Evidence of a thoracic crop in workers, soldiers, and queens of Carebara perpusilla ants (Formicidae: Myrmicinae).

The ability to share and store food is paramount in group-living animals, allowing a finely tuned distribution of resources over time and individuals and an enhanced survival over periods of food scarcity. Ants have several ways to store food: one of them is their gastral crop, also known as a "social stomach." Nutrients in the crop can be regurgitated to nestmates through oral trophallaxis (mouth-to-mouth) or proceed to the midgut by opening the proventriculus, a valve connecting the crop to the midgut. However, some ants are also known to have a so-called "thoracic crop," an extension of the esophagus that allows for additional storage space. In this study, we provide the first evidence of a thoracic crop in the genus Carebara, in reproductive (queen) and sterile (soldier and worker) castes. We discuss how the ant body plan allowed for the evolution of a novel food storage structure in the mesothorax.

Complex co-evolutionary relationships between cicadas and their symbionts.

Previous evidence suggests that cicadas lacking Hodgkinia may harbour the yeast-like fungal symbionts (YLS). Here, we reinforce an earlier conclusion that the pathogenic ancestor of YLS independently infected different cicada lineages instead of the common ancestor of Cicadidae. Five independent replacement events in the loss of Hodgkinia/acquisition of YLS and seven other replacement events of YLS (from an Ophiocordyceps fungus to another Ophiocordyceps fungus) are hypothesised to have occurred within the sampled cicada taxa. The divergence time of YLS lineages was later than that of corresponding cicada lineages. The rapid shift of diversification rates of YLS and related cicada-parasitizing Ophiocordyceps began at approximately 32.94 Ma, and the diversification rate reached the highest value at approximately 24.82 Ma, which corresponds to the cooling climate changes at the Eocene-Oligocene boundary and the Oligocene-Miocene transition respectively. Combined with related acquisition/replacement events of YLS occurred during the cooling-climate periods, we hypothesise that the cooling-climate changes impacted the interactions between cicadas and related Ophiocordyceps, which coupled with the unusual life cycle and the differentiation of cicadas may finally led to the diversification of YLS in Cicadidae. Our results contribute to a better understanding of the evolutionary transition of YLS from entomopathogenic fungi in insects.

Linking the Morphology of Sternal Glands to Rubbing Behavior by Vespa soror (Hymenoptera: Vespidae) Workers During Recruitment for Group Predation.

The activities of social insect colonies are supported by exocrine glands and the tremendous functional diversity of the compounds that they secrete. Many social wasps in the subfamilies Vespinae and Polistinae have two sternal glands-the van der Vecht and Richards' glands-that vary in their features and function across the species in which they are found. Field observations suggest that giant hornets use secretions from the van der Vecht gland to chemically mark targeted nests when workers initiate group attacks on social insect prey. However, descriptions of giant hornets' sternal glands and details about their recruitment behavior are lacking. We describe the morphology of the sternal glands of the giant hornet Vespa soror du Buysson and consider their potential to contribute to a marking pheromone. We also assess the gastral rubbing behavior of workers as they attacked Apis cerana F. (Hymenoptera: Apidae) colonies. V. soror workers have well-developed van der Vecht and Richards' glands on their terminal gastral sternites, with morphologies that robustly support the synthesis, storage, and dissemination of their secretory products. Observations confirm that the van der Vecht gland is exposed during gastral rubbing, but that the Richards' gland and glands associated with the sting apparatus may also contribute to a marking pheromone. Workers briefly but repeatedly rubbed their gasters around hive entrances and on overhead vegetation. Colonies were heavily marked over consecutive attacks. Our findings provide insight into the use of exocrine secretions by giant hornets as they recruit nestmates to prey colonies for group attacks.

Structural diversity of symbionts and related cellular mechanisms underlying vertical symbiont transmission in cicadas.

