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Lachesilla anura Badonnel and L. grandis Badonnel are recorded from Yemen and both sexes are illustrated based on this newly-stu­died material. Males of these species were previously unknown. A male specimen described and tentatively assigned to L. anura by Badonnel is herein described as Lachesilla guineensis Lienhard, n. sp. The female genitalia of L. anura are unique in the genus Lache­silla Westwood, especially due to the complete absence of gonapophyses and the presence of particularly complex sclerotizations sur­rounding the spermapore.

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.

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 not be excluded. Cormopsocus groehni retains many plesiomorphic features, which will contribute greatly to elucidating the ancestral state of Psocodea.

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Lienhard, Charles (2021): A new species of Prionoglaris Enderlein (Psocodea: 'Psocoptera': Prionoglarididae) from an Armenian cave, with an account of the distribution of the genus. Revue suisse de Zoologie 128 (2): 227-235, DOI: 10.35929/RSZ.0048

Background Rickettsia felis is emergent in tropical areas. Despite its high morbidity, its natural history has not yet been fully determined. We investigated the role of the common household booklouse, Liposcelis bostrychophila, recently found to harbor R. felis. Methods Blood samples from 372 febrile patients from Senegalese villages, as well as nasal and skin samples from 264 asymptomatic individuals, were tested for cat flea–associated and booklice-associated strains of R. felis. Dust samples from beds were collected to isolate booklice and R. felis. Mice were infected with aerosol of R. felis strain from naturally infected booklice. Results Forty febrile patients (11%) were infected by R. felis, including 26 (7%) by the booklice-associated strain. Nine nasal samples (3.4%) and 28 skin samples (10.6%) contained R. felis, including 7 and 24, respectively, with the booklice-associated strain. The presence of live L. bostrychophila was observed in 32 dust samples (16.8%); R. felis was identified in 62 dust samples (32.5%). Several mice samples were positive for R. felis; interstitial lymphohistiocytic infiltrates were identified in lungs. Conclusions Liposcelis bostrychophila may be a reservoir of R. felis. The booklice-associated strain is pathogenic in mammals, causing pneumonia. Human infection may be acquired via inhalation of infected booklice particles.

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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.

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.

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 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.).

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.

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 penis in animals with normal copulatory organs. Here, we present the first molecular phylogeny of Sensitibillini and analyse the evolutionary pattern of their genitalia. Afrotrogla and Neotrogla did not form a monophyletic clade, and their female penis structures are significantly different, suggesting two independent origins of the female penis within Sensitibillini. In Neotrogla , the species that has a simple female penis is embedded among species that have an elaborate penis, and detailed structures of the female penis elaborations are in exact agreement among species, suggesting a secondary simplification of the female penis. A correlated evolutionary pattern between male and female genitalia was also detected. This coevolution of genitalia may suggest that sexual conflict or cryptic ‘male’ choice drove the diversity of the female penis, as is the case of male penile diversity in animals with conventional genitalia.

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, whose formation is initiated by seminal injection. It is not only used for fertilization but also consumed by the female as nutrition. The valve complex has two slots for insemination so that Neotrogla can continue mating while the first slot is occupied. In conjunction with the female penis, this switching valve is a morphological novelty enabling females to compete for seminal gifts in their nutrient-poor cave habitats through long copulation times and multiple seminal injections. The evolution of this switching valve may have been a prerequisite for the reversal of the intromittent organ in Neotrogla., eLife digest In dry caves of southeastern Brazil, live a group of insects named Neotrogla that are perhaps best known because the egg-producing females have penises while the sperm-producing males have vaginas. The sex roles of these Brazilian cave insects are also reversed: females compete over the males, who in turn are selective of their female partners. This sex role reversal likely evolved within Neotrogla because the males’ semen represents a rich and reliable source of energy within a nutrient-poor cave environment. When females are not using semen to fertilize their eggs, they consume it. Yet, while other animals show sex role reversal, Neotrogla species alone have reversed sexual organs. Neotrogla penises are spiky and may have evolved so that females can anchor themselves to males and then mate for prolonged periods. This would allow the females to stock up on the nutritious semen. Compared to their closest relatives, Neotrogla species can hold twice as much semen within their sperm storage organs. Scientists have speculated that a valve-like structure within this organ enables this extra storage by allowing the female to redirect semen into two separate chambers. But the organ’s small size has made it difficult to determine its inner workings, and scientists have yet to discover a switching valve that serves such a purpose within the animal kingdom. Yoshizawa et al. examined three Neotrogla species using advanced imaging technology and detected the first example of a biological switching valve. Neotrogla females can control this valve, switching the flow of semen between two slots. In this way, females can store two batches of semen at once. Seemingly exploiting this adaptation, the females’ spiky penises help them restrain males until they have received multiple semen injections. Yoshizawa et al. therefore suggest the emergence of this valve within the sperm storage organ may have promoted the evolution of the female penis. Along with giving insight into the lives of cave insects, these findings are of interest to engineers, who face challenges when constructing valves on a microscopic scale. The unique switching valve of female Neotrogla may one day inspire new man-made machinery that could advance a range of industries.

