrial genome are remarkably stable across bilaterian animals. Lice (Insecta: Phthiraptera) are a major exception to this genomic stability in that the canonical single chromosome with 37 genes found in almost all other bilaterians has been lost in multiple lineages in favour of multiple, minicircular chromosomes with less than 37 genes on each chromosome. Results: Minicircular mt genomes are found in six of the ten louse species examined to date and three types of minicircles were identified: heteroplasmic minicircles which coexist with full sized mt genomes (type 1); multigene chromosomes with short, simple control regions, we infer that the genome consists of several such chromosomes (type 2); and multiple, single to three gene chromosomes with large, complex control regions (type 3). Mapping minicircle types onto a phylogenetic tree of lice fails to show a pattern of their occurrence consistent with an evolutionary series of minicircle types. Analysis of the nuclear-encoded, mitochondrially-targetted genes inferred from the body louse, Pediculus, suggests that the loss of mitochondrial single-stranded binding protein (mtSSB) may be responsible for the presence of minicircles in at least species with the most derived type 3 minicircles (Pediculus, Damalinia). Conclusions: Minicircular mt genomes are common in lice and appear to have arisen multiple times within the group. Life history adaptive explanations which attribute minicircular mt genomes in lice to the adoption of blood-feeding in the Anoplura are not supported by this expanded data set as minicircles are found in multiple non-blood feeding louse groups but are not found in the blood-feeding genus Heterodoxus. In contrast, a mechanist explanation based on the loss of mtSSB suggests that minicircles may be selectively favoured due to the incapacity of the mt replisome to synthesize long replicative products without mtSSB and thus the loss of this gene lead to the formation of minicircles in lice.
  • Kazunori Yoshizawa
    SYSTEMATIC ENTOMOLOGY 36 (3) 377 - 394 0307-6970 2011/07 [Not refereed][Not invited]
     
    Phylogenetic relationships among the winged orders of Polyneoptera [Blattodea, Dermaptera, Embiodea (=Embioptera), Isoptera, Mantodea, Orthoptera, Phasmatodea, Plecoptera and Zoraptera] were estimated based on morphological data selected from the hindwing base structure. Cladistic analyses were carried out using hindwing base data alone and in combination with other, more general, morphological data. Both datasets resulted in similar trees and recovered the monophyly of Polyneoptera. Deepest phylogenetic relationships among the polyneopteran orders were not confidently estimated, but the monophyly of Mystroptera (= Embiodea + Zoraptera), Orthopterida (= Orthoptera + Phasmatodea) and Dictyoptera (= Blattodea + Mantodea + Isoptera) was supported consistently. In contrast, placements of Plecoptera and Dermaptera were unstable, although independent analysis of the wing base data supported their sister-group relationship with two nonho-moplasious synapomorphies (unique conditions in the ventral basisubcostale, and in the articulation between the antemedian notal wing process and first axillary sclerite). Results from the combined wing base plus general morphology data were consistent, even if the wingless orders Grylloblattodea and Mantophasmatodea were included in the analysis. Generally, trees obtained from the present analyses were concordant with the results from other morphological and molecular analyses, but Isoptera were placed inappropriately to be the sister of Blattodea + Mantodea by the inclusion of the wing base data, probably as a result of morphological regressions of the order.
  • Issei Ohshima, Kazunori Yoshizawa
    MOLECULAR PHYLOGENETICS AND EVOLUTION 59 (2) 469 - 476 1055-7903 2011/05 [Refereed][Not invited]
     
    We investigated the utility of indel data for genealogical and population genetic analyses using the Tpi intron of the leaf mining moth Acrocercops transecta (Insecta: Lepidoptera). Genealogical analyses revealed that indel data were less homoplasious than DNA sequence data and that indel data contained a sufficient signal to provide a high resolution tree that was highly congruent with the tree estimated from DNA sequences. Although some conflicts were identified in the distributions of multi-residue indels, such conflicts were especially useful for the unambiguous detection of recombinations. For the first time, we adopted a Bayesian clustering method for indel characters to infer genetic structure of the moth. We concluded that indel characters have the potential to be a powerful tool in the analysis of population genetics and population structure as well as in the detection of gene flow. (C) 2011 Elsevier Inc. All rights reserved.
  • Yoshizawa Kazunori
    タクサ : 日本動物分類学会誌 日本動物分類学会 0 (30) 17 - 29 1342-2367 2011/02/20 [Not refereed][Not invited]
     
    Problems in direct optimization method as implemented in the software POY were reviewed. Alignment of highly variable regions using POY is problematic because it likely provides artificial maximization of pseudo-homology within the alignment. If the variable regions are aligned with conserved regions, then the bias could be emerged in two ways. When the conserved regions contain only weak phylogenetic signal, then stochastic similarities within the variable regions strongly affect to the final alignment. This type of bias can be emerged even if the variable regions are aligned independently...
  • Kazunori Yoshizawa, Emilie Bess, Kevin P. Johnson
    INVERTEBRATE SYSTEMATICS 25 (2) 81 - 90 1445-5226 2011 [Not refereed][Not invited]
     
    The systematic status of Kaindipsocinae (formerly Kaindipsocini) is revised based on morphology of the male terminalia and on molecular data. The genera Clematostigma, Lasiopsocus and Tanystigma are newly assigned to this subfamily. The 'Blaste' lunulata species-group is also placed within Kaindipsocinae and is probably closest to Kaindipsocus. Both morphological and molecular data provide strong support for monophyly of Kaindipsocinae and molecular data support a sister relationship between this subfamily and the rest of Psocidae.
  • YOSHIZAWA KAZUNORI
    Insecta Matsumurana New Ser Research Faculty of Agriculture, Hokkaido University 66 (66) 11 - 36 0020-1804 2010/10 [Not refereed][Not invited]
  • Kazunori Yoshizawa, Charles Lienhard
    ARTHROPOD SYSTEMATICS & PHYLOGENY 68 (2) 181 - 195 1863-7221 2010/06 [Not refereed][Not invited]
     
    The taxonomy, fossil record, phylogeny, and systematic placement of the booklouse family Liposcelididae (Insecta: Psocodea: 'Psocoptera') were reviewed. An apterous specimen from lower Eocene, erroneously identified as Embidopsocus eocenicus Nel et al., 2004 in the literature, is recognized here as an unidentified species of Liposcelis Motsehulsky. 1852. It represents the oldest fossil of the genus. Phylogenetic relationships within the family presented in the recent literature were re-analyzed, based on a revised data matrix. The resulting tree was generally in agreement with that originally published. but the most basal dichotomy between the fossil taxon Cretoscelis Grimaldi & Engel, 2006 and the rest of the Liposcelididae was not supported. Monophyly of Liposcelis with respect to Troglotroctes Lienhard. 1996 is highly questionable, but the latter genus is retained because of lack of conclusive evidence. Paraphyly of Psocoptera (i.e., closer relationship between Liposcelididae and parasitic lice) is now well established, based on both morphological and molecular data. Monophyly of Phthiraptera is questionable, but support for the 'Polyphyly of Lice Hypothesis. is still not definitive. A checklist of valid names of all presently recognized Liposcelididae taxa (10 genera, 200 species) is also included with information on their geographical distribution. Because monophyly of the subfamily Embidopsocinae is highly questionable, we list the genera alphabetically without adopting the usual subdivision into two subfamilies.
  • Kazunori Yoshizawa, Kevin P. Johnson
    MOLECULAR PHYLOGENETICS AND EVOLUTION 55 (3) 939 - 951 1055-7903 2010/06 [Refereed][Not invited]
     