Many insects depend on symbiont(s) for survival. This is particularly the case for sap-feeding hemipteran insects. In this study, we revealed that symbionts harboured in cicadas are diverse and complex, and the yeast-like fungal symbionts (YLS) are present in most cicada species, but Hodgkinia is absent. During vertical transmission, Sulcia became swollen with the outer membrane drastically changed, while Hodgkinia became shrunken and changed from irregular to roughly spherical. Sulcia and/or Hodgkinia were exocytosed from the bacteriocytes to the intercellular space of bacteriomes, where they gathered together and were extruded to haemolymph. YLS and associated facultative symbiont(s) in the fat bodies were released to the haemolymph based on bacteriocyte disintegration. The obligate symbiont(s) were endocytosed and exocytosed successively by the epithelial plug cells of the terminal oocyte, while associated facultative symbiont(s), and possibly also YLS, may take a 'free ride' on the transmission of obligate symbiont(s) to gain entry into the oocyte. Then, the intermixed symbionts formed a characteristic 'symbiont ball' in the oocyte. Our results suggest that YLS in cicadas represent a new example of a relatively early stage of symbiogenesis in insects and contribute to a better understanding of the diversity and transmission mechanisms of symbionts in insects.

Effects of the Argentine ant venom on terrestrial amphibians.

Invasive species have major impacts on biodiversity and are one of the primary causes of amphibian decline and extinction. Unlike other top ant invaders that negatively affect larger fauna via chemical defensive compounds, the Argentine ant (Linepithema humile) does not have a functional sting. Nonetheless, it deploys defensive compounds against competitors and adversaries. We estimated levels of ant aggression toward 3 native terrestrial amphibians by challenging juveniles in field ant trails and in lab ant foraging arenas. We measured the composition and quantities of toxin in L. humile by analyzing pygidial glands and whole-body contents. We examined the mechanisms of toxicity in juvenile amphibians by quantifying the toxin in amphibian tissues, searching for histological damages, and calculating toxic doses for each amphibian species. To determine the potential scope of the threat to amphibians, we used global databases to estimate the number, ranges, and conservation status of terrestrial amphibian species with ranges that overlap those of L. humile. Juvenile amphibians co-occurring spatially and temporally with L. humile die when they encounter L. humile on an ant trail. In the lab, when a juvenile amphibian came in contact with L. humile the ants reacted quickly to spray pygidial-gland venom onto the juveniles. Iridomyrmecin was the toxic compound in the spray. Following absorption, it accumulated in brain, kidney, and liver tissue. Toxic dose for amphibian was species dependent. Worldwide, an estimated 817 terrestrial amphibian species overlap in range with L. humile, and 6.2% of them are classified as threatened. Our findings highlight the high potential of L. humile venom to negatively affect amphibian juveniles and provide a basis for exploring the largely overlooked impacts this ant has in its wide invasive range.

