Genital Capsule Research Articles

“Picking up signals” in male genital morphospace and integrating phylogenomics to delimit Neotropical Nylanderia Emery species (Hymenoptera: Formicidae)

Abstract Genital morphology, a cornerstone in taxonomy that predates Linnaeus’s Systema Naturae, is vital for species delimitation. However, the widely accepted paradigm that genitalia are taxonomically informative lacks robust testing between closely related species, and supporting evidence is often limited to taxonomic literature in which genitalia are assumed a priori to be species-specific. The cosmopolitan ant genus Nylanderia Emery includes 123 described species, with most in the Neotropics still undescribed. Workers are often morphologically cryptic, and males are rare in collections but required for morphological delimitation. Using Ultraconserved Elements (UCEs) from 236 samples, including 53 Neotropical Nylanderia species, we reconstructed a phylogenetic framework to compare the genitalia (gonopods) of males collected alongside workers. We used geometric morphometrics on images of slide-mounted genitalia from 16 species and nano-CT scans of Nylanderia fulva (Mayr) and Nylanderia pubens (Forel) genital capsules and interpreted results considering phylogeny under maximum likelihood and the multispecies coalescent. We found strong morphological and molecular support for 2 distantly related American clades, identifiable by gonopod shape, with significant differences observed among most species. Three previously reported COI clades of N. fulva were not supported as monophyletic, nor were their gonopods significantly different. However, N. pubens was supported as distinct by all phylogenetic and 3DGM results. Our findings emphasize the importance of male genitalia for delimiting species boundaries and revising Neotropical Nylanderia. Given their importance, particularly in morphologically cryptic taxa, we recommend a greater focus on linking male and worker phenotypes, which can be facilitated through comprehensive nest series collection.

Discovery of the male of Augochloropsis (Glyptochlora) Moure (Hymenoptera, Apidae, Halictinae), with a revised diagnosis for the subgenus

Augochloropsis (Glyptochlora) Moure is a morphologically homogeneous and rare subgenus of Augochlorini, endemic to the Amazon Basin. So far, the subgenus was known only from females. Herein, we identified and studied the male of Augochloropsis (Glyptochlora) peruviana Celis & Melo. This specimen represents the first opportunity to document the male morphology of Augochloropsis (Glyptochlora). In addition to the description of the male, including genital capsule and metasomal sterna, we provide an updated diagnosis for A. (Glyptochlora).

Two New Species of Melucha Amyot and Serville, 1843 from French Guiana (Hemiptera: Heteroptera: Coreidae: Nematopodini), with a Revised Key to the Species

Two new species of Melucha Amyot and Serville, 1843 collected in French Guiana are described: M. edictum new species and M. flatilis new species. Dorsal photographs are provided as well as photos of male genital capsules and parameres where available. A key to the 15 known species is included.

Historical Note on the Genus Trichogramma (Hymenoptera, Trichogrammatidae) in Brazil, Focusing on Taxonomy and Diversity.

The history of the taxonomy of Trichogramma parasitoid wasps can be divided into two phases. In the first phase, species identifications were based on external morphological characters; however, these identifications proved to be confusing and inaccurate. In the second phase, starting in the 1970s, taxa were identified based on charcteristics of the male genital capsule, leading to a major advance in Trichogramma taxonomy. The history of Trichogramma taxonomy in Brazil is recent and mainly related to species that parasitize agricultural pests. In Brazil, the first phase of Trichogramma taxonomy occurred in the 1960s, while the second phase occurred from the 1980s onward. In this second phase, Trichogramma taxonomy progressed significantly and knowledge of Trichogramma diversity as well as associations with lepidopteran pests improved markedly in Brazil as well as worldwide. The last five decades have seen significant progress in studies in Brazil, with taxonomy evolving from identifications based exclusively on morphological characters to integrative taxonomy, encompassing biology (crosses) and morphometry. This historical outline presents the phases of Trichogramma taxonomy in Brazil, addressing the hurdles encountered in the first descriptions, erroneous records of the species, and species descriptions since the 1980s. We highlight the importance of accurately identifying Trichogramma taxa for their use in biological control, as well as species diversity and associations with lepidopteran hosts.