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.

SummarySex-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 (∼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.

The unusually developed head surface sculpture of a Pakistan strain of the stored-product psocid Liposcelis paeta was discovered using scanning electron microscopy (SEM). The morphologically unique serrate tubercle sculpture observed was previously unknown in the genus Liposcelis. The typical morphological female and male characteristics, which are important for species diagnosis, were described and visualised with SEM micrographs. The vertex sculptures of various geographic strains of L. paeta and of some species belonging to all known species-groups of the genus Liposcelis were compared to assess the extent of variability. Molecular data for L. paeta geographic strains are analysed and the taxonomic positions of these geographic strains are also discussed.

Psyllipsocus stupendus sp. nov. Figs 1-4 Holotype: MHNG; male (slide-mounted); Mexico, Morelos state, Biosphere Reserve Sierra de Huautla, 2.5 km N and 4 km W of the Centro de Educación Ambiental e Investigación Sierra de Huautla (CEAMISH), 940 m; 12.-17.iii.1996, Malaise trap 2; 18˚ 27’.671 N, 99˚ 02’.475 W [right wings and left maxillary palp lacking]. Paratypes: MHNG and CNIN (slide-mounted or in alcohol). – 6 males, 6 females (one of them allotype); same locality as for holotype; ii.-vi.1996, Malaise traps. – 1 female, 1 nymph; same locality, on rock outcrop; 9.vi.1996, leg. T. Martínez. – 1 female; Mexico, Morelos state, Biosphere Reserve Sierra de Huautla, Municipio de Tepalcingo, El Limón de Cuauchichinola, 1200 m, on rock outcrop; 11.v.1996, leg T. Martínez. Diagnosis: This new species can be distinguished from all known species of the genus Psyllipsocus by the very particular male and female genitalia, the general structure of which is already observable in nondissected specimens in alcohol. Etymology: The species epithet (Latin: stupendus, -a, -um) refers to the astonishing male and female genitalic structures. Description: General colouration yellowish to medium brown. Head without particular facial markings, compound eye brown, maxillary palp uniformly light to medium brown, antenna light brown. Legs: femur light brown, tibia medium brown with whitish sub-basal zone, tarsus medium brown. Forewing hyaline with more or less intense brown markings along several veins in apical half (Fig. 1 A), hindwing hyaline (Fig. 1 B). Abdomen whitish, basally and laterally with some brown hypodermal pigment, terminalia brown. Both sexes macropterous (Fig. 1 AB). Forewing: Rs and M fused for a length; distal closed cell slightly shorter than marginal length of pterostigma and much shorter than basal closed cell (bcc/dcc ≈ 3.6); first portion of pterostigmal R1 about equal in length to R1-Rs crossvein; CuA1 strongly curved, AP relativly short and high. Hindwing: Basal portion of Rs not differentiated and R1 originating from Rs-M fusion, thus closed cell triangular. Three ocelli present. Pilosity of frons and vertex almost uniform. Antenna with at least 23 segments (most antennae damaged); antennal flagellomeres with uneven surface (due to insertion points of long and relatively thick setae), in basal half of antenna maximal length of flagellar hairs about 5x greatest width of their flagellomeres; pedicellar microspades organ weakly developed (with only one unit). P2 with a weakly differentiated stout sensillum in basal half (somewhat thicker than other setae of similar length); P4 slender hatchet-shaped, with a prominent subapical sensillum (Fig. 1 F). Lacinial tip as in Fig. 1 E. Pretarsal claws simple, symmetrical, with a small preapical denticle; hind legs with well-developed coxal organ. Epiproct and paraproct simple in both sexes (Fig. 1 G), anal spine long, setal organ consisting of two fine setae of about equal length, but the ventral seta somewhat thicker than the dorsal one, paraproctal sensorium with 6-7 fine trichobothria on basal florets and one normal seta. Male genitalia: Hypandrium and phallosome forming a ventrally bulged structure of about half the length of the abdomen, laterally articulated to the sclerotized rod-like antero-lateral margin of the clunium (Fig. 1 C). Hypandrium with a distinct posterior lobe (Fig. 1 D); some long hairs on the lobe and near its base, remaining pilosity shorter (pilosity not shown in the figures); near base of the posterior lobe a pair of small dorsal hooked sclerites (Figs 1 D, 2B), probably linked by ligaments to the phallic cradle and/or to the endophallic tube. Phallic cradle differentiated as a well-sclerotized frame on inner wall of hypandrium (Fig. 2 A). Phallosome complex (Figs 1 D, 2C), with a pair of slender, movable, slightly curved and acutely pointed claspers; endophallic tube basally on each side with a half-moon shaped plate and distally tapering towards a pair of narrowly rounded and slightly curved tips; a pair of posteriorly directed basal struts inserted medially near anterior margin of hypandrium (forming a posteriorly opened V in ventral view; see Fig. 1 CD, interrupted