Kazunori Yoshizawa

Hokkaido University | Hokudai · Graduate School of Agriculture

Genitalia are known to evolve rapidly and are among the most variable structures in insect morphology, making them a target of active research. However, function and evolutionary significance of internal genital structures remain less well understood. Here, we report the morphology and mechanism of a novel ejaculatory system that has evolved in the...

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...

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 fi...

Females of the silver-washed fritillary butterfly, Argynnis paphia (Linnaeus), and its relatives are known to have a unique genital projection called the cornucopia. Previous observations showed that the cornucopia partly penetrates the male genitalia during copulation, but its detailed structures and functions are unknown to date. Our observations...

Species of the Brazilian cave barklouse genus Neotrogla (Psocodea: “Psocoptera”: Trogiomorpha: Prionoglarididae: Sensitibillini) are known to have a “female penis (gynosome)” that functions as an intromittent organ inserted into the membranous pouches in the simple male genital chamber during copulation to receive semen. However, the functions of o...

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 understoo...

The widely distributed tropical species Psyllipsocus yucatan Gurney is recorded from several localities in Thailand. P. yongi New & Lee, previously only known from Peninsular Malaysia, is recorded from Thailand and its male is described for the first time. Four new species are described and illustrated: P. formosus Lienhard & Yoshizawa n. sp. (Peni...

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...

Brazil’s caves, home to diverse species and minerals, were stripped of protections by a recent presidential decree.

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...

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.

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...

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 wi...

Palaeosiamoglaris hkamtiensis sp. nov. of the family Prionoglarididae is described from Hkamti amber (ca. 110 Ma), together with Psyllipsocus myanmarensis sp. nov., the third fossil psyllipsocid species described from Noije Bum Hill (ca. 98 Ma). Based on the current observations of Palaeosiamoglaris hkamtiensis sp. nov., we discuss the synapomorphi...

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 Homil...

The insect order Psocodea is a diverse lineage comprising both parasitic (Phthiraptera) and non-parasitic 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 no...

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 a higher paternity Nuptial gifting sometimes causes severe female-female competition for obtain...

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...

Cormopsocidae n. fam. of the psocid suborder Trogiomorpha was proposed for a fossil species from mid‐Cretaceous Burmese amber, Cormopsocus groehni n. gen. & n. sp. This family was estimated to be the sister group of all other trogiomorphan taxa, but the possibility of much deeper divergence (i.e. placement as a sister group of all Psocodea) could n...

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 tha...

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) wa...

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...

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 t...

Although the great genital diversity of the barklouse genus Trichadenotecnum has been described in previous studies, the specific function of the genital structures during the copulation process has received less investigative attention. We reconstructed 3D‐models of each structure of the male and female genitalia of Trichadenotecnum incognitum in...

The Japanese white-toothed shrew (Crocidura dsinezumi) is a species endemic to Japan. For this species, only minimal phylogeographic investigations have been conducted. We obtained DNA sequences of mitochondrial cytochrome b and control region and nuclear ApoB genes for 191 individuals of C. dsinezumi from 107 locations collected throughout its kno...

Hemipteroid insects (Paraneoptera), with over 10% of all known insect diversity, are a major component of terrestrial and aquatic ecosystems. Previous phylogenetic analyses have not consistently resolved the relationships among major hemipteroid lineages. We provide maximum likelihood-based phylogenomic analyses of a taxonomically comprehensive dat...

The cave-dwelling psocid tribe Sensitibillini (Afrotrogla, Neotrogla and Sensitibilla) is of special morphological and evolutionary interest because of its possession of reversed copulatory organs: i.e. females of Afrotrogla and Neotrogla have a penis-like organ. The female penis structure is highly variable among taxa, as is the case of the male p...

The genus Neotrogla (Psocodea: Prinoglarididae) comprises four named species from Brazil. Females of this cave‐dwelling insect are characterized by a conspicuous penis‐like intromittent organ, termed a gynosome, which is inserted into the vagina‐like male genitalia during copulation. Another evolutionarily novel structure, the spermathecal plate, e...

Two new oversize troglobitic species of diplurans campodeids, Pacificampadaidarabotchi Sendra, sp. n. and Pacificampanipponica Sendra, sp. n. , found in three caves in two southern Japanese islands are described. It is the first record of cave-dwelling Diplurans from Japan and more specifically these are two Campodeinae of Pacificampa Chevrizov, 19...

We report a functional switching valve within the female genitalia of the Brazilian cave insect Neotrogla. The valve complex is composed of two plate-like sclerites, a closure element, and in-and-outflow canals. Females have a penis-like intromittent organ to coercively anchor males and obtain voluminous semen. The semen is packed in a capsule, who...

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 lic...

The elytral base sclerites (= sclerites located at the articular region between the forewing and thorax in Coleoptera) of selected taxa were examined and homologized. Although the elytral base sclerites are highly modified compared to the wing base sclerites of the other neopterans, they can be homologized by using the conservative wing flapping an...

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...

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 ex...

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 diff...

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...

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 Auchen...

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 Trichadenotecn...

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 sugges...

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...

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 c...

Yoshizawa, K. and Lienhard, C. 2015. The alinguum group, a new species group of the genus Trichadenotecnum, with descriptions of two new species from Thailand (Psocodea: 'Psocoptera': Psocidae). Ins. matsum. n. s. 71: 179-188, 5 figs. The alinguum species group is newly proposed in the genus Trichadenotecnum (Psocidae: Ptyctini), and diagnosis of t...

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 la...

The genus Cryptopsocus Li, 2002 is synonymized with Trichadenotecnum Enderlein, 1909. The type species of Crypto-psocus, 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 marginat...

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 stat...

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...

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...

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 st...

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....

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 im...

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 colle...

A very unusual genital apparatus of a species of the enigmatic insect order Zoraptera is described. It contains a unique configuration of two different intromittent organs, one of them coiled, very narrow, and strongly elongated. Hyper elongated genitalia are known in different groups of insects. Males have to accommodate these unwieldy structures...

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 t...

Sex-specific elaborations are common in animals and have attracted the attention of many biologists, including Darwin [1 • Darwin C.R. The Descent of Man and Selection in Relation to Sex. John Murray, London1871 • Crossref • Google Scholar ]. It is accepted that sexual selection promotes the evolution of sex-specific elaborations. Due to the fast...

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 ma...

Phylogenetic relationships among major groups of Criocerinae were reconstructed using molecular data (mitochondrial cytochrome oxi-dase 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 i...

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 u...

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 n...

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.

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 morpho...

All fossil psocid species ('Psocoptera', i. e. free living, mostly bark-dwelling members of the insect order Psocodea) known from Cretaceous amber are listed and their systematic placement is discussed. This critical evaluation of published data resulted in a list of 32 species assignable to 27 genera and 11 families. Each genus could be assigned t...

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...

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 molecular...