    Recent molecular phylogenetic analyses of 18S rDNA have indicated that parasitic lice (order Phthiraptera) are possibly polyphyletic. These analyses recovered one of the parasitic louse suborders, Amblycera, as the sister group to the free-living booklouse family Liposcelididae. We further tested this hypothesis using DNA sequences from five genes: nuclear 18S rDNA, Histone 3, and wingless and mitochondrial 16S rDNA and COL Combined analyses of these five genes provided reasonably strong support for the Amblycera + Liposcelididae clade, supporting the polyphyly of lice hypothesis. To explore the robustness of this result, we examined the phylogenetic signal contained in each gene independently (except for wingless, which could not be readily amplified in many target taxa). Analyses of each gene separately and in various combinations with other genes revealed that clear signal supporting Amblycera + Liposcelididae only existed in the 18S data, although no analysis supported monophyly of parasitic lice. Nevertheless, combined analyses of all genes provided stronger support for this relationship than that obtained from 18S data alone. The increase in support for this clade was mostly explained by the stabilization of other parts of the tree and potentially inappropriate substitution modeling. These findings demonstrate that the increased support values provided by combined data set does not always indicate corroboration of the hypothesis. (C) 2010 Elsevier Inc. All rights reserved.
  • Issei Ohshima, Kazunori Yoshizawa
    MOLECULAR ECOLOGY 19 (10) 2106 - 2119 0962-1083 2010/05 [Refereed][Not invited]
     
    Recently diverged populations often exhibit incomplete reproductive isolation, with a low level of gene flow continuing between populations. Previous studies have shown that, even under a low level of gene flow, genetic divergence between populations can proceed at the loci governing local adaptation and reproductive isolation but not at other neutral loci. A leaf-mining moth, Acrocercops transecta, consists of Juglans- and Lyonia-associated host races. The two host races differ in host preferences of ovipositing females and in larval adaptation to host plants but mate readily in the laboratory, producing fertile hybrids. The Juglans and Lyonia races are often sympatric in the wild, implying that gene introgression could occur in nature between the two host races. We tested this hypothesis by combining phylogenetic analyses with coalescent simulations, focusing on mitochondrial genes (COI and ND5) and the nuclear Tpi, Per and Ldh genes located on the Z-chromosome. The mitochondrial genes clearly distinguished the Lyonia race from the Juglnas race, whereas the Tpi, Per and Ldh genealogies did not reflect the two host races. Coalescent simulations indicated gene flow at the three Z-linked genes in both directions, whereas there was no introgression in the mitochondrial genes. The lack of introgression in mitochondrial genes suggests that female host preference is the primary force leading to the bifurcation of maternally inherited loci. Thus, the results show that a low level of gene flow coupled with the inflexible female host preference differentiates histories of divergence between maternally and biparentally inherited genes in this host race system.
  • Kazunori Yoshizawa
    SYSTEMATIC ENTOMOLOGY 35 (2) 199 - 206 0307-6970 2010/04 [Not refereed][Not invited]
  • Yoko Matsumura, Kazunori Yoshizawa
    BIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY 99 (3) 512 - 520 0024-4066 2010/03 [Not refereed][Not invited]
     
    Complicated genital structures are widely known in many animals. It is assumed that an increasing morphological complexity of genital structures would also increase the difficulty of inserting and withdrawing the structures. We examined the insertion and withdrawal mechanisms of extremely elongated genitalia in the Japanese leaf beetle, Lema (Lema) coronata. Insertion and withdrawal processes are completed quickly. Investigation of genital morphology showed that there are no protractor or retractor muscles attached to the elongated part. Instead, the elongated part is tightly enveloped by a membrane. Because of the close fit between the elongated part and its surrounding membrane, eversion of the membrane allows for insertion of the elongated part, and retraction of the membrane induces withdrawal of the elongated part. This surrounding membrane makes it possible to store the median ejaculatory guide (MEG) in a small genital cavity, with the basal portion of the elongated part extending internally against the entrance. This unusual character state is observed only in members of the subgenus Lema, which also have the elongated part. This condition can be considered to be a preadaptation for extreme elongation of genitalia. (C) 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 512-520.
  • Kazunori Yoshizawa, Alfonso Neri Garcia Aldrete
    STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT 45 (1) 61 - 66 0165-0521 2010 [Not refereed][Not invited]
     
    A new species of the genus Trichadenotecnum, T. shawi n. sp., belonging in the roesleri species group, is here described and illustrated. The species exhibits several morphological features that are valuable to revise the ancestral condition of the genus and the definition of the species group. Based on morphological observations, the phylogenetic position of the species within the roesleri group is also discussed.
  • YOSHIZAWA KAZUNORI
    Insecta Matsumurana New Ser Research Faculty of Agriculture, Hokkaido University 65 (65) 149 - 155 0020-1804 2009/08 [Not refereed][Not invited]
     
    A new genus and species of the tribe Ptyctini, Kimunpsocus takumai, was described based on the specimens collected in Hokkaido, Japan. Morphologically, Kimunpsocus is apparently a member of the tribe Ptyctini and is similar to Loensia in the forewing markings and venation. However, independence of this new genus was supported morphologically and molecularly.
  • Tomoya Ninomiya, Kazunori Yoshizawa
    SYSTEMATIC ENTOMOLOGY 34 (2) 334 - 345 0307-6970 2009/04 [Not refereed][Not invited]
     
    Homology of the wing base structure in the Odonata is highly controversial, and many different interpretations of homology have been proposed. In extreme cases, two independent origins of insect wings have been suggested, based on comparative morphology between the odonate and other pterygote wing bases. Difficulties in establishing homology of the wing base structures between Odonata and other Pterygota result mainly from their extreme differences in morphology and function. In the present paper, we establish homology of the wing base structures between Neoptera, Ephemeroptera and Odonata using highly conservative and unambiguously identifiable characters (the basal wing hinge and subcostal veins) as principal landmarks. Homology of the odonate wing base structure with those of Ephemeroptera and Neoptera can be identified reliably. Based on this interpretation, the ancestral condition of the insect wing base structure is discussed.
  • Kazunori Yoshizawa, Alfonso Neri Garcia Aldrete, Edward L. Mockford
    ZOOLOGICAL JOURNAL OF THE LINNEAN SOCIETY 153 (4) 651 - 723 0024-4082 2008/08 [Not refereed][Not invited]
     
    The taxonomy of the New World species of the genus Trichadenotecnum is revised. A total of 44 species, including 29 new species, were treated. These species are classified into 12 monophyletic species groups, eight of them newly proposed here. Two species previously assigned to Trichadenotecnum, T. pichincha New & Thornton and T. sylvaticum Turner, are recognized as not belonging to this genus. Phylogenetic relationships among 16 previously and presently proposed species groups are estimated based on a data matrix of 58 morphological characters. Trees from these analyses support monophyly of Trichadenotecnum and the proposed species groups. The New World species were divided into three major clades. Based on the phylogenetic hypothesis and distributional pattern of the species groups, the biogeographical history of the New World Trichadenotecnum is discussed. (C) 2008 The Linnean Society of London.
  • YOSHIZAWA KAZUNORI
    Insecta Matsumurana New Ser RESEARCH FACULTY OF AGRICULTURE, HOKKAIDO UNIVERSITY 64 (64) 23 - 34 0020-1804 2008/03 [Not refereed][Not invited]
  • Kazunori Yoshizawa, Kevin P. Johnson
    MOLECULAR PHYLOGENETICS AND EVOLUTION 46 (2) 547 - 559 1055-7903 2008/02 [Not refereed][Not invited]
     
    We evaluated the higher level classification within the family Psocidae (Insecta: Psocodea: 'Psocoptera') based on combined analyses of nuclear 18S, Histone 3, wingless and mitochondrial 12S, 16S and COI gene sequences. Various analyses (inclusion/exclusion of incomplete taxa and/or rapidly evolving genes, data partitioning, and analytical method selection) all provided similar results, which were generally concordant with relationships inferred using morphological observations. Based on the phylogenetic trees estimated for Psocidae, we propose a revised higher level classification of this family, although uncertainty still exists regarding some aspects of this classification. This classification includes a basal division into two subfamilies, 'Amphigerontiinae' (possibly paraphyletic) and Psocinae. The Amphigerontiinae is divided into the tribes Kaindipsocini (new tribe), Blastini, Amphigerontini, and Stylatopsocini. Psocinae is divided into the tribes 'Ptyctini' (probably paraphyletic), Psocini, Atrichadenotecnini (new tribe), Sigmatoneurini, Metylophorini, and Thyrsophorini (the latter includes the taxon previously recognized as Cerastipsocini). We examined the evolution of symmetric/asymmetric mate genitalia over this tree and found this character to be quite homoplasious. (C) 2007 Elsevier Inc. All rights reserved.
  • Kazunori Yoshizawa, Kevin P. Johnson
    MOLECULAR PHYLOGENETICS AND EVOLUTION 46 (2) 547 - 559 1055-7903 2008/02 [Refereed][Not invited]
     