Efectos del Veneno de la Hormiga Argentina sobre los Anfibios Terrestres Resumen Las especies invasoras tienen un impacto importante sobre la biodiversidad y son una de las causas principales del declive y extinción de los anfibios. A diferencia de otras hormigas super-invasoras que afectan negativamente a animales más grandes por medio de compuestos químicos de defensa, la hormiga argentina (Linepithema humile) no tiene unaguijón funcional. Sin embargo, esta hormiga despliega compuestos defensivos contra sus competidores y adversarios. Estimamos los niveles de agresión de las hormigas hacia tres anfibios terrestres nativos exponiendo a los anfibios juveniles en pistas de hormigas en el campo y en las arenas de forrajeo de las hormigas en el laboratorio. Medimos la composición y las cantidades de toxina que presenta L. humile por medio del análisis de las glándulas pigidiales y el contenido en el cuerpo completo. Examinamos los mecanismos de la toxicidad en los anfibios juveniles cuantificando la toxina en el tejido del anfibio, buscando daños histológicos y calculando las dosis tóxicas para cada especie de anfibio. Para determinar el alcance potencial de la amenaza para los anfibios usamos bases de datos mundiales para estimar el número, distribución y estado de conservación de las especies terrestres de anfibios con distribuciones que se solapan con la de L. humile. Los anfibios juveniles que co-ocurren temporal y espacialmente con L. humile mueren al encontrarse con esta especie de hormiga en sus pistas. En el laboratorio, cuando un anfibio juvenil entró en contacto con L. humile, las hormigas reaccionaron rápidamente rociando a estos juveniles con veneno proveniente de las glándulas pigidiales. La iridomyrmecina fue el compuesto tóxico que encontramos en las glándulas pigidiales. Después de ser absorbida por la piel del anfibio, se acumuló en el cerebro, los riñones y el hígado. La dosis tóxica para los anfibios depende de la especie. A nivel mundial, se estima que 817 especies de anfibios terrestres tienen una distribución que se solapa con la de L. humile, y el 6.2% de estas especies se encuentran clasificadas como amenazadas. Nuestros hallazgos resaltan el potencial alto del veneno de L. humile para tener efectos negativos sobre los anfibios juveniles y también proporcionan una base para la exploración de los impactos de esta hormiga en su amplio rango invasivo, los cuales generalmente son ignorados.

Skeletomuscular adaptations of head and legs of Melissotarsus ants for tunnelling through living wood.

BACKGROUND: While thousands of ant species are arboreal, very few are able to chew and tunnel through living wood. Ants of the genus Melissotarsus (subfamily Myrmicinae) inhabit tunnel systems excavated under the bark of living trees, where they keep large numbers of symbiotic armoured scale insects (family Diaspididae). Construction of these tunnels by chewing through healthy wood requires tremendous power, but the adaptations that give Melissotarsus these abilities are unclear. Here, we investigate the morphology of the musculoskeletal system of Melissotarsus using histology, scanning electron microscopy, X-ray spectrometry, X-ray microcomputed tomography (micro-CT), and 3D modelling. RESULTS: Both the head and legs of Melissotarsus workers contain novel skeletomuscular adaptations to increase their ability to tunnel through living wood. The head is greatly enlarged dorsoventrally, with large mandibular closer muscles occupying most of the dorsal half of the head cavity, while ventrally-located opener muscles are also exceptionally large. This differs from the strong closing: opening asymmetry typical of most mandibulated animals, where closing the mandibles requires more force than opening. Furthermore, the mandibles are short and cone-shaped with a wide articulatory base that concentrates the force generated by the muscles towards the tips. The increased distance between the axis of mandibular rotation and the points of muscle insertion provides a mechanical advantage that amplifies the force from the closer and opener muscles. We suggest that the uncommonly strong opening action is required to move away crushed plant tissues during tunnelling and allow a steady forward motion. X-ray spectrometry showed that the tip of the mandibles is reinforced with zinc. Workers in this genus have aberrant legs, including mid- and hindlegs with hypertrophied coxae and stout basitarsi equipped with peg-like setae, and midleg femura pointed upward and close to the body. This unusual design famously prevents them from standing and walking on a normal two-dimensional surface. We reinterpret these unique traits as modifications to brace the body during tunnelling rather than locomotion per se. CONCLUSIONS: Melissotarsus represents an extraordinary case study of how the adaptation to - and indeed engineering of - a novel ecological niche can lead to the evolutionary redesign of core biomechanical systems.

Distinctive fungal communities in an obligate African ant-plant mutualism.

