The evolution of morphological castes under decoupled control.

Eusociality, where units that previously reproduced independently function as one entity, is of major interest in evolutionary biology. Obligate eusociality is characterized by morphologically differentiated castes and reduced conflict. We explore conditions under which morphological castes may arise in the Hymenoptera and factors constraining their evolution. Control over offspring morphology and behaviour seems likely to be decoupled. Provisioners (queens and workers) can influence offspring morphology directly through the nutrition they provide, while adult offspring control their own behaviour. Provisioners may, however, influence worker behaviour indirectly if offspring modify their behaviour in response to their morphology. If manipulation underlies helping, we should not see helping evolve before specialized worker morphology, yet empirical observations suggest that behavioural castes precede morphological castes. We use evolutionary invasion analyses to show how the evolution of a morphologically differentiated worker caste depends on the prior presence of a behavioural caste: specialist worker morphology will be mismatched with behaviour unless some offspring already choose to work. A mother's certainty about her offspring's behaviour is also critical-less certainty results in greater mismatch. We show how baseline worker productivity can affect the likelihood of a morphological trait being favoured by natural selection. We then show how under a decoupled control scenario, morphologically differentiated castes should be less and less likely to be lost as they become more specialized. We also suggest that for eusociality to be evolutionarily irreversible, workers must be unable to functionally replace reproductives and reproductives must be unable to reproduce without help from workers.

Body Part Selection for Size Classification of Two Trichogramma Species.

Body size is an important morphological characteristic that covaries with the quality of parasitoids and predators. Data show that the larger the organism is, the better the biological parameters and the host location by natural enemies in the field. The standard way of evaluating the size of parasitoids of the genus Trichogramma (Hymenoptera: Trichogrammatidae) is by measuring the tibia, but using only one body part to estimate the size of organisms can lead to miscalculations. In this paper, commercial Trichogramma pretiosum Riley, 1879 (Hymenoptera: Trichogrammatidae) and Trichogramma galloi Zucchi, 1988 (Hymenoptera: Trichogrammatidae) were mounted on slides for microscopy and photographed, and the photographs were used to measure their antennae, scutellum, ovipositor, tibia, and wing. Principal component analysis (PCA) and linear discriminant analysis (LDA) were performed to select the body part that best represents their size. PCA showed that all body parts represented size in a similar way, and LDA showed that the ovipositor was the most representative. We conclude that the best body parts for representing the size of the Trichogramma species studied are the wing and ovipositor, and at least two body parts are needed to detect two size groups.

Using DNA barcoding to identify high-priority taxa (Hymenoptera: Ichneumonidae) from Great Smoky Mountains National Park.

The All Taxa Biodiversity Inventory (ATBI) in Great Smoky Mountains National Park (GSMNP) seeks to document every species of living thing in the park. The ATBI is decades in progress, yet some taxa remain virtually untouched by taxonomists. Such "high priority" taxa include the hyper-diverse parasitoid wasp family Ichneumonidae. Despite the positive and multifaceted effects ichneumonids have on their environment, only a small percentage of those collected in the park have been identified as species, mostly to their complex morphology and overwhelming diversity. Recently, DNA barcoding has transformed biodiversity inventories, streamlining the process to be more rapid and efficient. To test the effectiveness of barcoding 20 + year-old specimens of Ichneumonidae and catalog new records for GSMNP, COI was amplified from 95 ichneumonid morphospecies collected from Andrew's Bald, NC. Species identifications were confirmed morphologically. Eighty-one ichneumonids generated sequence data, representing 16 subfamilies and 44 genera. The subfamily Oxytorinae is newly recorded from GSMNP, along with 10 newly recorded genera and 23 newly recorded species across Ichneumonidae. These results contribute significantly to the ATBI by adding new park records for a high-priority taxon and demonstrate the effectiveness of applying DNA barcoding to samples in long-term storage or those lacking immediate taxonomic expertise.

Insights from the past: the work of Hans von Alten on the evolution of brain structure, ecological adaptation, and cognition in hymenopteran species.