    We evaluated the higher level classification within the family Psocidae (Insecta: Psocodea: 'Psocoptera') based on combined analyses of nuclear 18S, Histone 3, wingless and mitochondrial 12S, 16S and COI gene sequences. Various analyses (inclusion/exclusion of incomplete taxa and/or rapidly evolving genes, data partitioning, and analytical method selection) all provided similar results, which were generally concordant with relationships inferred using morphological observations. Based on the phylogenetic trees estimated for Psocidae, we propose a revised higher level classification of this family, although uncertainty still exists regarding some aspects of this classification. This classification includes a basal division into two subfamilies, 'Amphigerontiinae' (possibly paraphyletic) and Psocinae. The Amphigerontiinae is divided into the tribes Kaindipsocini (new tribe), Blastini, Amphigerontini, and Stylatopsocini. Psocinae is divided into the tribes 'Ptyctini' (probably paraphyletic), Psocini, Atrichadenotecnini (new tribe), Sigmatoneurini, Metylophorini, and Thyrsophorini (the latter includes the taxon previously recognized as Cerastipsocini). We examined the evolution of symmetric/asymmetric mate genitalia over this tree and found this character to be quite homoplasious. (C) 2007 Elsevier Inc. All rights reserved.
  • Systematic positions of the species currently placed under the holding genus Psocidus s.l. described by Okamoto (Psocodea: 'Psocoptera': Psocidae).
    Insecta Matsumurana, New Series 64 23 - 34 2008 [Not refereed][Not invited]
  • Takema Fukatsu, Ryuichi Koga, Wendy A. Smith, Kohjiiro Tanaka, Naruo Nikoh, Kayoko Sasaki-Fukatsu, Kazunori Yoshizawa, Colin Dale, Dale H. Clayton
    APPLIED AND ENVIRONMENTAL MICROBIOLOGY 73 (20) 6660 - 6668 0099-2240 2007/10 [Refereed][Not invited]
     
    The current study focuses on a symbiotic bacterium found in the slender pigeon louse, Columbicola columbae (Insecta: Phthiraptera). Molecular phylogenetic analyses indicated that the symbiont belongs to the gamma subdivision of the class Proteobacteria and is allied to Sodalis glossinidius, the secondary symbiont of tsetse flies (Glossina spp.) and also to the primary symbiont of grain weevils (Sitophilus spp.). Relative-rate tests revealed that the symbiont of C. columbae exhibits accelerated molecular evolution in comparison with the tsetse fly symbiont and the weevil symbiont. Whole-mount in situ hybridization was used to localize the symbiont and determine infection dynamics during host development. In first- and second-instar nymphs, the symbionts were localized in the cytoplasm of oval bacteriocytes that formed small aggregates on both sides of the body cavity. In third-instar nymphs, the bacteriocytes migrated to the central body and were finally located in the anterior region of the lateral oviducts, forming conspicuous tissue formations called ovarial ampullae. In adult females, the symbionts were transmitted from the ovarial ampullae to developing oocytes in the ovarioles. In adult males, the bacteriocytes often disappeared without migration. A diagnostic PCR survey of insects collected from Japan, the United States, Australia, and Argentina detected 96.5% (109/113) infection, with a few uninfected male insects. This study provides the first microbial characterization of a bacteriocyte-associated symbiont from a chewing louse. Possible biological roles of the symbiont are discussed in relation to the host nutritional physiology associated with the feather-feeding lifestyle.
  • Emilie C. Bess, Kazunori Yoshizawa
    ENTOMOLOGICAL SCIENCE 10 (3) 279 - 290 1343-8786 2007/09 [Not refereed][Not invited]
     
    The genera Ptycta Enderlein, 1925, and Copostigma Enderlein, 1903, are defined as a monophyletic complex based on the morphology of the male terminalia. Ptycta is redefined as those species of the Copostigma-Ptycta complex with forewing veins Rs+M fused for a short distance. Two new species of Ptycta from Japan are described, P. recava sp. nov. and P. johnsoni sp. nov., increasing the number of Japanese species to four, along with P. parvidentata Tsutsumi, 1964, and P. micromaculata Thornton, Lee, and Chui, 1972. Distributional information and illustrations of each species, and a key to Japanese species of Ptycta are included.
  • Kazunori Yoshizawa, Tomoya Ninomiya
    ARTHROPOD STRUCTURE & DEVELOPMENT 3 36 (3) 277 - 279 1467-8039 2007/09 [Refereed][Not invited]
     
    We revised the homology of wing base structure in Ephemeroptera (Insecta: Pterygota) proposed by Willkommen and Hornschemeyer in a recent issue of Arthropod Structure and Development. The first free sclerite (s1) in Ephemeroptera should be homologized with a part of the first axillary sclerite (1Ax) of Neoptera, together with the second free sclerite, whereas the authors recognized s1 as a detached part of the anterior notal wing process. The fifth free sclerite of Ephemeroptera should be homologized with the median notal wing process of Neoptera, rather than it being homologous with a part of 1 Ax in Neoptera, as the authors postulated. Hypothesized secondary fusion of the axillary sclerites in Ephemeroptera and Odonata proposed by the authors is premature, because the basal phylogeny of Pterygota is still poorly understood, and an alternative interpretation of morphological evolution (i.e., that undifferentiated axillary sclerites represent the ground plan of Pterygota) can also be drawn from the Ephemeroptera + Neoptera hypothesis. (c) 2007 Elsevier Ltd. All rights reserved.
  • Kazunori Yoshizawa
    SYSTEMATIC ENTOMOLOGY 32 (2) 197 - 204 0307-6970 2007/04 [Not refereed][Not invited]
     
    The order Zoraptera is one of the most enigmatic insect groups. Its phylogenetic position is far from settled, and more than ten different placements have been discussed since the insects were first discovered. This problem is also difficult to resolve with molecular studies because of the unusual characteristics of zorapteran 18S rDNA sequences, which are the most widely used genetic markers for the estimation of the deep phylogeny of insects. In this study, the wing base structures of Zoraptera and various potential sister taxa were examined. Numbers of unique modifications were detected in the wing base structure of Zoraptera, and six were also observed in the wing base of Embiodea (Embioptera, Embiidina; webspinners). No possible synapomorphies supporting the other relationships were detected. This is the second unambiguous morphological synapomorphy providing strong evidence for the phylogenetic position of Zoraptera.
  • Shigeyuki Sekimoto, Kazunori Yoshizawa
    ENTOMOLOGICAL SCIENCE 10 (1) 81 - 86 1343-8786 2007/03 [Not refereed][Not invited]
     
    Suhpalacsa iriomotensis sp. nov. (Neuroptera: Ascalaphidae: Ascalaphinae), is described from Iriomotejima Island, Japan. This new species can be easily distinguished from other species of the genus by the unique ventrolateral prominences of the male ectoproct. This is the first record of the genus Suhpalacsa from Japan. The morphology of the male ectoproct and the monophyly of Suhpalacsa are discussed briefly.
  • YOSHIZAWA KAZUNORI, LIENHARD CHARLES, THAPA VASANT KUMAR
    Insecta Matsumurana New Ser FACULTY OF AGRICULTURE, HOKKAIDO UNIVERSITY 63 (63) 1 - 33 0020-1804 2007/02 [Not refereed][Not invited]
     
    Nepalese species of the genus Trichadenotecnum Enderlein are revised. A total of 16species are recognized of which six are newly described: Trichadenotecnum isseii n. sp.,T. suwai n. sp., and T. danieli n. sp. belonging to the spiniserrulum group, T. nepalense n.sp. and T. depitarense n. sp. belonging to the newly established distinctum group, and T.malickyi n. sp. belonging to the newly established digitatum group. T. distinctum Datta,originally described from India, and T. sclerotum New, originally described from Bhutan,are newly recorded from Nepal. Four possibly undescribed species are also recognized;however, these species are not named here because they are represented only by femalespecimens. One of these species belongs to the newly established longimucronatumgroup, which was formerly recognized as an independent genus, Conothoracalis Li, herenewly synonymized with Trichadenotecnum. The holotype male of T. masoni New andthe allotype male of T. godavarense New are considered to be "chimeras" composed ofgeneral body parts of one species and male terminalia of the other species. Therefore,terminal parts of the holotype male of T. masoni are here excluded from the holotype.Several new generic combinations and species group assignments are also proposed fornon-Nepalese species.
  • Kazunori Yoshizawa, Courtenay N. Smithers
    Records of the Australian Museum 58 (3) 411 - 415 2201-4349 2006/11/29 [Refereed][Not invited]
     