Three ant species nest obligately in the swollen-thorn domatia of the African ant-plant Vachellia (Acacia) drepanolobium, a model system for the study of ant-defence mutualisms and species coexistence. Here we report on the characteristic fungal communities generated by these ant species in their domatia. First, we describe behavioural differences between the ant species when presented with a cultured fungal isolate in the laboratory. Second, we use DNA metabarcoding to show that each ant species has a distinctive fungal community in its domatia, and that these communities remain characteristic of the ant species over two Kenyan sampling locations separated by 190 km. Third, we find that DNA extracted from female alates of Tetraponera penzigi and Crematogaster nigriceps contained matches for most of the fungal metabarcodes from those ant species' domatia, respectively. Fungal hyphae and other debris are also visible in sections of these alates' infrabuccal pockets. Collectively, our results indicate that domatium fungal communities are associated with the ant species occupying the tree. To the best of our knowledge, this is the first record of such ant-specific fungal community-level differences on the same myrmecophytic host species. These differences may be shaped by ant behaviour in the domatia, and by ants vectoring fungi when they disperse to establish new colonies. The roles of the fungi with respect to the ants and their host plant remain to be determined.

Worker reproduction of the invasive yellow crazy ant Anoplolepis gracilipes.

BACKGROUND: Reproductive division of labor is one of the key features of social insects. Queens are adapted for reproduction while workers are adapted for foraging and colony maintenance. In many species, however, workers retain functional ovaries and can lay unfertilized male eggs or trophic eggs. Here we report for the first time on the occurrence of physogastric workers and apparent worker reproduction in the invasive yellow crazy ant Anoplolepis gracilipes (Fr. Smith). We further examined the reproductive potential and nutritional role of physogastric workers through multidisciplinary approaches including morphological characterization, laboratory manipulation, genetic analysis and behavioral observation. RESULTS: Egg production with two types of eggs, namely reproductive and trophic eggs, by physogastric workers was found. The reproductive egg was confirmed to be haploid and male-destined, suggesting that the workers produced males via arrhenotokous parthenogenesis as no spermatheca was discovered. Detailed observations suggested that larvae were mainly fed with trophic eggs. Along with consumption of trophic eggs by queens and other castes as part of their diet, the vital role of physogastric workers as "trophic specialist" is confirmed. CONCLUSION: We propose that adaptive advantages derived from worker reproduction for A. gracilipes may include 1) trophic eggs provisioned by physogastric workers likely assist colonies of A. gracilipes in overcoming unfavorable conditions such as paucity of food during critical founding stage; 2) worker-produced males are fertile and thus might offer an inclusive fitness advantage for the doomed orphaned colony.

Metformin promotes lifespan through mitohormesis via the peroxiredoxin PRDX-2.

The antiglycemic drug metformin, widely prescribed as first-line treatment of type II diabetes mellitus, has lifespan-extending properties. Precisely how this is achieved remains unclear. Via a quantitative proteomics approach using the model organism Caenorhabditis elegans, we gained molecular understanding of the physiological changes elicited by metformin exposure, including changes in branched-chain amino acid catabolism and cuticle maintenance. We show that metformin extends lifespan through the process of mitohormesis and propose a signaling cascade in which metformin-induced production of reactive oxygen species increases overall life expectancy. We further address an important issue in aging research, wherein so far, the key molecular link that translates the reactive oxygen species signal into a prolongevity cue remained elusive. We show that this beneficial signal of the mitohormetic pathway is propagated by the peroxiredoxin PRDX-2. Because of its evolutionary conservation, peroxiredoxin signaling might underlie a general principle of prolongevity signaling.

Degeneration patterns of the worker spermatheca during morphogenesis in ants (Hymenoptera: Formicidae).