In his treatise on arthropod brains, Hans von Alten (1910) focuses on a specific functional group of insects-the flying Hymenoptera-which exhibit a spectrum of lifestyles ranging from solitary to social. His work presents a distinctive comparative neuro-anatomical approach rooted in an eco-evolutionary and eco-behavioral background. We regard his publication as an exceptionally valuable source of information and seek to inspire the research community dedicated to the study of the insect brain to explore its insights further, even after more than 110 years. We have translated and annotated his work, expecting it to engage researchers not just with its remarkable drawings but also with its substantive content and exemplary research strategy. The present text is designed to complement von Alten's publication, situating it within the temporal context of nineteenth-century and early twentieth-century studies, and to draw connections to contemporary perspectives, especially concerning a central brain structure: the mushroom body.

Lestiphorus bilunulatus yamatonis Tsuneki, 1963 (Hymenoptera, Crabronidae) raised to full species status, with description of the hitherto unknown male.

Lestiphorus yamatonis Tsuneki, 1963, stat. nov. is resurrected from synonymy and raised to full species level. The species diagnosis is updated and the hitherto unknown male is described. Illustrations of morphological details of L. yamatonis and related species are given. An updated key of the all twelve Palaearctic Lestiphorus is provided.

DNA barcoding of Sclerogibbidae and confirmation of the opposite sexes of Sclerogibba berlandi Benoit (Hymenoptera: Sclerogibbidae).

The female and male of Sclerogibba berlandi Benoit, 1963 (Hymenoptera: Sclerogibbidae), collected in Abu Dhabi (United Arab Emirates) and characterized by a pronounced sexual dimorphism, were associated for the first time by mitochondrial COI sequences. A comparison with COI sequences of S. crassifemorata Riggio De Stefani-Perez, 1888 and S. talpiformis Benoit, 1950 from Abu Dhabi proved the diversity of S. berlandi. The COI sequence of a female of S. rossi Olmi, 2005 from Japan, Okinawa-jima, was analyzed for the first time and compared with the other sequences.

Ultra-Conserved Elements and morphology reciprocally illuminate conflicting phylogenetic hypotheses in Chalcididae (Hymenoptera, Chalcidoidea).

Recent technical advances combined with novel computational approaches have promised the acceleration of our understanding of the tree of life. However, when it comes to hyperdiverse and poorly known groups of invertebrates, studies are still scarce. As published phylogenies will be rarely challenged by future taxonomists, careful attention must be paid to potential analytical bias. We present the first molecular phylogenetic hypothesis for the family Chalcididae, a group of parasitoid wasps, with a representative sampling (144 ingroups and seven outgroups) that covers all described subfamilies and tribes, and 82% of the known genera. Analyses of 538 Ultra-Conserved Elements (UCEs) with supermatrix (RAxML and IQTREE) and gene tree reconciliation approaches (ASTRAL, ASTRID) resulted in highly supported topologies in overall agreement with morphology but reveal conflicting topologies for some of the deepest nodes. To resolve these conflicts, we explored the phylogenetic tree space with clustering and gene genealogy interrogation methods, analyzed marker and taxon properties that could bias inferences and performed a thorough morphological analysis (130 characters encoded for 40 taxa representative of the diversity). This joint analysis reveals that UCEs enable attainment of resolution between ancestry and convergent/divergent evolution when morphology is not informative enough, but also shows that a systematic exploration of bias with different analytical methods and a careful analysis of morphological features is required to prevent publication of artifactual results. We highlight a GC content bias for maximum-likelihood approaches, an artifactual mid-point rooting of the ASTRAL tree and a deleterious effect of high percentage of missing data (>85% missing UCEs) on gene tree reconciliation methods. Based on the results we propose a new classification of the family into eight subfamilies and ten tribes that lay the foundation for future studies on the evolutionary history of Chalcididae.

Integrative taxonomy and phylogeography of Telenomus remus (Scelionidae), with the first record of natural parasitism of Spodoptera spp. in Brazil.