    Trichadenotecnum enderleini (Roesler, 1943) is the only native species of the genus known from Australia to date. However, its exact systematic position is not firmly fixed mainly due to the lack of information on taxonomically and phylogenetically important morphological characters. In this study, T. enderleini is examined morphologically to clarify further its systematic position. The forewing markings and male terminal structures clearly show that the species lacks all apomorphies of Trichadenotecnum Enderlein, 1909 and is thus only distantly related to the genus. Therefore, T. enderleini is excluded from Trichadenotecnum. Ptycta floresensis Endang, Thornton & New, 2002 from Indonesia is considered to be the closest relative of T. enderleini, and T. enderleini is consequently transferred to the genus Ptycta Enderlein, 1925. As a result, Australia is excluded from the distributional range of Trichadenotecnum, except for T. circularoides Badonnel, 1955 considered to be introduced. © Copyright Australian Museum, 2006.
  • Kazunori Yoshizawa, Courtenay N. Smithers
    RECORDS OF THE AUSTRALIAN MUSEUM 58 (3) 411 - 415 0067-1975 2006/11 [Not refereed][Not invited]
     
    Trichadenotecnum enderleini (Roesler, 1943) is the only native species of the genus known from Australia to date. However, its exact systematic position is not firmly fixed mainly due to the lack of information on taxonomically and phylogenetically important morphological characters. In this study, T enderleini is examined morphologically to clarify further its systematic position. The forewing markings and male terminal structures clearly show that the species lacks all apomorphies of Trichadenotecnum Enderlein, 1909 and is thus only distantly related to the genus. Therefore, T enderleini is excluded from Trichadenotecnum. Ptycta floresensis Endang, Thornton & New, 2002 from Indonesia is considered to be the closest relative of T enderleini, and T enderleini is consequently transferred to the genus Ptycta Enderlein, 1925. As a result, Australia is excluded from the distributional range of Trichadenotecnum, except for T. circularoides Badonnel, 1955 considered to be introduced.
  • Kayoko Sasaki-Fukatsu, Ryuichi Koga, Naruo Nikoh, Kazunori Yoshizawa, Shinji Kasai, Minoru Mihara, Mutsuo Kobayashi, Takashi Tomita, Takerna Fukatsu
    APPLIED AND ENVIRONMENTAL MICROBIOLOGY 72 (11) 7349 - 7352 0099-2240 2006/11 [Refereed][Not invited]
     
    The symbiotic bacteria associated with the stomach disc, a large aggregate of bacteriocytes on the ventral side of the midgut, of human body and head lice were characterized. Molecular phylogenetic analysis of 16S rRNA gene sequences showed that the symbionts formed a distinct and well-defined clade in the Gammaproteobacteria. The sequences exhibited AT-biased nucleotide composition and accelerated molecular evolution. In situ hybridization revealed that in nymphs and adult males, the symbiont was localized in the stomach disc, while in adult females, the symbiont was not in the stomach disc but in the lateral oviducts and the posterior pole of the oocytes due to female-specific symbiont migration. We propose the designation "Candidatus Riesia pediculicola" for the louse symbionts.
  • K Yoshizawa, KP Johnson
    SYSTEMATIC ENTOMOLOGY 31 (2) 350 - 361 0307-6970 2006/04 [Not refereed][Not invited]
     
    Lice (Insecta: Phthiraptera) have long been considered to compose a monophyletic group of insects on the basis of external morphological characteristics. However, a recent phylogenetic analysis of 18S rDNA sequences suggested that 'Phthiraptera' have arisen twice within the order Psocoptera (booklice and barklice). The external features of lice are highly specialized to a parasitic lifestyle, and convergence may be frequent for such characters. To provide a further test between traditional and recent molecular-based phylogenetic hypotheses, a phylogenetic analysis of lice and relatives based on morphological characters that are independent from the selective pressures of a parasitic lifestyle is needed. Here, we examined the morphology of the male phallic organ in lice and relatives ('Psocoptera': suborders Troctomorpha and Psocomorpha) and detected some novel modifications that were stable within each group and useful for higher level phylogenetic reconstruction. Phylogenetic analysis based on these characters provided a concordant result with the 18S-based phylogeny. In particular, the apomorphic presence of articulations between the basal plate, mesomere and ventral plate (= sclerite on the permanently everted endophallus) is observed consistently throughout the psocid families Pachytroctidae and Liposcelididae and the louse suborder Amblycera, providing support for a clade composed of these three groups, although possible homoplasy was detected in some Ischnocera. This is the first study to provide morphological support for the polyphyly of lice.
  • K Yoshizawa, C Lienhard, KP Johnson
    ZOOLOGICAL JOURNAL OF THE LINNEAN SOCIETY 146 (2) 287 - 299 0024-4082 2006/02 [Not refereed][Not invited]
     
    Phylogenetic relationships among extant families in the suborder Trogiomorpha (Insecta: Psocodea: 'Psocoptera') were inferred from partial sequences of the nuclear 18S rDNA and Histone 3 and mitochondrial 16S rDNA genes. Analyses of these data produced trees that largely supported the traditional classification; however, monophyly of the infraorder Psocathropetae (= Psyllipsocidae + Prionoglarididae) was not recovered. Instead, the family Psyllipsocidae was recovered as the sister taxon to the infraorder Atropetae (= Lepidopsocidae + Trogiidae + Psoquillidae), and the Prionoglarididae was recovered as sister to all other families in the suborder. Character states previously used to diagnose Psocathropetae are shown to be plesiomorphic. The sister group relationship between Psyllipsocidae and Atropetae was supported by two morphological apomorphies: the presence of a paraproctal anal spine and an anteriorly opened phallosome. Based on these sequence data and morphological observations, we propose a new classification scheme for the Trogiomorpha as follows: infraorder Prionoglaridetae (Prionoglarididae), infraorder Psyllipsocetae (Psyllipsocidae), infraorder Atropetae (Lepidopsocidae, Trogiidae, Psoquillidae). (c) 2006 The Linnean Society of London.
  • Ohshima, I, K Yoshizawa
    MOLECULAR PHYLOGENETICS AND EVOLUTION 38 (1) 231 - 240 1055-7903 2006/01 [Refereed][Not invited]
     
    Insect herbivores such as gall formers and leaf miners are often highly specialized and adapted to their respective natal host plants. Due to the specialization and adaptation, it is presumed that host shifts readily occur among closely related plant species. Leaf-mining moths, the Acrocercops leueophaea complex, consist of three species, A. leucophaea, A. defigurata, and A. transecta. Larvae of all the species of the complex feed on Juglandaceae plants, but A. leucophaea and A. transecta are also associated with an Ericaceae plant, which is quite distantly related to Juglandaceae. Such a host utilization as in this species complex is very rare among phytophagous insects. In the present study, we estimate the history of host shifts by reconstructing the phylogeny of the A. leucophaea complex using molecular data (partial sequence of mitochondrial COI, 12S rDNA, and ND5). Parsimony and maximum likelihood analyses indicated that the common ancestor of the A. leucophaea complex used Juglandaceae only, and that the association with Ericaceae has evolved in A. leucophaea and A. transecta independently. Parametric bootstrap analysis also supported multiple origins of the association with Ericaceae in this complex. These results imply that there are ecological and biochemical factors that promote host shifting between Juglandaceae and Ericaceae despite the two families being not closely related. (c) 2005 Elsevier Inc. All rights reserved.
  • K Yoshizawa, C Lienhard, AB Idris
    REVUE SUISSE DE ZOOLOGIE 112 (4) 831 - 848 0035-418X 2005/12 [Not refereed][Not invited]
     