lines), each strut linked to phallic cradle by a less sclerotized foliaceous structure (Fig. 2 C, dotted lines) attached on a small lateral denticle situated in about one third of its length. Female genitalia: Complex sclerotized structures of v1, v2, vagina and spermapore plate readily visible through the largely membranous subgenital plate in ventral view (Fig. 1 G). Subgenital plate membranous dorsally, at most weakly sclerotized ventrally, roughly triangular, bearing hairs of medium length on ventral surface but no particularly long setae on hind margin. Gonapophysis v3 typical for the genus (Fig. 3 A), almost rectangular, weakly sclerotized, bearing some stouter laterodistal setae (somewhat thicker than other setae of same length); v1 and v2 well-developed, elongated and strongly sclerotized (Fig. 3 AB), basally widened and connected to the sclerotized and longitudinally folded dorsal wall of the vagina (Fig. 3 B). Posterior part of this sclerotized vaginal tube laterally articulated to its anterior part by a joint-like connection (x in Fig. 3 B). Dorsal wall of the genital chamber transversally double-folded, resulting in a ventral main compartment, dorsally delimited by the sclerotized vaginal wall, and a dorsal secondary compartment, dorsally delimited by the spermapore plate (Fig. 4 B). Spermapore plate elongated and wellsclerotized, with a longitudinal ventral bulge on each side covering a region of glandular tissue (Fig. 3 B, interrupted lines); spermapore opened into an ovally semicircular sclerite at the anterior end of the spermapore plate (Figs 1 H, 3B), situated proximally in the dorsal compartment of the genital chamber (Fig. 4 B). Spermatheca thinwalled (Fig. 1 G), very fragile, its duct greatly coiled; spermatophore only observed in one female (Fig. 1 G: the structure figured may correspond to two simple spermatophores or to one more complex spermatophore). Dimensions: Male holotype: BL = 1.7 mm; FW = 1820 μm; FWw = 650 μm; FW/FWw = 2.8; HW = 1510 μm; F = 342 μm; T = 720 μm; t1 = 270 μm; t2 = 45 μm; t3 = 54 μm; IO/D = 1.41. – Female allotype: BL = 1.8 mm; FW = 1750 μm; FWw = 650 μm; FW/FWw = 2.7; HW = 1440 μm; F = 348 μm; T = 715 μm; t1 = 260 μm; t2 = 47 μm; t3 = 51 μm; IO/D = 1.44. Distribution: Mexico, Morelos state, mountains of the Sierra de Huautla Biosphere Reserve, on rock outcrops (but most known specimens collected in Malaise traps without indication of biotope or microhabitat). Type locality: Mexico, Morelos state, Sierra de Huautla Biosphere Reserve, 2.5 km N and 4 km W of the Centro de Educación Ambiental e Investigación Sierra de Huautla (CEAMISH), 940 m, 18˚ 27’.671 N, 99˚ 02’.475 W. Remarks: The new species seems to be related to a group of North and Middle American species characterized by distally elongated phallic structures with proximal insertion of a V-shaped pair of basal struts. The following species listed by Mockford (2011) show this type of male genitalia: P. albipalpus Mockford, 2011 (Mexico), P. apache Mockford, 2011 (Southwestern USA), P. flexuosus Mockford, 2011 (Guatemala), P. maculatus Garcia Aldrete, 1993 (Mexico, USA), P. oculatus Gurney, 1943 (Mexico, USA), P. poblanus Mockford, 2011 (Mexico), P. regiomontanus Mockford, 2011 (Mexico), P. squamatus Mockford, 2011 (Mexico), P. subterraneus Mockford, 2011 (USA: Texas). In these species the arms of the basal struts are anteriorly directed, as in all other species of the genus that have well-developed basal struts. In P. stupendus, the structures which are here considered as homologous to the basal struts (see description) are posteriorly directed (forming a posteriorly opened V), probably due to the presence of extremely elongated phallic sclerites. The brachypterous species P. kintpuashi Mockford, 2011, the male of which is not known, may also be related to P. stupendus. Its female has some stouter laterodistal setae on v3 and the spermathecal duct is greatly coiled, similar to P. stupendus. A tendency to sclerotization of the vaginal wall has never previously been observed in the genus Psyllipsocus, and in all known species v2 and v3 are relatively short and membranous or only weakly sclerotized. In spite of the striking autapomorphic genital characters of P. stupendus it is at present not justified to consider this species as a representative of a new genus. Concerning the non-genitalic structures, it fits well in the genus Psyllipsocus, and it is almost certain that this genus would become paraphyletic by a generic separation of P. stupendus. For further discussion see below. DISCUSSION In two of the three dissected females of P. stupendus, the genitalia were in normal resting position (see Figs 1 G, 4B), as in the other alcohol-preserved females of the present material. That of the female no. 8164 show an evaginated dorsal compartment of the genital chamber (Fig. 4 D), although this may be an artifact, possibly due to stress during fixation in alcohol. Without detailed observations of copulation and oviposition it is impossible to make a sound functional interpretation of the unusual genital morphology in P. stupendus. It may nontheless be of interest to present some