Review of the Newly Established amictus-Subgroup of the Genus Leptocoris (Hemiptera: Heteroptera: Rhopalidae), with the Description of a New Species from the Arabian Peninsula

Leptocoris arabicus new species (Hemiptera: Heteroptera: Rhopalidae: Serinethinae) is described based on 55 specimens of both sexes collected in various localities of southern Oman and southeastern Yemen; a hypothesis is proposed as to its host plant. The new species is close to Leptocoris amictus Germar, 1838 and to Leptocoris ruber Göllner-Scheiding, 1980, new status, due to the presence of a produced horned process along the posterior ventral margin of the genital capsule. It is distinguished from both by differences in colour pattern and morphology. An informal L. amictus species-subgroup is proposed to accommodate these three species within the Leptocoris hexophthalmus species-group sensu Göllner-Scheiding (1980). The following new faunistic records are also provided: L. amictus from Ethiopia, Mozambique and Zambia; L. ruber from Central African Republic. The occurrence of L. chevreuxi (Noualhier, 1893) in Oman is confirmed. The list of all available species-group names assigned to Leptocoris Hahn, 1833 (gender masculine) and their synonyms is compiled and the gender agreement is corrected in the species names when appropriate. The nomenclatural problem concerning Leptocoris living on Marshall Islands discovered by Kerzhner (2003) is discussed and the following corrections are provided: Leptocoris rufus rufus Hahn, 1833, new status = Leptocoris lariversi Usinger, 1952, new synonymy, and Leptocoris rufus isolatus (Distant, 1914) new status.

Descriptions of Four New Western Hemisphere Palm Bugs (Heteroptera: Thaumastocoridae: Xylastodorinae): A New Species of Xylastodoris from Haiti and Two Remarkable New Genera, One with Two New Species from Ecuador, and One with A New Species from Peru

Four new Western Hemisphere xylastodorine palm bugs (Thaumastocoridae: Xylastodorinae) are recognized. Neolatebracoris new genus is described to accommodate Neolatebracoris erwini new species from Peru; Xylastodoris cornutus new species is described from specimens intercepted at Miami, Florida, from Haiti; and Xylastotylus new genus is described to accommodate Xylastotylus emarginatus new species (as the type species) and X. minutus new species from Ecuador. Dorsal and lateral habitus and available genital capsule images, diagnoses, and descriptions are provided for each new genus and species. Relationships and biogeography of the new taxa are discussed, and revised keys to the New World genera of the subfamily and the species of Xylastodoris and Xylastotylus are provided.

Phylogenetic Trends in the Dissymmetrisation of Genitalia in Hadenini (Lepidoptera, Noctuidae)

The external genital appendages of Noctuidae represent correlated–coevolved elements of a complex structure. The pathways of changes are delimited by some constraints, and they are parallelly evolved in different phyletic lines. Asymmetrical male external genitalia were found in Hadenini, in Polia and its closely related genera, but also in Anarta (s. l.). In Poliina, the asymmetry of the genital capsule is mostly expressed in the Holarctic Polia nebulosa species group, forming the basal split within Polia (s. str.). Higher species diversity, as a consequence of asymmetry connected with the split of functions between the right and left sides, was observed in Polia (s. str.) and in Anarta (s. l.). The highest level of expansivity was observed in the Holarctic Anarta, where some species are migrating and becoming invasive ones, while a bulk of species populates the steppic and mountainous habitats of North America and Central Asia. Parallelly, while some genera/subgenera of the subtribe Poliina are oligotypic and/or consist of strictly localised species, the sensu stricto Polia species with asymmetrical genital capsules are widely distributed. The diversification of the Hadulina phylogenetic line should have been initiated by the Messinian aridisation crisis. The core area of diversification of Poliina was connected with the Sino-Himalayan region, followed by expansion into the Holarctic boreo-nemoral zone.