    The genus Podopterocus Banks has been known only from male specimens and has been characterized by the thickened first antennal flagellomere and the expanded hind tibia. The genus Dinopsocus Banks has been characterized by the thickened first antennal flagellomere. Detailed morphological examinations and brief molecular-based identification (only for one species) suggest that both taxa are congeneric. Based on characters of female terminalia and fore wing venation, both taxa are furthermore considered to be closely related to the genus Sigmatoneura Enderlein. The discovery of two new species and observations on Sigmatoneura spp. reveal the limited significance of the diagnostic characters of Podopterocus and Dinopsocus. Therefore we consider both Podopterocus and Dinopsocus as junior synonyms of Sigmatoneura. Monophyly of the genus Sigmatoneura, including Podopterocus, Dinopsocus, and the subgenus Longifolia Li, is well supported by the unique fore wing venation of females and by sexually dimorphic fore wing venation and coloration. Two new species, Sigmatoneura kakisayap sp. n. and Sigmatoneura lemahsayap sp. n., which would be classified under Podopterocus or Dinopsocus by the previous generic definitions, are described. Sigmatoneura longicornis comb. n., the type species of Podopterocus, is redescribed and transferred to Sigmatoneura and the female of this species is described for the first time. Dinopsocus atratus, the type species of Dinopsocus, is synonymized with S. longicornis. Sigmatoneura semicolorata comb. n. is redescribed and transferred from Dinopsocus to Sigmatoneura.
  • YOSHIZAWA KAZUNORI
    Insecta Matsumurana New Ser 北海道大学 62 (62) 1 - 44 0020-1804 2005/12 [Not refereed][Not invited]
     
    Adult integumental morphology of the suborder Psocomorpha (Psocodea:'Psocoptera') was examined, and homologies and transformation series of charactersthroughout the suborder and Psocoptera were discussed. These examinations formed thebasis of the recent morphology-based cladistic analysis of the Psocomorpha (Yoshizawa,2002, Zool. J. Linn. Soc. 136: 371–400).
  • K Yoshizawa, KP Johnson
    MOLECULAR PHYLOGENETICS AND EVOLUTION 37 (2) 572 - 580 1055-7903 2005/11 [Refereed][Not invited]
     
    The order Zoraptera (angel insects) is one of the least known insect groups, containing only 32 extant species. The phylogenetic position of Zoraptera is poorly understood, but it is generally thought to be closely related to either Paraneoptera (hemipteroid orders: booklice, lice, thrips, and bugs), Dictyoptera (blattoid orders: cockroaches, termites, and mantis), or Embioptera (web spinners). We inferred the phylogenetic position of Zoraptera by analyzing nuclear 18S rDNA sequences, which we aligned according to a secondary structure model. Maximum likelihood and Bayesian analyses both supported a close relationship between Zoraptera and Dictyoptera with relatively high posterior probability. The 18S sequences of Zoraptera exhibited several unusual properties: (1) a dramatically increased substitution rate, which resulted in very long branches; (2) long insertions at helix E23; and (3) modifications of secondary structures at helices 12 and 18. (c) 2005 Elsevier Inc. All rights reserved.
  • A report on the faunal survey of insects and arachnids living on the rainforest canopy of the Malay Peninsula.
    Serangga 10 37 - 48 2005 [Not refereed][Not invited]
  • SATOYOSHI Ayako, KABAYA Hajime, HAGIWARA Ko, TANIYAMA Hiroyuki, YOSHIZAWA Kazunori, TUJI Masayoshi, HAGIWARA Kasturo, MURAMATSU Yasukazu, ASAKAWA Mitsuhiko
    Japanese Journal of Zoo and Wildlife Medicine 日本野生動物医学会 9 (2) 79 - 83 1342-6133 2004/09 [Not refereed][Not invited]
     
    Because alien macaques (probably, Macaca mulatta) occur in the southern part of the Boso Peninsula, Japan, there is a possibility that their parasites and/or other pathogens may infect Japanese macaques (M.fuscata). To verify the possibility, this study was conducted mainly on parasites that included protozoan, helminth and arthropod, and other pathogens including Rickettsia and the Borna virus of 50 free ranging Japanese macaques in the peninsula. Consequently, 5 arthropod and nematode parasite species common to the Japanese macaque, namely Pedicinus obtusus, P.eurygaster, Streptopharagus ...
  • KP Johnson, K Yoshizawa, VS Smith
    PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES 271 (1550) 1771 - 1776 0962-8452 2004/09 [Refereed][Not invited]
     
    A major fraction of the diversity of insects is parasitic, as herbivores, parasitoids or vertebrate ectoparasites. Understanding this diversity requires information on the origin of parasitism in various insect groups. Parasitic lice (Phthiraptera) are the only major group of insects in which all members are permanent parasites of birds or mammals. Lice are classified into a single order but are thought to be closely related to, or derived from, book lice and bark lice (Psocoptera). Here, we use sequences of the nuclear 18S rDNA gene to investigate the relationships among Phthiraptera and Psocoptera and to identify the origins of parasitism in this group (termed Psocodea). Maximum-likelihood (ML), Bayesian ML and parsimony analyses of these data indicate that lice are embedded within the psocopteran infraorder Nanopsocetae, making the order Psocoptera paraphyletic (i.e. does not contain all descendants of a single common ancestor). Furthermore, one family of Psocoptera, Liposcelididae, is identified as the sister taxon to the louse suborder Amblycera, making parasitic lice (Phthiraptera) a polyphyletic order (i.e. descended from two separate ancestors). We infer from these results that parasitism of vertebrates arose twice independently within Psocodea, once in the common ancestor of Amblycera and once in the common ancestor of all other parasitic lice.
  • K Yoshizawa
    SYSTEMATIC ENTOMOLOGY 29 (3) 383 - 394 0307-6970 2004/07 [Not refereed][Not invited]
     
    Phylogenetic relationships among species groups of Trichadenotecnum were inferred based on morphology and the partial sequences of five gene regions (mitochondrial 12S rDNA, 16S rDNA, cytochrome oxidase I, NADH dehydrogenase subunit 5 and nuclear 18S rDNA). All analyses supported the monophyly of Trichadenotecnum and all previously proposed species groups, except that T. circularoides was excluded from the spiniserrulum group. To examine the phylogenetic usefulness of morphological data, the morphological characters used in the construction of an earlier taxonomic system for Trichadenotecnum were mapped parsimoniously on the molecular tree. As a result: (1) commonly used forewing marking features (sparsely or extensively spotted) are considered to be very homoplastic and less informative of higher-level phylogenetic relationships; (2) a broadly expanded epiproct lobe is considered to be independently evolved at least two or three times, and a detailed morphological re-examination allows recognition of these convergent structures; (3) the short ventral valve of gonapophyses independently evolved at least three or four times, although this character was used initially to diagnose the spiniserrulum group.
  • Y Shinozaki, K Yoshizawa, K Murata, T Shiibashi, J Kimura, S Maruyama, Y Hayama, H Yoshida, S Nogami
    JOURNAL OF VETERINARY MEDICAL SCIENCE 66 (3) 333 - 335 0916-7250 2004/03 [Not refereed][Not invited]
     
    A species of sucking louse, Neohaematopinus callosciuri, was found for the first time in Japan. The species was found on an invasive species of squirrel, Pallas squirrel, Callosciurus etythraeus, in the Kamakura district, Kanagawa Prefecture, Japan. A total of 52 lice were obtained from 22 of 104 squirrels captured. The lice were about three times more prevalent in male squirrels than in females and were detected most frequently in the winter. As N. callosciuri has never been reported on wild animals in Japan, this species probably was introduced into Japan along with their host, Pallas squirrels.
  • Y Shinozaki, T Shiibashi, K Yoshizawa, K Murata, J Kimura, S Maruyama, Y Hayama, H Yoshida, S Nogami
    MEDICAL AND VETERINARY ENTOMOLOGY 18 (1) 61 - 63 0269-283X 2004/03 [Not refereed][Not invited]
     