preliminary hypotheses about the functional morphology of the complex male and female genitalic structures in this species. Fig. 4A shows the standard structure of female terminalia in Psocoptera. In some psocids the vagina (genital chamber) is subdivided by a membranous transversal fold of its dorsal wall in a ventral main compartment and a smaller dorsal compartment (e. g. Stenopsocus, see Badonnel, 1934). The spermathecal duct opens directly into the main compartment (Weber, 1931) or into the dorsal compartment near the distal part of the ventral compartment (Badonnel, 1934, 1951). For fertilization the sperm is released from the spermatheca and reaches the egg when it moves past the spermapore (Fig. 4A). In the female of P. stupendus the dorsal wall of the genital chamber is transversally double-folded; the ventral compartment is dorsally delimited by the sclerotized part of the vagina; the dorsal compartment is ventrally delimited by a thin membrane and dorsally by the elongate spermapore plate which bears proximally the opening of the spermathecal duct (Fig. 4 B). In the resting position, the distal end of the spermapore plate slightly surpasses the posterior end of the sclerotized vaginal wall (Figs 1 G, 4B). The situation after dissection of the terminalia (Fig. 4 C) and the observation of the evaginated dorsal compartment in the female no. 8164 (Fig. 4 DE) show that the membranous ventral wall of the dorsal compartment is attached to the distal end of the sclerotized part of the vaginal wall and entirely separated from the sclerotized dorsal wall of the ventral compartment. Thus, the position of the spermapore plate may vary during copulation and oviposition depending on the extent of evagination of the eversible dorsal compartment. Assuming that the egg passes through the ventral compartment while female genitalia are in resting position (Fig. 4 B), it cannot enter in direct contact with the spermapore for fertilization. In this case, the sperm released from the spermatheca would have to swim toward the distal end of the spermapore plate before reaching the egg, probably passing between the lateral bulges of the spermapore plate which contain glandular tissue (Fig. 3 B). Fertilization of the eggs seems therefore more complicated than in the standard situation, due to the presence of the sclerotized proximal part of the vaginal wall. The presence of a pair of strongly sclerotized, apically curved and pointed phallic claspers (Fig. 2 C), previously unknown in the genus Psyllipsocus, suggests that the sclerotized part of the vagina may have a protective function against traumatic effects of copulation, reminiscent of particular sclerotized structures associated with the vulvar area in Miridae (Heteroptera) which are interpreted as a defence system for the female against potential wounding by the male organ (Pluot-Sigwalt & Matocq, 2006). But the sclerotized part of the vagina may also provide new anchoring sites on which males can hold the mate by using their unique phallic claspers. Alternatively, it is also possible that the sclerotized vagina forms a tube-like functional unit together with the sclerotized first and second ovipositor valvulae. The presence of much reduced and weakly sclerotized v1 and v2 in all other species of Psyllipsocus suggests that these ovipositor valvulae may have a particular function in P. stupendus, possibly in combination with the adjacent sclerotized vaginal wall. In the genus Neotrogla Lienhard, 2010, belonging to the related trogiomorphan family Prionoglarididae, structures associated with the spermapore are inserted deep into the male genital chamber during copulation (Yoshizawa et al., 2014), which suggests the possibility of a somewhat similar mechanism in P. stupendus. The close contact between the female spermapore and the opening of the male seminal duct, indispensable for the transmission of the spermatophore in the suborder Trogiomorpha (Klier, 1956; Yoshizawa et al., 2014) may be obtained in P. stupendus by evagination of the dorsal compartment of the female genital chamber and the shallow insertion of the spermapore plate into the male genitalia. During copulation the phallic claspers may drag the extendable part of the female genitalia into the male body. An example of female genitalia shallowly inserted into the male abdomen in this way is known in the cricket Gryllus bimaculatus De Geer, 1773; in this species the female copulatory papilla, bearing the opening of the spermathecal duct, is protruded and enters the male genital cavity where genital coupling is achieved (Sakai & Kumashiro, 2004). Thus, the unique structures of male and female genitalia in P. stupendus may result from a reproductive biology which differs considerably from that of the other species of the genus. A special form of selective pressure based on sexual selection may be at the origin of the very particular genital structures in this species, while the natural (ecological) selection remains similar to most other species of Psyllipsocus, favouring the uniform general morphology (see the similar hypotheses for Zoraptera in Mashimo et al., 2014).