Descriptions of a new species and of three unknown males of Xylocopa Latreille, 1802 from the Neotropical region (Hymenoptera: Apidae: Xylocopini).

A new species of the previously monotypic subgenus Xylocopa (Xylocopina), X. asedothae Zama & Silveira sp. nov., and males of three species previously known only from females, X. (Diaxylocopa) truxali Hurd & Moure, 1963, X. (Dasyxylocopa) fortuita Melo, 2017 and X. (Schonnherria) ornata Smith, 1874, are described and illustrated for the first time. Diagnoses, comments on morphological variation, photographs and updated geographical distributions for all them are also provided. An identification key for both sexes of the species of X. (Dasyxylocopa) is updated and photographs of the genital capsules for males specimens used here are also provided.

Description of three new species of Carvalhygia Brailovsky, 1995 from The Republic of the Philippines and a key to species (Heteroptera: Coreidae: Colpurini).

Three new species of Carvalhygia Brailovsky, 1995 collected in The Republic of the Philippines are described in the tribe Colpurini (Heteroptera: Coreidae)C. complanus sp. nov., C. herczekia sp. nov. and C. stellata sp. nov. A key to the seven described species is included, illustrated with photomicrographs of dorsal and lateral habitus, female genital plates, male genital capsule of C. camarinesa, C. carvalhoi and C. herczekia, and parameres of C. camarinesa and C. herczekia.

Illustrated key to the species in the genus Laminiceps Costa, 1863 (Hemiptera: Heteroptera: Coreidae: Acanthocephalini), with descriptions of twelve new species.

The genus Laminiceps Costa, 1863 (Coreidae: Acanthocephalini) is redescribed, and 12 new species are described: L. aburtonia (Brazil), L. curiosus (French Guiana), L. eustoliana (Paraguay), L. excelsus (Peru), L. factiosus (Brazil), L. fundatus (Trinidad), L. gelidus (Brazil), L. genuinus (Peru), L. halatus (Brazil), L. infrequens (Brazil), L. lentus (Venezuela), and L. navus (Bolivia). A revised key to all 30 species is included. Dorsal and lateral photos of all species are provided as well as photos of male genital capsules and parameres where available.

Reproductive Apparatus, Gonadic Maturation, and Allometry of Cyclocephala barrerai Martínez (Coleoptera: Melolonthidae: Dynastinae).

Simple SummaryCyclocephala barrerai larvae feed on the roots of ornamental and commercial grasses. Most of their biology is still unknown, and there are no tools to manage their populations. The objective of this work was to study the reproductive system, gonadic maturation, and allometry of C. barrerai. This is the first report into the Melolonthidae family that characterizes the reproductive apparatus, gonadic maturation of both sexes, and allometric relationships of virgin F1 specimens. Adults do not feed, so gonadic maturation depends on larvae reserves. Females present sclerotized accessory glands and a genital chamber with muscular fibers and bacteria. The morphology of the parameres and the antennae are under sexual selection. This is the first report of two main differences among sexes: females are heavier while males have longer antennae. Males and females exhibit allometric relationships that can be used to predict the bodyweight of field-collected specimens.The Order Coleoptera provides good examples of morphological specializations in the reproductive apparatus, gonadic maturation, and allometry differing between the sexes. The female and male reproductive apparatus has been modified to ensure reproduction between individuals of the same species. The genus Cyclocephala has more than 500 species distributed in America, and Cyclocephala barrerai Martínez is an economically important species in the central part of Mexico. The objective of this work was to study the reproductive system, gonadic maturation, and allometry of C. barrerai. We used light, scanning electron, and laser scanning confocal microscopy to describe the reproductive apparatus and gonadic maturation of females and males. The relationship between adult weight and different parts of the body was established by linear regression. Regardless, the reproductive apparatuses of C. barrerai are like those of other Melolonthidae: the genital chamber, the type II accessory glands, and the ventral plaques of the female and the ejaculator bulb and genital capsule of the males are specific to C. barrerai. The gonads are fully developed when 18 d old. The weight of adult C. barrerai has a positive linear relationship with distinct parts of its body, while the antennae of males are larger than those of the females.