    The squirrel Callosciurus erythraeus (Pallas) (Rodentia: Sciuridae) was intentionally introduced to Japan in 1935 and has become established throughout much of the country. Although they live mainly in forests, Pallas squirrels come into gardens and are frequently fed by people or kept as pets, so their ectoparasites could be of potential medical as well as veterinary importance. During 2001-2003 we conducted the first ectoparasite survey of Pallas squirrels in Japan. From 105 C. erythraeus captured in Kamakura District of Kanagawa Prefecture on Honshu Island, three types of ectoparasite were found: 52 specimens of the sucking louse Neohaematopinus callosciuri Johnson (Anoplura: Haematopinidae), 26 fleas Ceratophyllus (Monopsyllus) anisus Rothschild (Siphonaptera: Ceratophyllidae) and four nymphs of the tick Haemaphysalis flava Neumann (Acari: Ixodidae) on 22, 13 and one squirrels, respectively. Evidently in Japan C. erythraeus carries relatively few ectoparasite species; this may be a contributory factor to their invasive success. Further investigations are needed to assess risks of zoonotic transmission of plague or murine typhus by C. anisus, of louse-borne typhus by N. callosciuri and of tularaemia and especially Japanese spotted fever (Rickettsia japonica) by H. flava.
  • Y Shinozaki, K Yoshizawa, K Murata, T Shiibashi, J Kimura, S Maruyama, Y Hayama, H Yoshida, S Nogami
    JOURNAL OF VETERINARY MEDICAL SCIENCE 66 (3) 333 - 335 0916-7250 2004/03 [Refereed][Not invited]
     
    A species of sucking louse, Neohaematopinus callosciuri, was found for the first time in Japan. The species was found on an invasive species of squirrel, Pallas squirrel, Callosciurus etythraeus, in the Kamakura district, Kanagawa Prefecture, Japan. A total of 52 lice were obtained from 22 of 104 squirrels captured. The lice were about three times more prevalent in male squirrels than in females and were detected most frequently in the winter. As N. callosciuri has never been reported on wild animals in Japan, this species probably was introduced into Japan along with their host, Pallas squirrels.
  • Systematics of Trichadenotecnum (Psocoptera: Psocidae) in Hong Kong
    Publicaciones Especiales del Instituto de Biologia, Universidad Nacional Autonoma de Mexico 20 121 - 149 2004 [Not refereed][Not invited]
  • K Yoshizawa, KP Johnson
    MOLECULAR PHYLOGENETICS AND EVOLUTION 29 (1) 102 - 114 1055-7903 2003/10 [Refereed][Not invited]
     
    Phthiraptera (chewing and sucking lice) and Psocoptera (booklice and barklice) are closely related to each other and compose the monophyletic taxon Psocodea. However, there are two hypotheses regarding their phylogenetic relationship: (1) monophyletic Psocoptera is the sister group of Phthiraptera or (2). Psocoptera is paraphyletic, and Liposcelididae of Psocoptera is the sister group of Phthiraptera. Each hypothesis is supported morphologically and/or embryologically, and this problem has not yet been resolved. In the present study, the phylogenetic position of Phthiraptera was examined using mitochondrial 12S and 16S rDNA sequences, with three methods of phylogenetic analysis. Results of all analyses strongly supported the close relationship between Phthiraptera and Liposcelididae. Results of the present analyses also provided some insight into the elevated rate of evolution in mitochondrial DNA (mtDNA) in Phthiraptera. An elevated substitution rate of mtDNA appears to originate in the common ancestor of Phthiraptera and Liposcelididae, and directly corresponds to an increased G + C content. Therefore, the elevated substitution rate of mtDNA in Phthiraptera and Liposcelididae appears to be directional. A high diversity of 12S rDNA secondary structure was also observed in wide range of Phthiraptera and Liposcelididae, but these structures seem to have evolved independently in different clades. (C) 2003 Elsevier Science (USA). All rights reserved.
  • K Yoshizawa, T Saigusa
    ACTA ZOOLOGICA 84 (1) 33 - 40 0001-7272 2003/01 [Not refereed][Not invited]
     
    Morphology of the head and mouthparts in Psocoptera was investigated, and revised interpretations for clypeus and maxilla were proposed. The convex plate in the frontal region of the head capsule is the postclypeus, rather than the frons; the galea is clearly differentiated from the stipes and the origin of the stipito-lacinial muscle is partly shifted from the stipes to the base of the galea; the cardo is completely fused with the stipes without any suture or sulcus. Brief discussions on the evolution of piercing and sucking mouthparts and on the phylogeny of Paraneoptera were provided, based on these revised interpretations.
  • Entomological Science 6 301 - 308 2003 [Not refereed][Not invited]
  • K Yoshizawa
    ZOOLOGICAL JOURNAL OF THE LINNEAN SOCIETY 136 (3) 371 - 400 0024-4082 2002/11 [Not refereed][Not invited]
     
    Phylogenetic relationships among all 24 families of suborder Psocomorpha (Insecta: Psocodea: 'Psocoptera') are inferred based on adult morphology. Monophyly of Psocomorpha is strongly supported by six autapomorphies. The presently accepted four infraorders - Psocetae, Homilopsocidea, Epipsocetae and Caeciliusetae - are regarded as monophyletic, but Archipsocidae and Hemipsocidae, previously assigned to Homilopsocidea and Psocetae, respectively, are regarded as the basalmost clades of the suborder. Based on the results of the cladistic analysis, a higher classification for Psocomorpha is proposed. Six infraorders (two new - Archipsocetae, Hemipsocetae - and the four aforementioned) are recognized. Four new superfamilies are recognized within Homilopsocidea: Elipsocoidea, Lachesilloidea, Pseudocaecilioidea and Peripsocoidea. Two superfamilies are recognized within Caeciliusetae: Asiopsocoidea and Caeciliusoidea. Descriptions of taxa above family level are provided. (C) 2002 The Linnean Society of London.
  • YOSHIZAWA KAZUNORI
    Insecta matsumurana. Series entomology. New series : journal of the Faculty of Agriculture Hokkaido University 北海道大学 59 29 - 38 0020-1804 2002 [Not refereed][Not invited]
     
    YOSHIZAWA, K. 2002. Type specimens of Psocoptera described by H. Okamoto in Hokkaido University Insect Collection. Ins. matsum. n. s. 59: 29-38. Psocid species described by Hanjiro Okamoto are listed. Type specimens of 20 species of the altogether 34 Okamoto's psocid species are confirmed to exist in the Hokkaido University Insect Collection.
  • Yoshizawa Kazunori
    Insecta matsumurana. Series entomology. New series : journal of the Faculty of Agriculture Hokkaido University 北海道大学 58 1 - 25 0020-1804 2001 [Not refereed][Not invited]
  • K Yoshizawa, T Saigusa
    SYSTEMATIC ENTOMOLOGY 26 (1) 1 - 13 0307-6970 2001/01 [Not refereed][Not invited]
     
    Phylogenetic relationships among three paraneopteran clades (Psocodea, Hemiptera and Thysanoptera) were analysed based on the morphology of forewing base structure, Monophyly of Paraneoptera was supported by nine autapomorphies, monophyly of Condylognatha (= Thysanoptera + Hemiptera) by two autapomorphies, monophyly of Thysanoptera by five autapomorphies and monophyly of Hemiptera by one autapomorphy. Thus, (Psocodea + (Thysanoptera + Hemiptera)) were proposed to be the phylogenetic relationships within Paraneoptera. A homoplastic similarity of the third axillary sclerite was observed between Thysanoptera and Heteroptera, and a possible evolutionary factor providing this homoplasy was discussed. The present analysis also suggested a monophyletic Auchenorrhyncha, and reduction of the proximal median plate was considered as an autapomorphy of this dade.
  • K Yoshizawa
    INVERTEBRATE TAXONOMY 15 (2) 159 - 204 0818-0164 2001 [Not refereed][Not invited]
     