Charles Lienhard, Alfonso N. García Aldrete (2016): An extraordinary new species of Psyllipsocus (Psocodea: 'Psocoptera': Psyllipsocidae) from the Biosphere Reserve Sierra de Huautla, Morelos, Mexico. Revue suisse de Zoologie 123 (1): 105-112, DOI: 10.5281/zenodo.46291

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.

For the sterile insect technique, and other related biological control methods where large numbers of the target mosquito are reared artificially, production efficiency is key for the economic viability of the technique. Rearing success begins with high quality eggs. Excess eggs are often stockpiled and stored for longer periods of time. Any pests that prey on these eggs are detrimental to stockpiles and need to be avoided. Psocids of the genus Liposcelis (Psocoptera, Liposcelididae) are common scavengers consuming various types of organic material that are distributed globally and thrive in warm damp environments, making insectaries ideal habitats. In this short report, we investigated the species that has been found scavenging stored mosquito eggs in our insectary and identified it to be Liposcelis bostrychophila Badonnel, 1931. Additional observations were made to determine whether these predators indeed feed on mosquito eggs, and to suggest simple, effective ways of avoiding infestation.Élevage de masse de moustiques : mais qui mange les œufs ?Pour la technique des insectes stériles et les autres méthodes de lutte biologique associées, dans lesquelles un grand nombre de moustiques cibles sont élevés artificiellement, l’efficacité de la production est essentielle pour la viabilité économique de la technique. Le succès de l’élevage commence par des œufs de bonne qualité. Les œufs excédentaires sont souvent stockés pendant de longues périodes. Tous les organismes nuisibles qui exploitent ces œufs nuisent à ces stocks et doivent être évités. Les psoques du genre Liposcelis (Psocoptera, Liposcelididae) sont des charognards répandus qui consomment diverses matières organiques, sont répartis dans le monde entier et prospèrent dans des environnements chauds et humides, ce qui rend les insectariums des habitats idéaux pour eux. Dans ce court rapport, nous avons étudié l’espèce qui mangeait des œufs de moustiques stockés dans notre insectarium et nous avons déterminé qu’il s’agissait de Liposcelis bostrychophila Badonnel, 1931. D’autres observations ont été faites pour déterminer si ces prédateurs se nourrissent effectivement des œufs de moustiques et suggérer des moyens simples et efficaces pour éviter l’infestation.