The egg parasitoid Telenomus remus (Hymenoptera: Scelionidae) has been investigated for classical and applied biological control of noctuid pests, especially Spodoptera (Lepidoptera: Noctuidae) species. Although T. remus was introduced into Brazil over three decades ago for classical biological control of S. frugiperda, this wasp has not been recorded as established in corn or soybean crops. We used an integrative approach to identify T. remus, combining a taxonomic key based on the male genitalia with DNA barcoding, using a cytochrome c oxidase subunit I mitochondrial gene fragment. This is the first report of natural parasitism of T. remus on S. frugiperda and S. cosmioides eggs at two locations in Brazil. We also confirmed that the T. remus lineage in Brazil derives from a strain in Venezuela (originally from Papua New Guinea and introduced into the Americas, Africa, and Asia). The occurrence of T. remus parasitizing S. frugiperda and S. cosmioides eggs in field conditions, not associated with inundative releases, suggests that the species has managed to establish itself in the field in Brazil. This opens possibilities for future biological control programs, since T. remus shows good potential for mass rearing and egg parasitism of important agricultural pests such as Spodoptera species.

Notes on the genus Netelia, subgenus Parabates (Ichneumonidae, Tryphoninae), with description of a new species from Oriental Region.

The characters that define the subgenus Parabates of the genus Netelia are investigated. The shape of the basal apodeme of the aedeagus is considered to be a synapomorphy of this subgenus, suggesting the monophyly of Parabates. A new species, Netelia (Parabates) fusciapicalis n. sp. belonging to the ishiharai group is described from Taiwan, Vietnam and Laos. This is the sixth extant species of the subgenus, the first record of the genus Netelia from Vietnam and Laos, and the first record of the subgenus Parabates from Taiwan.

Diaparsis phobos sp. nov. and D. deimos sp. nov., two amazing new species of Tersilochinae (Hymenoptera: Ichneumonidae) from Madagascar.

Two species of the genus Diaparsis Förster, D. phobos sp. nov. and D. deimos sp. nov., are described from Madagascar. The genus Diaparsis, as well as the subfamily Tersilochinae, is recorded from Madagascar for the first time. Both new species possess unuque morphological features previously unknown in the subfamily Tersilochinae: D. deimos sp. nov. has a remarkable slightly decurved ovipositor, and D. phobos sp. nov. possesses deep impressions on outer sides of ocelli.

Body and Wing Allometries Reveal Flight-Fecundity Tradeoff in Response to Larval Provisioning in Osmia lignaria (Hymenoptera: Megachilidae).

Variation in body size has important implications for physical performance and fitness. For insects, adult size and morphology are determined by larval growth and metamorphosis. Female blue orchard bees, Osmia lignaria, (Say) provision a finite quantity of food to their offspring. In this study, we asked how provision-dependent variation in size changes adult morphology. We performed a diet manipulation in which some larvae were starved in the final instar and some were given unlimited food. We examined the consequences on adult morphology in two ways. First, allometric relationships between major body regions (head, thorax, abdomen) and total body mass were measured to determine relative growth of these structures. Second, morphometrics that are critical for flight (wing area, wing loading, and extra flight power index) were quantified. Head and thorax mass had hyperallometric relationships with body size, indicating these parts become disproportionately large in adults when larvae are given copious provisions. However, abdominal mass and wing area increased hypoallometrically with body size. Thus, large adults had disproportionately lighter abdomens and smaller wing areas than smaller adults. Though both males and females followed these general patterns, allometric patterns were affected by sex. For flight metrics, small adults had reduced wing loading and an increased extra flight power index. These results suggest that diet quantity alters development in ways that affect the morphometric trait relationships in adult O. lignaria and may lead to functional differences in performance.

The genus Odontepyris/ Kieffer (Hymenoptera, Bethylidae) from China.

Ten new species of Odontepyris Kieffer from China are described and illustrated: O. angustus sp. nov., O. ecarinatus sp. nov., O. eminens sp. nov., O. flavipedis sp. nov., O. hirtus sp. nov., O. latifrontalis sp. nov., O. occiputalis sp. nov., O. xiaoi sp. nov., O. xui sp. nov., and O. yunnanensis sp. nov.. A key to the known Chinese species is provided.

A review of the subfamily Methochinae from Taiwan (Hymenoptera: Tiphiidae) with description of a new species and redescription of the known species.

Eleven species of Methocha Latreille from Taiwan are revised. Methocha cirrhocrus Narita Mita, sp. nov. is described and illustrated. The previously unknown male of M. maai Lin, 1966 is described. Methocha taoi Lin, 1966 is newly synonymized under Methocha areolata Lin, 1966. The genus Karlissa Krombein, 1979 is newly recorded from Taiwan, and a new combination is proposed for Methoca (sic!) tricha Strand, 1913, which is transferred to the genus Karlissa Krombein. A key to the species based on males and females is given.