Reproductive division of labor is one of the crucial features in social insects, however, the developmental mechanisms leading to modifications in the reproductive apparatus of workers are still not very clear. Ants show a remarkable diversity in the morphological specialization of the worker's reproductive apparatus, that allows to distinguish four types, type 1: workers that have ovaries and a functional spermatheca, and that reproduce like queens, type 2: workers have ovaries and a vestigial spermatheca, type 3: workers have ovaries but no spermatheca, and type 4: workers lost both ovaries and spermatheca. We investigated morphogenesis of the worker spermatheca in 28 ant species by histological examination. In workers of type 1, the morphogenesis of the spermatheca is very similar to that in ant queens. In type 2, the spermathecal disc also differentiates, however, the development is interrupted and remains vestigial. In types 3 and 4, the absence of the spermatheca in the adult phase is caused by a degeneration after initial formation of the spermathecal disc or by a complete lack of the spermathecal discs. The timing of these interruption and degeneration events varies among species. The species exhibiting an earlier interrupting point of spermatheca formation in workers have a larger queen-worker dimorphism, that seems to be independent from ant phylogeny.

Parthenogenetic Reproduction in Strumigenys Ants: An Update.

Parthenogenetic reproduction is a common feature for social Hymenoptera, as males typically develop from unfertilized eggs (arrhenotoky). Production of female offspring without the involvement of sperm (thelytoky) also exists but is rather exceptional as it has been reported for only 16 ant species so far. Three of these belong to the genus Strumigenys: S. hexamera, S. membranifera and S. rogeri. Our observations on the reproductive biology in various Oriental Strumigenys species extends this list of thelytokous ants with three more species: S. emmae, S. liukueiensis and S. solifontis. Of these six thelotykous species, S. emmae, S. membranifera and S. rogeri are known as tramp species. Reproduction without the need to fertilize eggs no doubt offers these species a considerable advantage when establishing colonies in new environments. Published histological data on S. hexamera and S. membranifera already showed that the queens possess a functional spermatheca. We now provide evidence that this is also the case for the four other thelytokous Strumigenys species. Retaining a functional spermatheca and reproductive system may keep the queens ready for the exceptional event of mating and hence increase genetic variability, as males do occur very rarely.

Comparative morphology and ultrastructure of the labial gland among castes of Camponotus japonicus (Hymenoptera: Formicidae).

The labial gland is a major exocrine gland in the thoracic cavity of ants, which is connected to the mouth area and opens at the base of the labium. The labial gland in the various castes and males of Camponotus japonicus were examined using dissection, light microscopy, scanning and transmission electron microscopy. Our findings reveal clear caste differences in the appearance of the labial gland, and suggest a relationship between gland structure and its behavioral role in queens, males and workers: queens possess the largest reservoir sac; males have the smallest; workers have abundant secretion droplets in the gland cells of the tubules; epithelial thickness of gland cells varies considerably between workers and reproductive ants. The apical cell region has a high density of microvilli and polymorphic mitochondria, whereas the central cell region is rich in rough endoplasmic reticulum (RER) which means its main secretions are proteinaceous compounds. We believe that the labial gland has a different function in the castes of C. japonicus: the gland of workers and queens may be used to feed the larvae; workers participate in trophallaxis in the nest, while the males are not involved in these activities. Calculations of the relative size of the labial gland for each caste and males indicated that minor workers have the relative highest developed labial gland, supporting a role in trophallaxis. In addition, we found a phenomenon of enlarged labial gland in minor workers, which accounted for almost 22%, but the reason for this is unknown.

Morphology and ultrastructure of the mandibular gland in Camponotus japonicus.

The mandibular gland is an important exocrine gland connected to the mandibles. In ants, it plays a crucial role in alarm communication and reproduction. While the overall morphology of the mandibular gland does not vary much between ant species, significant differences exist in the ultrastructural characteristics and gland size in some reported cases. However, there have been few systematic comparative studies on this subject. In this paper, we conducted microscopic observations to analyze the morphological and ultrastructural similarities and differences of the mandibular gland in different castes of Camponotus japonicus. Our findings revealed that the mandibular glands of males were significantly larger than those of females. In terms of relative size, within the female castes the mandibular glands of minor workers were the largest. Additionally, our observations of the ultrastructure showed that secretory substances were most abundant in queens. All female castes and males showed the presence of crystalline structures in the secretory cells. The content of these biocrystals decreased from minor workers to major workers, queens, and males. In males, however, crystalline structures were rare and the most abundant substances are droplet-like secretions. The differences in morphology and ultrastructure between different castes suggest that the function of the mandibular gland varies among castes. In conclusion, our study provides insights into the morphological and ultrastructural characteristics of the mandibular gland in different castes of C. japonicus. Further research is needed to fully understand the specific functions and mechanisms of this gland in ants.