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.

The unusually developed eyes of a Liposcelis silvarum strain from Croatia were discovered using scanning electron microscopy (SEM). Such a morphologically unique type of eye was previously unknown in the genus Liposcelis. The typical morphological characteristics of females and males, important for species diagnosis, were also described and visualised using SEM micrographs. This paper includes details of male genitalia and egg morphology not previously published for L. silvarum. Species status of this geographic strain is discussed.

Lienhard, Charles (2012): A new species of Austropsocus (Psocodea: 'Psocoptera': Pseudocaeciliidae) with a peculiar forewing venation from New Caledonia. Revue suisse de Zoologie 119 (2): 161-167, DOI: 10.5962/bhl.part.150022, URL: http://www.biodiversitylibrary.org/part/150022

is described for three new cave-dwelling species from Brazil: Neotrogla brasiliensis Lienhard sp. n. (from Minas Gerais State), N. aurora Lienhard sp. n. (from Tocantins State) and N. truncata Lienhard sp. n. (from Bahia State). These species are the first Neotropical representatives of the sub family Speleketorinae and the first New World representatives of the tribe Sensitibillini, previously known only from southern Africa. This distri butional pattern of Sensitibillini is tentatively interpreted as due to Western Gondwanan vicariance. In the females of Neotrogla a complex of acces sory structures to the spermathecal duct is described and denoted by the new term "gynosome". A hypothesis of functional complementarity, during copulation, between the "penis-like" gynosome and the strongly reduced male phallosome of Neotrogla is presented.

New and interesting psocids from Greece, Spain, and Portugal. — Six new species are described and illustrated: Asiopsocus meridionalis n. sp. (Spain), Caecilius ambulans n. sp. (Portugal), Caecilius arundinis n. sp. (Greece, Portugal), Lachesilla dimorpha n. sp. (Greece), Mesopsocus graecus n. sp. (Greece), Mesopsocus lusitanus n. sp. (Portugal). Trichopsocus fastuosus (NAVAS, 1915) nov. comb. (= Caecilius fastuosus NAVAS, 1915) is redescribed from Portugal. New distribution records are given for the following recently described species: Brachypsocus badonneli LIENHARD, 1979, Trimerocaecilius becheti MEINANDER, 1978, Hemineura bigoti BADONNEL, 1970, and Gibbopsocus pustulatus BADONNEL, 1975.

New records and synonymies in western Palaearctic Psocoptera. – Four new synonymies are established: Caecilius dahli Badonnel, 1963 = C. ambulans Lienhard, 1981 n. syn. (Caeciliidae); Trichopsocus dalii (Mclachlan, 1867) = T. kolosvaryi Danks, 1950 n. syn. (Trichopsocidae); Aaroniella badonneli (Danks, 1950) = A. eertmoedi Mockford, 1979 n. syn. (Philotarsidae); Loensia variegata (Latreille, 1799) = Psocus flavonimbatus Rostock, 1879 n. syn. (Psocidae). Asiopsocus meridionalis Lienhard (Asiopsocidae) and Caecilius dahli are recorded for the first time from Oman and Italy respectively. The nearctic species Eliopsocus guentheri Mockford (Elipsocidae) is recorded for the first time from the Palaearctic Region (Algeria).

About 2,000 specimens of Psocoptera were collected in Malta recently. Examination of this material revealed 21 new records for the Maltese archipelago, augmenting the known psocid fauna of these islands from 6 to 27 species. One of the most abundant species is Peripsocus stagnivagus Chapman, 1930 (= P. bivari Baz, 1988 = P. leleupi Badonnel, 1976, new synonymies), formerly considered to be a predominantly Nearctic species. The discovery in Malta, of one male of this usually parthenogenetic species enabled comparison of this first Palaearctic male with the well-documented, rare Nearctic male. The lack of any morphological difference between these males, or between females from the Nearctic, the western Palaearctic and several Atlantic islands, supports the proposed synonymies.