A New Species of the Stalk-Eyed Bug Genus Epipolops from Colombia, with Clarification on the Identity and a Redescription of Epipolops quadrispinus Stål (Hemiptera: Heteroptera: Geocoridae)

The long-misidentified stalk-eyed bug Epipolops quadrispinus Stål is redescribed and its identity is clarified, and Epipolops tuberculatus new species is described from Colombia based on specimens previously misidentified as E. quadrispinus. Dorsal and lateral habitus images of the adults and male genital capsules are provided to facilitate identification.

Illustrated key to the species in the genus Sephina (Hemiptera: Heteroptera: Coreidae: Coreinae: Spartocerini), with descriptions of three new species and new distributional records.

Three new species of Sephina Amyot and Serville 1843, are described: S. crassa from Ecuador, S. effrena from Colombia and S. ornata from Panama. A key to the 27 known species and photographs of the dorsal habitus are included. Also included are photographs of the male genital capsule of S. vinula Stl 1862, and parameres of S. indierae Wolcott 1924 and S. vinula. New distributional records for S. dorsalis (White 1842), S. esquivelae Brailovsky Schez 1983, S. faceta Brailovsky 2001, S. formosa (Dallas 1852), S. geniculata Distant 1881, S. limbata Stl 1862, S. nigripes Schmidt 1907, S. pagella Brailovsky Schez 1983, S. pubera (Erichson 1848), S. pustulata (Fabricius 1803), S. rogersi Distant 1881, S. subulata Brailovsky Schez 1983, and S. vinula are included. Sephina excellens Schmidt, 1907 is synonymized under S. pubera (Erichson 1848). The synonymy of Sephina grayi Van Duzee, 1909 under S. gundlachi (Gurin-Meville 1857) is here corroborated.

EmUrolabida/em emgraziae/em, new urostylidid species from the Philippines with comments on the current taxonomy and systematics of the family (Hemiptera: Heteroptera: Urostylididae).

A new species of Urostylididae from the Philippine islands (Mindanao) is herein described. This new species, similar to Urolabida bipunctata Stål, 1871, differs from the latter in the shape of the external male genitalia. Urolabida bipunctata and the new species, Urolabida graziae Roca-Cusachs, sp. nov., are extremely similar in external appearance, and fit into the (incomplete) description of genus Urolabida Westwood, 1837. The examination and comparison with the type specimens of other Urostylididae species, especially the male genital capsule, clearly separates this two species from Urolabida tenera Westwood, 1837, the type species of the genus Urolabida, however the taxonomy of this group is not resolved and needs a deep revision, therefore we refrain from description of a new genus-group taxon here. Additionally, images of the type specimens of Urolabida tenera, Urostylis histrionica Westwood, 1837, and Urostylis punctigera Westwood, 1837, are provided and discussion on the current systematics and classification of the family, and particularly of genus Urolabida are also given.

Revision of the New World leafhopper tribe Faltalini (Hemiptera: Cicadellidae: Deltocephalinae) and the evolution of brachyptery.