    The genus Trichadenotecnum Enderlein, 1909 is redefined as a monophyletic group. The following 22 Japanese species are treated and classified into five species-groups except one species not placed in a group: sexpunctatum- group - T. al bum, sp. nov., T. incognitum Roesler, 1939; medium-group - T. takahashii, sp. nov., T. mixtum, sp. nov., T. magnomixtum, sp. nov., T. amamiense, sp. nov., T. okinawense, sp. nov., T. yonaguniense, sp. nov., T. kumejimense, sp. nov., T. latebrachium, sp. nov., T. pseudomedium, sp. nov., T. yaeyamense, sp. nov.; alexanderae-group - T. alexanderae Sommerman, 1948, T. cast um Betz, 1983, T. sexpunctellum (Enderlein, 1907); majus-group - T. yamatomajus, sp. nov., T. nothoapertum, sp. nov.; spiniserrulum-group - T. falx, sp. nov., T. furcalingum, sp. nov., T. pardidum Thornton, 1961, T. circularoides Badonnel, 1955; incertae sedis - T. fuscipennis, sp. nov. The monophyly and phylogenetic relationships of these species-groups are discussed. A key to Japanese species of Trichadenotecnum is given.
  • 皇居の動物相調査で得られたチャタテムシ目昆虫
    国立科学博物館専報 36 29 - 34 2000 [Not refereed][Not invited]
  • YOSHIZAWA Kazunori
    Entomological science Entomological Society of Japan 3 (4) 669 - 674 1343-8786 2000 [Not refereed][Not invited]
     
    Mepleres suzukii (Okamoto, 1910)(Psocoptera : Pseudocaeciliidae) was redescribed. Mepleres Enderlein, 1926 had been incorrectly treated by some authors as an invalid synonym of Pseudoscottiella Badonnel, 1946. Thus, Pseudoscottiella was appropriately treated here as a junior synonym of Mepleres. Meniscopsocus Li, 1993 was newly synonymized with Mepleres. All new species combinations resulting from these taxonomic treatments were listed. Within Mepleres, African and Oriental-Pacific-Australian species groups had been recognized, with Mep. suzukii assigned to the latter group. However, monophyly of the latter group could not be justified, whereas the African group was regarded as monophyletic. A key to Japanese Mepleres was provided.
  • K Yoshizawa
    ENTOMOLOGICA SCANDINAVICA 29 (2) 199 - 209 0013-8711 1998 [Not refereed][Not invited]
     
    A new genus, Atrichadenotecnum, is described in the tribe Psocini of the family Psocidae (Psocoptera). Atrichadenotecnum ryukyuense sp. n., A. tayal sp. n., and A. quadripunctatum sp. n. are described and Atrichadenotecnum nudum (Thornton: 1961) comb, n. and Atrichadenotecnum quinquepunctatum (McLachlan, 1872) comb. n. are transferred from the genus Trichadenotecnum Enderlein. The new genus is regarded as the sister-group of the genus Psocus Latreille.
  • Taxonomic study of the genusMesopsocus (Psocoptera: Mesopsocidae) of Japan
    Entomological Science 1 265 - 269 1998 [Not refereed][Not invited]
  • YOSHIZAWA Kazunori, SAIGUSA Toyohei
    Japanese journal of entomology 東京昆蟲學會 65 (1) 168 - 170 0915-5805 1997 [Not refereed][Not invited]
  • Species Diversity 2 51 - 58 1997 [Not refereed][Not invited]
  • Psocoptera of Bangladesh collected by Prof. Emer. Katsura Morimoto, with descriptions of four new species
    Esakia 37 15 - 24 1997 [Not refereed][Not invited]
  • YOSHIZAWA Kazunori
    Japanese Journal of Entomology 東京昆蟲學會 64 (1) 3 - 16 0915-5805 1996 [Not refereed][Not invited]
     
    The genera Heterocaecilius and Phallocaecilius of Japan are dealt with. H. solocipennis is transferred from the genus Pseudocaecilius and redescribed. A new species, Heterocaecilius fuscus, is described. Phallocaecilius hirsutus is recorded from Japan for the first time and redescribed. This is also the first record of the genus Phallocaecilius from Japan. Male and female genitalia are illustrated for the three species. Diagnostic characters of the genera and a key to the Japanese species of the genus Heterocaecilius are provided. This paper is the second part of the study of the Japanese Pseudocaeciliidae.
  • Taxonomic study on the family Pseudocaeciliidae (Psocoptera: Psocomorpha) from Japan. 3. GeneraAllocaecilius,Ophiodoperma andPseudoscottiella
    Japanese Journal of Entomology 64 300 - 312 1996 [Not refereed][Not invited]
  • YOSHIZAWA K.
    Jpn. J. Ent. 64 187 - 188 1996 [Not refereed][Not invited]
  • YOSHIZAWA Kazunori
    Jpn. J. Ent. 東京昆蟲學會 63 (4) 703 - 721 0915-5805 1995 [Not refereed][Not invited]
     
    The genus Pseudocaecilius of Japan is revised. Three species, P. citricola, P. kagoshimensis and P. maculosus are redescribed and a new subspecies, P. kagoshimensis yaeyamensis, is described. Male and female genitalia are illustrated for each taxon, as far as the materials are available. Generic diagnostic characters and a key to Japanese species and subspecies are provided. This paper is the first part of the study of the Japanese Pseudocaeciliidae.
  • New records of the Psocoptera from Yakutia, Russia
    Bulletin of the Biogeographical Society of Japan 50 19 - 22 1995 [Not refereed][Not invited]

MISC

Books etc

  • 節足動物の多様性と進化
    裳華房 2008

Association Memberships

  • Royal Entomological Society, London   Society of Systematic Biologists (US)   日本昆虫学会   日本進化学会   Willi Hennig Society (US)   

Research Projects

  • Japan Society for the Promotion of Science：Grants-in-Aid for Scientific Research
    Date (from‐to) : 2021/04 -2025/03 
    Author : 工藤 慎一, 吉澤 和徳, 大庭 伸也
     
    キンカメムシ科とツノカメムシ科を用いた予備的な比較分析の結果，保護の進化に伴って卵のアスペクト比が高くなる相関進化が検出された。すなわち，保護を行う種は，保護を行わない種に比べて細長い卵を生産する。次に楕円フーリエ記述子に対して主成分分析を行った結果, 第１主成分（以下PC1）と第２主成分（PC2）で卵形状における分散の大部分を説明できることが明らかとなった（累積寄与率＝98.6％）。PC1とPC2を軸とする形態空間を構築したところ, 保護を行う種の凸包は, 保護を行わない種の凸包とは重ならなかった。つまり，アスペクト比解析で明らかになった細長い形状に加えて，卵の上部が鈍端な形であると解釈される領域に位置した。 メス親が基質上に産み付けた卵塊を体で覆う姿勢で保護を行う種では，卵塊周辺部に対する捕食・寄生圧が高まることが明らかにされている（Eberhard 1975; Mappes & Kaitala 1994）。その結果，卵塊周辺部の卵に対する資源投資が減少し小型化する「卵サイズのクラッチ内変異」が生じていることが報告された（Mappes et al. 1997; Kudo 2001, 2006）。今年度得られた予備的な結果は，卵塊エリアを圧縮する方向に働く選択圧が，そのような卵サイズ変異だけでなく，卵形状の進化的変化にも関連していることを強く示唆している。すなわち，キンカメムシ科並びにツノカメムシ科において，メス親の防衛行動が細長い卵の進化を促したと考えられる。一方, 楕円フーリエ解析を用いた卵形状の定量化の結果, 保護を行う種の卵は上部がより鈍端な形であることが明らかとなったが，保護を行う種において, この卵の形状がどのような適応的意義をもつのか，現状では不明である。
  • 半翅系昆虫の全形態学：ゲノム系統の検証と新奇形質の進化プロセス解明
    Japan Society for the Promotion of Science：Grants-in-Aid for Scientific Research Grant-in-Aid for Scientific Research (B)
    Date (from‐to) : 2019/04 -2024/03 
    Author : 吉澤 和徳, 山田 量崇, 塘 忠顕
  • 文部科学省：基盤研究(C)
    Date (from‐to) : 2016/04 -2019/03 
    Author : 工藤慎一
  • 文部科学省：科学研究費補助金（基盤B)
    Date (from‐to) : 2015/04 -2019/03 
    Author : 吉澤 和徳
  • Japan Society for the Promotion of Science：Grants-in-Aid for Scientific Research
    Date (from‐to) : 2012/04 -2016/03 
    Author : YOSHIZAWA Kazunori
     