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). © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 146, 287–299.

Two new apterous species of Psocoptera (Insecta) are described and illustrated, each belonging to a new genus: Helenatropos abrupta gen. n., sp. n. (female) (Trogiidae) from a semidesertic biotope on the mid-Atlantic island of St Helena, and Thoracotroctes spinifer gen. n., sp. n. (male) (Pachytroctidae) from the endemic Didiereaceae vegetation in southern Madagascar. The systematic positions of the new genera within their families are discussed and a new subfamily is defined for Thoracotroctes gen. n.: Thoracotroctinae subfam. n. A key to the subfamilies of Pachytroctidae is provided.

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 Sismatoneura.

Reticulopsocus besucheti gen. n., sp. n. is described and illustrated from a series of specimens of both sexes collected under stones in southern Turkey. The new genus is closely related to the Mediterranean genera Chelyopsocus Lienhard and Philedaphia Lienhard. Some illustrations of the type genus of the family, Protroctopsocus Mockford, are also provided. Both sexes of the new genus are elytropterous and beetle-like in habitus but the fore wing venation is strongly sexually dimorphic: males have a normal venation, though somewhat simplified, females however, have a highly apomorphic, reticulate venational pattern. A key to the genera of the family Protroctopsocidae is provided.

Siamo glaris zebrina gen. n., sp. n. is described and illustrated from a male specimen collected in Thailand. The new genus is closely related to the Palaearctic genus Prionoglaris Enderlein. Illustrations of the type species of that genus, P. stygia Enderlein, are also given. The systematic position of these two genera within the family Prionoglarididae is discussed and the family is subdivided into two subfamilies: Prionoglaridinae Badonnel and Speleketorinae Smithers. A key to the adults of the genera of the family Prionoglarididae is provided.

After having analysed the most recent literature on Chinese Psocoptera, in particular the important monograph published by Li Fasheng in 2002, the following 174 nomenclatural acts are proposed, as a contribution to stabilize the nomenclature of Chinese Psocoptera. New genus names (replacement names for junior homonyms): Lifashengia nom. nov. (Amphientomidae) for Obeliscus Li Fasheng, 2002 nec Beck, 1837, nec Popofsky, 1913, nec Graybill, 1923; Licaecilius nom. nov. (Caeciliusidae) for Neocaecilius Li Fasheng, 2002 nec Li Fasheng, 2000; Fashengocaecilius nom. nov. (Pseudocaeciliidae) for Calocaecilius Li Fasheng, 2002 nec Mockford, 1974; Fashenglianus nom. nov. (Psocidae) for Lativalva Li Fasheng, 2002 nec Amsel, 1956; Lipsocus (Psocidae) nom. nov. for Cyclotus Li Fasheng, 2002 nec Edwards, 1852. New species name (replacement name for junior homonym): Peripsocus lifashengi nom. nov. for P. orientalis Li Fasheng, 2002 nec P. orientalis Garcia Aldrete, 1999 (Peripsocidae). New synonymies: Lachesilla Westwood, 1840 (Lachesillidae) (=Dicrolachesillus Li Fasheng, 2002), Ectopsocus McLachlan, 1899 (Ectopsocidae) (=Ectotrichus Li Fasheng, 2002), Smithersia Thornton, 1992 (Myopsocidae) (=Polygonomyus Li Fasheng, 2002). Re-established synonymies: Heterocaecilius Lee & Thornton, 1967 (Pseudocaeciliidae) (=Orbocaecilius Li Fasheng, 1999), Trichadenotecnum Enderlein, 1909 (Psocidae) (=Trichadenopsocus Roesler, 1943). 43 new combinations are proposed and 30 combinations are re-established. For 19 genus names and 71 species names the correct original spelling is selected. Incorrect subsequent spellings are only mentioned in some particular cases, concerning 25 genus names and 46 species names. Up to the end of the year 2002 a total of about 5470 valid species and 440 valid genera of Psocoptera have been described worldwide. 2600 species (almost half of the known world fauna) have been recorded from Asia, in particular due to the activity of the Chinese entomologist Li Fasheng, who described himself about 1500 of the approximately 1530 species known from China.