The taxonomy of the deltocephaline leafhopper tribe Faltalini (13 genera, 63 species) is revised. A key to all genera and keys to species for each genus are provided. All genera are described and at least one species of each genus is illustrated including images of the habitus, male and female genitalia, and scanning electron microscope (SEM) images of the head, face, and other characters. The new genera Dietrichana n.gen. [type species D. pampas n. sp.] and Paraclorindaia n.gen. [type species P. pinguis n. sp.] are described and the following 36 new species are described: Ackbaria jujuyensis n. sp. [Argentina], A. ojosverdes n. sp. [Argentina], Aequcephalus laplata n. sp. [Chile], Bonamus jussaral n. sp. [Brazil], Clorindaia obrienorum n. sp. [Uruguay], C. otamendi n. sp. [Argentina], Dietrichana pampas n. sp. [Argentina], Faltala catalanoae n. sp. [Argentina, Uruguay], F. paradellae n. sp. [Argentina], F. viscacha n. sp. [Argentina, Uruguay], Hecullus mexicanus n. sp. [Mexico], Kramerana saltensis n. sp. [Argentina], Paraclorindaia pinguis n. sp. [Argentina, Uruguay], P. vesperis n. sp. [Argentina], Tenucephalus admirabilis n. sp. [Argentina], T. bartletti n. sp. [Peru], T. brevilamina n. sp. [Argentina], T. cerrado n. sp. [Brazil], T. ecuadorensis n. sp. [Ecuador], T. flagellifer n. sp. [Peru], T. heppneri n. sp. [Peru], T. iguazu n. sp. [Argentina], T. interstinctus n. sp. [Peru], T. johnsoni n. sp. [Brazil], T. lingulatus n. sp. [Bolivia, Peru], T. manuensis n. sp. [Peru], T. misiones n. sp. [Argentina], T. musettiae n. sp. [Brazil], T. nielsoni n. sp. [Brazil], T. novafriburgo n. sp. [Brazil], T. parana n. sp. [Argentina], T. spectabilis n. sp. [Peru], T. takiyae n. sp. [Brazil, Peru], T. umbrinus n. sp. [Brazil], T. viperinus n. sp. [Argentina], and Virganana danasensis n. sp. [Ecuador]. The new combinations Paraclorindaia brasileira (Zahniser) n.comb. [transferred from Clorindaia] and Tenucephalus amabilis (Linnavuori Heller) n.comb. [transferred from Bonamus] are recognized. A phylogenetic analysis based on 51 morphological characters scored for 42 taxa including two outgroup taxa was performed. The resulting phylogeny resolved Faltalini as monophyletic, but the tribe remains difficult to diagnose because it encompasses a large amount of morphological diversity in characters traditionally used to define tribes of Deltocephalinae including head morphology, wing length and venation, and male genitalia. Despite this diversity, members of the tribe share the following characters: 1) phragma of male genital capsule partly developed into sclerotized plates or setose bulbous lobes (exception: Acrolithus); 2) first valvula shape relatively straight; 3) first valvula dorsal sculpturing separated from dorsal margin by unsculptured band; 4) first valvula dorsal sculpturing pattern maculose, with maculae separate or only slightly overlapping, or granulose; 5) first valvula with distinctly delimited ventral sculptured area; and 6) second valvula without dorsal teeth. Two major clades within the tribe were resolved and strongly supported, consisting of (Bonamus, Tenucephalus) and (Hecalocorica, (Hecullus, (Acrolithus, (Ackbaria, (Aequcephalus, Dietrichana, Virganana, Clorindaia, Faltala (Kramerana, Paraclorindaia)))))). A stepwise evolution of brachyptery in the latter clade is suggested by the phylogeny, beginning with evolution of subbrachypterous females in Hecullus and Acrolithus and culminating in completely brachypterous males and females in the monophyletic "Faltala group" of eight genera. It is hypothesized that a transition from an ancestral forested habitat to an open or grassland habitat facilitated the evolution of brachyptery. The hindwings and associated thoracic structures of eight species of Faltalini and three other Deltocephalinae were imaged with SEM. Brachyptery in Cicadellidae and other insects are briefly reviewed and discussed considering the hypothesis proposed here.

The case of <em>Holopyga gogorzae</em> Trautmann, 1926 and revision of the <em>H. miranda</em> group (Hymenoptera, Chrysididae)

The species treated as Holopyga gogorzae Trautmann, 1926 by Linsenmaier (1959) and subsequent authors was found to be misidentified. The first available name for this West Mediterranean species, thus the valid one, is Holopyga calida Linsenmaier, 1951, herewith raised to specific rank. Surprisingly enough, the true H. gogorzae Trautmann, whose female is still unknown, does not belong to the H. miranda group; it conversely belongs to the H. fervida group, and is most closely related to H. rubra Linsenmaier, 1999 according to genital capsule and tarsal claws. A key to the species of the H. miranda group, and relevant illustrations, are here provided. During this study, Hedychridium planatum Bischoff, 1910 and H. planatum var. auratum Bischoff, 1910 both proved Holopyga, not Hedychridium, most likely synonyms of H. fervida (Fabricius, 1781). The name Holopyga amoenula group is here proposed to replace the name Holopyga gloriosa group given by Linsenmaier (1959), because the name gloriosa Fabricius, 1793 has been suppressed by the ICZN - International Commission on Zoological Nomenclature (1998) and is therefore invalid.