    Trichadenotecnum species of Southeast Asia were revised taxonomically. During the project period, a total of nine new species were described and a genus, Cryptopsocus from China, was synonymized with Trichadenotecnum. World-based phylogeny of Trichadenotecnum was estimated. Most species groups recognized morphologically were identified as monophyletic group. In contrast, extensive morphological convergences/reversals were detected in a few species group, which caused inappropriate species-group classification. Dating and biogeographical analyses suggested that the tans-continental dispersal via Bering Strait during the Tertiary period and reduction of distributional area during the last gracious period affected strongly in forming the present distributional pattern of Trichadenotecnum.
  • 日本学術振興会：科学研究費助成事業
    Date (from‐to) : 2013/04 -2015/03 
    Author : 吉澤 和徳, TSAI Jing-Fu
     
    親による子の保護行動の進化の要因の解明は，進化生物学で最も興味深いテーマの一つとなっている．本研究では母親による卵や子の保護行動の研究の進んでいる半翅目ツノカメムシ科ツノカメムシ亜科昆虫を対象に以下の研究を行った．１）ツノカメムシ亜科の分子系統関係の解明，２）卵保護行動の進化パターンの解明，３）卵保護行動の進化要因． １）核 18S rDNA, Histone 3，ミトコンドリア 12S rDNA, 16S rDNA, COI 遺伝子を用い，ツノカメムシ亜科44種の系統関係を推定した．外群にはツノカメムシ科の他の亜科の８種を用いた．解像度の高い系統仮説が得られ，Acanthosoma 属が非単系統群であることなど新たな知見も得られた． ２）１で得られた系統樹上に卵保護行動の有無を再構成した結果，保護行動が独立に少なくとも４回（亜科内３回，外群１回）進化していることが明らかとなった．亜科内での保護行動の単一起源の可能性は，尤度比検定により有意に却下された． ３）卵保護行動を行わない種で，腹部にある特殊な器官からの分泌物を卵に塗り付ける行動が確認された．また，この分泌器官は，卵保護行動を行う種では，消失または退化していることも明らかになった．分泌器官の有無を系統樹上に再構成し，卵保護行動の有無の再構成結果と比較したところ，卵保護の進化と分泌器官の退化／消失は常に同じ系統枝で起こっており，両形質の進化は系統的に有意に相関していることが明らかとなった．分泌器官が消失した種で，行動情報が得られていない種が多数存在するが，それらの種においてもらん保護行動が生じている可能性は高い．
  • 文部科学省：基盤研究 (B)
    Date (from‐to) : 2011/04 -2015/03 
    Author : 秋元信一
  • Japan Society for the Promotion of Science：Grants-in-Aid for Scientific Research
    Date (from‐to) : 2009/04 -2012/03 
    Author : YOSHIZAWA Kazunori
     
    (1) Genealogies of sika deer and sika deer lice were estimated using partial sequence of mitochondrial COI gene. Three lineages were identified for sika deer(Continental, Northern and Southern clades) whereas two lineages were identified for deer lice(Continent+southern Kyushu and others);(2) analyses of topological congruence revealed that there is no significant pattern of codivergence between deer and lice. The present analyses only used the maternally inherited gene marker so that male-deer mediated lice dispersal is considered to be the main source of the incongruence ;(3) long distance dispersal of lice apart from their host is quite unlikely. In such case, louse population genetic tree can be good indicators of hidden population structure in their host ;(4) using morphometric analyses, microevolutionary changes in deer lice morphology is shown to accumulate along phylogenetic lines. The neutral characters appeared to comprise a larger fraction of the variation compared to those expected to be more functionally significant. This suggests that random drift of more neutral characters plays a major role in the microevolution of morphology of deer lice
  • Ministry of Education, Culture, Sports, Science and Technology：Grants-in-Aid for Scientific Research(若手研究(B))
    Date (from‐to) : 2006/04 -2009/03 
    Author : Kazunori YOSHIZAWA
     
    (1)シラミ,チャタテムシの詳細な高次系統関係を推定した(2)シラミとコナチャタテにおいて,塩基置換速度の加速,塩基含有量の変化,リボソームRNA二次構造の変化と言った特異的な進化現象が,互いに相関して起こっていることを明らかにした(3)シカハジラミのミトコンドリアゲノムが,複数の微小環から構成されていることを明らかにした(4)特異的な進化傾向によって,遺伝子情報のアライメントが困難となるが,このような領域の解析に広く使われている方法論である直接最適化法に重大な問題点が存在することを明らかにした(5)ジュズヒゲムシの特異的な分子進化傾向を明らかにするとともに,このことによって塩基配列データがこの昆虫の高次系統推定に当たって有用な情報をもたらさないことを明らかにした.
  • 文部科学省：科学研究費補助金(若手研究(B))
    Date (from‐to) : 2003/04 -2006/03 
    Author : 吉澤 和徳
     
    国内各地およびマレーシアにおいて,研究に用いる半翅系昆虫のサンプリングを行った.これらのフィールドワークにより,分子系統解析に必要なサンプルの収集はほぼ完了した.チャタテムシ目の頭部の形態学的研究を行い,頭盾および小顎において相同性の解釈の誤りを見いだした・これらの形質は,穿孔-吸汁式口器の進化のキーとなる形質であり,今後この研究結果に基づき,半翅目および総翅目の頭部の形態学的再検討を行う予定である.特に,小顎は半翅目+総翅目の単系統性を支持する形質でもあるため,発生過程も含めた綿密な調査を行う.分子系統解析に用いるDNAマーカーの探索を行った.ミトコンドリアの12S,16SrDNAと,核の18SrDNA, Histone3,Winglessが,進化速度が遅く,かつ幅広い分類群において比較的容易に増幅できる事から,高次系統関係の探索に有効なDNAマーカーの候補として挙げられた.これらの領域を増幅するためのPCRプライマーの設計は完了し,半翅系昆虫の幅広いサンプリングに基づく予備実験では,良好な結果が得られている.また,ミトコンドリア領域の塩基配列情報に基づき,チャタテムシ目及びシラミ目の分子系統解析を行った.この結果,チャタテムシ目が側系統群である事が明らかとなるとともに,シラミ目のミトコンドリアゲノムに,他の昆虫には見られない特異な進化傾向が見られる事も明らかとなった.現在...
  • 文部科学省：科学研究費補助金(奨励研究(A), 若手研究(B))
    Date (from‐to) : 2001/04 -2003/03 
    Author : 吉澤 和徳
     
    1.塩基配列情報に基づくチャタテムシおよびシラミの系統解析昨年度および本年度収集したDNA解析用の約50サンプルからDNAを抽出し,ミトコンドリアの12Sおよび16SrDNAをPCR法を用いて増幅し,これに基づき系統解析を行った.最節約法,近隣結合法,最尤法いずれの方法を用いても,シラミ目の姉妹群がチャタテムシ目コナチャタテ科である事を強く支持する結果が得られた.さらに,シラミとコナチャタテのミトコンドリアゲノムが,置換速度,二次構造,GC含有量において,他の昆虫のミトコンドリアには見られない特異な進化傾向を持つ事を明らかにした.これらの結果は,アメリカIllinois Natural History SurveyのKevin P. Johnson博士との共著論文としてMolecular Phylogenetics and Evolution誌に投稿中である.2.シラミの分類学的再検討日本産フクロウハジラミ属を,上記サンプルおよび九州大学から借用した内田コレクションに基づき分類学的に再検討した.その結果,多くの同定ミスと未記載種の存在を明らかにした.さらに,フクロウハジラミ属が,寄主-寄生者の関係において,多くのハジラミ類に見られるような一対一の関係が見られず,さらには亜科を越えるような非寄主特異性が存在する事を明らかにした.これらの結果は,宮城教育大学の溝田浩二氏と共同で,...
  • Japan Society for the Promotion of Science：Grants-in-Aid for Scientific Research Grant-in-Aid for JSPS Fellows
    Date (from‐to) : 1999 -2000 
    Author : 吉澤 和徳
  • Japan Society for the Promotion of Science：Grants-in-Aid for Scientific Research Grant-in-Aid for JSPS Fellows
    Date (from‐to) : 1998 -1998 
    Author : 吉澤 和徳

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