The following species are described and illustrated: Odontopsocus compactus sp. n. (Colombia), Lachesilla hirsuta sp. n. (Thailand), Ectopsocus venosus sp. n. (East Malaysia: Sabah). Each of them is characterized by a morphological structure which is unusual within the genus concerned and, in the case of Odontopsocus compactus, unique within the order Psocoptera. The presence of a cup-like, unsegmented and heavily sclerotized abdominal carapace in this species is discussed in the light of its mode of life in humid moss and litter and is compared with similar convergent structures in other psocids.

a checklist of the Western Palaearctic species of the family. - Valenzuela labrostylus sp. n. is described and illustrated (female only). The generic placement of this species remains doubtful because of the presence of labral stylets, which are absent in all other known Valenzuela species. The holotype of Caecilius corsicus Kolbe, 1882 is redescribed and the synonymy of this name with rhenanus Tetens, 1891 is proposed: Valenzuela corsicus (Kolbe) becomes the valid name of the most common Mediterranean species of Caeciliusidae. The 15 presently recognized Western Palaearctic species of this family are listed.

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 NewThornton is not included because its generic assignment is questionable. Females of T. cinnamonum EndangNew, T. imrum NewThornton and T. sibolangitense Endang, ThorntonNew, and the male of T. kerinciense EndangNew are described for the first time. A new species group is defined for T. krucilense Endang, ThorntonNew.

A new genus, Idatenopsocus, represented by Idatenopsocus orientalis (Vishnyakova, 1986), comb, nov., is described and the species is redescribed. Within the family Mesopsocidae, this new genus is considered as the sister group of a monophyletic group composed of the genera Mesopsocus, Microtrichipsocus, Metapsocus, Cyrtopsochus, Hexacyrtoma, and Rhinopsocus.

Two specimens of Psyllipsocus yucatan with black wings were found with normal individuals of this species on guano piles produced by the common vampire bat Desmodus rotundus. These specimens have both pairs of wings dorsally and ventrally covered by a black crystalline layer. They did not exhibit any signs of reduced vitality in the field and their morphology is completely normal. This ultrathin (1.5 µm) crystalline layer, naturally deposited on a biological membrane, is documented by photographs, SEM micrographs, energy dispersive spectroscopy (EDS) and X-ray diffractometry (XRD). The crystalline deposit contains iron, carbon and oxygen, but the mineral species could not be identified. Guano probably played a role in its formation; the presence of iron may be a consequence of the excretion of iron by the common vampire bat. This enigmatic phenomenon lacks obvious biological significance but may inspire bionic applications. Nothing similar has ever been observed in terrestrial arthropods.

The bark lice Psocus flavonimbatus Rostock, 1879 and Ptycta chubsugulensis Gunther, 1982 are assigned to the genus Kimunpsocus Yoshizawa, 2009. The male terminalia of K. flavonimbatus are described for the first time and compared to the corresponding structures of the other known species of the genus. Previously only known from the type locality in Estonia and the municipality of Kuhmo in Finland, K. flavonimbatus is now reported from 13 new sites and as a new species for four municipalities in eastern Central Finland. Analyses of an extensive material strongly suggest that K. flavonimbatus is a habitat-specialist species, confining its occurrence to pristine and semi-natural spruce-dominated forests. Nonetheless, its occurrence seems to be sporadic even in the old-growth forests. Forest structure of a subset of occupied and unoccupied sites is described and the conservation biology of K. flavonimbatus discussed.

A new species of Siamoglaris from Thailand with complementary description of the type species (Psocodea: ‘Psocoptera’: Priono glari didae). A new species of the previously monotypic genus Siamoglaris Lienhard, S. theresiae sp. n., is described and illustrated, based on three males from Thailand (female unknown). The female of the type species, S. zebrina Lienhard, is described for the first time and complements to the description of the male of this species are given. The generic diagnosis is revised. For the first time in Psocoptera, numerous thin-walled papilliferous spatulate setae (scent setae?) were observed on female gonapophyses and paraprocts; they are illustrated by scanning electron micrographs. Figures of female terminalia of the closely related genus Prionoglaris Enderlein are also presented.