The terminalia of the superfamily Scarabaeoidea (Coleoptera): specific glossary, dissecting methodology, techniques and previously unrecorded sexual dimorphism in some difficult groups

AbstractThe terminology used for the morphological structures of the abdomen as well as male and female genitalia across the superfamily Scarabaeoidea is discussed, based on a literature review across the group. Issues relating to the orientation of the male terminalia and potential homologies between the ‘genital capsule’- and ‘spiculum gastrale’-bearing taxa are discussed and standardisation of terminology pertaining to some ambiguous terms is proposed. A Scarabaeoidea-specific glossary is presented, and synonyms across the literature are given. Schematic illustrations of the abdomen and terminalia are provided. Some new morphological structures are here described for the first time. A standardised dissection protocol for male and female Scarabaeoidea is proposed. Sexually dimorphic structures in the abdomen of Pleurosticti (Dynastinae, Rutelinae, Melolonthinae and Cetoniinae), Glaphyridae, Phaenomeridinae, Passalidae and Omorgus (Trogidae) are described and illustrated for the first time, providing future workers with the possibility to determine the sex of specimens based on external characters without the need for dissection.

Description of the male of Lepidotrigona nitidiventris (Smith, 1857), redescription of the female holotype and additional morphological data on the workers (Hymenoptera: Apidae: Meliponini)

We describe for the first time the male of Lepidotrigona nitidiventris (Smith, 1857), emphasizing the structure of the male genital capsule and metasomal sterna four through seven. Our identification of the male as L. nitidiventris is based on our examination of the worker holotype (type locality Mt. Ophir, Peninsular Malaysia) which we found to match workers from the same nest as males found in western Thailand. The species belongs to the L. nitidiventris species group, comprising L. latipes (Friese), L. palavanica (Cockerell), L. trochanterica (Cockerell), with L. nitidiventris as the type species of the genus. No males of these species were previously known. We examined the holotype workers of the three other species and confirm all four as clearly different species. We propose, however, that the obviously smaller L. palavanica should not be included in the large-sized group of Lepidotrigona.

New Taxa of Neotropical Coreini (Hemiptera: Heteroptera: Coreidae)

Two new genera, Parypheroides new genus, and Sundarellus new genus, and two new species, Anasa albicans new species, from Argentina, and Sundarellus tiputinus new species, from Ecuador are described. Parypheroides is used to accomodate a species previously included in the binomen Paryphes imperialisStal, 1859. Photographs of the dorsal and ventral habitus, a close up of the head and pronotum, genital capsule and parameres of some species are included.

Review of the genus Bubaces (Hemiptera: Heteroptera: Rhyparochromidae: Lethaeini) with descriptions of three new species, new distributional records, and key to species.

The genus Bubaces Distant, 1893 is reviewed and three new species are described: B. gloriosus sp. nov., from Costa Rica (Puntarenas Province) and Mexico (Campeche, Chiapas, Guerrero, Oaxaca, and Veracruz), B. occidentalis sp. nov., from Mexico (Colima, Guerrero, Jalisco, and Michoacan) and B. rostralis sp. nov., from Mexico (Quintana Roo). Additional state faunistic records of the previously described species B. castaneus, B. convergens, B. enatus and B. uhleri are provided. Photographs of the dorsal habitus, the male genital capsule, and the parameres are included, as well as close up photos of the metathoracic scent glands of both sexes. A key to species of the genus is provided.