Morphology of the metapleural gland and its associated novel atrial cone gland in Strumigenys ants.

The metapleural gland is a unique feature of the ant phenotype, but diversity in its anatomy and function across the ants is not well documented or understood. We studied the morphology of the metapleural gland in 20 mainly Oriental Strumigenys species using histology, scanning and transmission electron microscopy, and micro-CT. The gland is formed by a cluster of class-3 cells, their secretion is transported through a bundle of ducts into the sclerotized atrium and is guided along a series of parallel cuticular ridges towards the atrial opening. Among the examined species, queens have more gland cells than conspecific workers, while the examined males do not have the gland. The social parasite S. mutica has the most developed metapleural gland. In addition, we describe a novel class-1 atrial cone gland which is associated with the metapleural gland. The epithelium of this cone gland forms an invagination into the ventral atrium of the metapleural gland. The cuticular cone may be lacking in some Afrotropical and Neotropical species, although these may still contain the epithelial gland. The functional and evolutionary drivers of morphological variation in the exocrine system across species form interesting questions for future work.

The spongiform tissue in Strumigenys ants contains exocrine glands.

The insect cuticle is multifunctional and often includes projections used for support, communication or protection. Ants in the genus Strumigenys exhibit a peculiar honeycomb-like spongiform tissue that covers their petiole, postpetiole and sometimes also the posterior mesosoma and anterior part of the first gastral segment. The tissue is abundantly developed in workers and queens, and much reduced in males. We found this spongiform tissue is associated with a novel exocrine gland that is made up by class-3 secretory cells that are clustered underneath the major pillars of the cuticular extensions, their associated narrow ducts enter these extensions and open at the surface through small pores. The chemical nature and function of the secretion are still unknown. The honeycomb texture may act in the storage and dispersion of the glandular secretions. In addition to the spongiform tissue gland, the posterior region of the petiole and postpetiole also contain intersegmental petiole and postpetiole glands, of which the ducts open through the intersegmental membrane that forms the connection with the next segment. Future work aimed at identifying the chemicals secreted by these glands will shed light onto the function of these unusual structures.

Histology and ultrastructure of glands associated with the sting apparatus of the swarm-founding wasp Protopolybia exigua (Hymenoptera: Vespidae: Polistinae).

Social insects present a great diversity of exocrine glands, which are linked to fundamental roles of social life, and their morphological characterization represents the first step toward the knowledge of their function. We here describe the exocrine glands present in the sting apparatus of queens and workers of the wasp Protopolybia exigua. Histological analysis of serial sections under light microscopy revealed a glandular epithelium in the spiracular plate, the quadrate plate, and the gonostyli, while the last two in addition also contain clusters of class-3 cells. A big cluster of class-3 cells was also found in the oblong plate of some workers and queens. These findings reveal that a variety of glands, in addition to the common venom gland and Dufour gland, are present in association with the sting apparatus, of which three glands are novel reports for the exocrine repertoire of social insects. The position of the glands suggests a role in lubrication to provide mobility of the sting and venom ejection and/or participation in alarm behavior, while a function related to reproduction is also possible. The peculiar characteristics displayed by swarm-founding wasps are linked with exocrine gland secretions, making their investigation in Epiponini wasps a vast and interesting field.

Morphology and ultrastructure of the infrabuccal pocket in Strumigenys ants.