Review of Holophris Mocsáry/ (Hymenoptera: Chrysididae) from Vietnam, with description of a new species.

The genus Holophris Mocsáry, 1890 of the tribe Elampini (Chrysidinae) is reviewed in Vietnam for the first time. A new species, H. melinh Nguyen Wisniowski, sp. nov. is described and illustrated, and two species, H. taiwana Tsuneki (1970) and H. thailandica Rosa, Wei, Notton Xu, 2016 are newly recorded. An updated key to all species of the genus known from Oriental region is presented.

Courtship, Mating Behavior, and Ovary Histology of the Nymph Parasitoid Psyllaephagus bliteus (Hymenoptera: Encyrtidae).

The potential of the parasitoid Psyllaephagus bliteus Riek for the biological control of the eucalyptus pest Glycaspis brimblecombei Moore (Hemiptera: Aphalaridae) nymphs is high. This research sought to analyze the courtship, male competition, and mating behavior of P. bliteus at the proportions of 1:1 and 2:1 males to female in a Petri dish (5 cm diameter), and to describe the ovary histology of virgin and mated females of this parasitoid. At 1:1, males touch the antennae and thorax-abdomen of females during courtship, but females avoid mate attempts before they are 48 h old. At 2:1, the competition between male parasitoids inhibits mating. The histology of ovaries of virgin and mated P. bliteus females is similar, with two well-defined germarium and vitellarium regions, with oocytes at different developmental stages, including mature ones rich in yolk and with eggshell. A clearer understanding of the reproductive behavior and histology of P. bliteus aids in the use of this parasitoid for the biological control of G. brimblecombei.

Revision of the Old World genus Mesocomys Cameron (Hymenoptera: Eupelmidae).

The Old World genus Mesocomys Cameron (1905) (Hymenoptera: Eupelmidae: Eupelminae) is revised. Eleven species, including two newly described species, are recognized and keyed in two previously established species groups, the albitarsis and the pulchriceps species groups sensu Gibson (1995), but with additional features provided to distinguish members of the two groups. Five species are recognized in the pulchriceps group-Mesocomys anelliformis n. sp., M. longiscapus n. sp., M. orientalis Ferrière, 1935, M. pauliani Ferrière, 1951, and M. pulchriceps Cameron, 1905. Seven species are assigned to the albitarsis group, but one, M. aegeriae Sheng, 1996 is treated as a nomen dubium; the six recognized and keyed species in the albitarsis group are M. albitarsis (Ashmead, 1904), M. breviscapis Yao, Yang Zhao, 2009, M. menzeli (Ferrière, 1930b), M. obscurus (Ferrière, 1930b) revised stat., M. superansi Yao, Yang Zhao, 2009, and M. trabalae Yao, Yang Zhao, 2009. Within the albitarsis group, the species are further discussed relative to two newly established species subgroups, the albitarsis subgroup for M. albitarsis, M. menzeli and M. obscurus, and the aegeriae subgroup for M. aegeriae, M. breviscapis, M. superansi and M. trabalae. Females of the albitarsis subgroup possess a finely sculptured mesoscutal medial lobe in combination with partly infuscate fore wings and/or at least partly pale flagellum, whereas females of the aegeriae subgroup possess a much more coarsely sculptured mesoscutal medial lobe and hyaline fore wings in combination with a dark flagellum. Members of the albitarsis species group are restricted to the Oriental and eastern Palaearctic regions except for a single female of the aegeriae subgroup seen from Algeria that is provisionally identified as M. breviscapis; members of the pulchriceps group are restricted to the Afrotropical region except for M. orientalis from the Oriental region. Newly placed in synonymy are M. aegeriae Sheng, 1998 under M. aegeriae Sheng, 1996 n. syn., M. sinensis Yao, Yang Zhao, 2009 under M. breviscapis Yao, Yang Zhao, 2009 n. syn., M. atulyus Narendran, 1995 under M. orientalis Ferrière, 1935 n. syn., M. vuilleti (Crawford, 1912) under M. pulchriceps Cameron, 1905 n. syn., and Semianastatus orientalis Kalina, 1984 and Mesocomys kalinai Özdikmen, 2011 under M. albitarsis (Ashmead, 1904) n. syns. Lectotypes are newly designated for M. menzeli, M. obscurus, M. orientalis, M. pauliani, M. pulchriceps and M. vuilleti. Morphological features characteristic of the genus and of the highly dimorphic sexes are described and illustrated, and the species are keyed, described, and illustrated through macrophotography. Phylogenetics are discussed for the genus, the two species groups, and species within the pulchriceps group. Distribution and host records are also summarized for each species.

Parasitoids (Hymenoptera) of Mealybug Pests (Hemiptera: Pseudococcidae) from Southern Brazil: Molecular and Morphological Characterization.

Parasitoids of three mealybug pests (Hemiptera: Pseudococcidae), Planococcus ficus (Signoret), Pseudococcus sociabilis Hambleton, and Pseudococcus viburni (Signoret) have been identified for the first time in Brazil. Mealybugs were collected in fruit-growing areas along southern Brazil during 2013-2016. An integrative approach, combining morphological and molecular methods, was used to identify the Brazilian parasitoids to the species level. Fifteen species were recorded, including 14 primary parasitoids belonging to Encyrtidae and Platygastridae and a single secondary parasitoid species belonging to Signiphoridae. The encyrtid parasitoids Acerophagus flavidulus (Brèthes), Anagyrus calyxtoi Noyes and Zaplatycerus sp., and the signiphorid secondary parasitoid Chartocerus axillaris De Santis are reported for the first time in Brazil.

The origin of wing polyphenism in ants: An eco-evo-devo perspective.

The evolution of eusociality, where solitary individuals integrate into a single colony, is a major transition in individuality. In ants, the origin of eusociality coincided with the origin of a wing polyphenism approximately 160 million years ago, giving rise to colonies with winged queens and wingless workers. As a consequence, both eusociality and wing polyphenism are nearly universal features of all ants. Here, we synthesize fossil, ecological, developmental, and evolutionary data in an attempt to understand the factors that contributed to the origin of wing polyphenism in ants. We propose multiple models and hypotheses to explain how wing polyphenism is orchestrated at multiple levels, from environmental cues to gene networks. Furthermore, we argue that the origin of wing polyphenism enabled the subsequent evolution of morphological diversity across the ants. We finally conclude by outlining several outstanding questions for future work.

Morphometric based differentiation among Trichogramma spp.

Genus Trichogramma Westwood is being utilized in various biological control programme worldwide. Trichogramma spp. are egg parasitoids which lay their eggs inside the host eggs. More than 240 species are known, of which, 45 species are recorded from India. It is quite difficult to identify and differentiate among the species of Trichogramma due to smaller size. In present study, we hypothesized a methodology to identify the Trichogramma species based on potential morphometric characteristics. The males and females of five different species of Trichogramma namely, T. brassicae, T. chilotraeae, T. danaidiphaga, T. danausicida and T. dendrolimi were morphometrically analysed. A total of 33 characters in males and 27 in females were taken into consideration for morphometric identification and analysis using principal component method. It was observed that few characters of male like body length, length of flagellar hair, maximum marginal hair length of fore wing, genitalia characters including aedeagus length, distance between chelate structure and gonoforceps, and others were identified as important morphometric characters. Similarly, in females, ovipositor length, body and head length, eye width, antennal club width and number of setae in forewings were found important for taxonomic identification. Male and female of each species are clearly identified from high definition microscopic images.

Relationship Between Body Mass and Forewing Length in Neotropical Ichneumonidae (Insecta: Hymenoptera).

Biomass is an important indicator of various ecological factors in insect populations and communities. Although the utility of this variable is proven, it is usually dismissed in ecological studies, generally, because of the difficulty of measuring it. Considering that insect biomass is apparently declining worldwide, here, we aimed to test the accuracy of forewing length as an estimator of body mass in Neotropical Ichneumonidae to help in monitoring insect biomass. Forewing length and dry body mass were measured in 728 individuals from distinct localities in Venezuela and Ecuador. A clear log-linear relationship existed between the two variables, and a log-linear regression had significant and strong predictive power of mass based on wing length. To derive the final values of mass from predictions made using our equation, we used an exponential transformation. We present an R function for predicting mass from wing length. We consider our findings applicable to many Neotropical species of Ichneumonidae, and helpful especially when specimens are measured in dry collections.