The morphology of the infrabuccal pocket has been studied with light and electron microscopy (SEM, TEM) in 19 species of Strumigenys ants. The structural organization is similar in workers, queens and males, and supports the involvement of the pocket in the filtration of food particles before they can enter the digestive tract. A carpet of posteriorly oriented bristle hairs on the hypopharynx first guide ingested food into the pocket, where large solid particles are compacted into a pellet that will be regurgitated. The remaining products enter the digestive tract through a filtering wall of parallel hair combs lining the pharynx interior that are directed against the food flow. This mechanical filtering allows only liquids and sufficiently small food particles to enter the digestive system. The wall of the infrabuccal pocket is differentiated into a conspicuous glandular epithelium, of which the ultrastructural characteristics can be understood as an adaptation against the frequent shape changes of the pocket. The gland elaborates a non-proteinaceous secretion, although its functional significance still remains unknown.

Comparison of the External Morphology of the Sternal Glands for Hornets in the Genus Vespa.

Many social wasps in the speciose subfamilies Polistinae and Vespinae have two sternal glands-the van der Vecht gland and the Richards gland-that are not found in other insects. The presence of these glands has been confirmed in only 6 of 22 hornet species (genus Vespa) and images of their fine structure have not been produced. Here; we characterize the external morphology associated with both glands for workers of nine Vespa species using scanning electron microscopy. All hornets had similar gland configurations; although gland-associated external features differed among species. Scaled for size, glands were equivalently sized for the giant hornets (V. mandarinia and V. soror) and their closest phylogenetic relatives (V. tropica and V. ducalis). Relative size of gland-associated structures was reduced by half for V. simillima; V. velutina; and V. affinis workers. The remaining species (V. crabro and V. analis) had intermediately sized features. Differences among species in external gland structure were best explained by selective pressures related to predatory behavior, rather than defense of nests against ants. However, a lack of information about how Vespa workers use their van der Vecht and Richards glands limits a comparative interpretation of the function of their external gland morphology.

The ant abdomen: The skeletomuscular and soft tissue anatomy of Amblyopone australis workers (Hymenoptera: Formicidae).

Recent studies of insect anatomy evince a trend towards a comprehensive and integrative investigation of individual traits and their evolutionary relationships. The abdomen of ants, however, remains critically understudied. To address this shortcoming, we describe the abdominal anatomy of Amblyopone australis Erichson, using a multimodal approach combining manual dissection, histology, and microcomputed tomography. We focus on skeletomusculature, but additionally describe the metapleural and metasomal exocrine glands, and the morphology of the circulatory, digestive, reproductive, and nervous systems. We describe the muscles of the dorsal vessel and the ducts of the venom and Dufour's gland, and characterize the visceral anal musculature. Through comparison with other major ant lineages, apoid wasps, and other hymenopteran outgroups, we provide a first approximation of the complete abdominal skeletomuscular groundplan in Formicidae, with a nomenclatural schema generally applicable to the hexapod abdomen. All skeletal muscles were identifiable with their homologs, while we observe potential apomorphies in the pregenital skeleton and the sting musculature. Specifically, we propose the eighth coxocoxal muscle as an ant synapomorphy; we consider possible transformation series contributing to the distribution of states of the sternal apodemes in ants, Hymenoptera, and Hexapoda; and we address the possibly synapomorphic loss of the seventh sternal-eighth gonapophyseal muscles in the vespiform Aculeata. We homologize the ovipositor muscles across Hymenoptera, and summarize demonstrated and hypothetical muscle functions across the abdomen. We also give a new interpretation of the proximal processes of gonapophyses VIII and the ventromedial processes of gonocoxites IX, and make nomenclatural suggestions in the context of evolutionary anatomy and ontology. Finally, we discuss the utility of techniques applied and emphasize the value of primary anatomical research.

Morphology and ultrastructure of the prepharyngeal and pharyngeal glands in the ant Camponotus japonicus.

The prepharyngeal gland (prePG) and pharyngeal gland (PG) make up the largest exocrine structures in the head of the ant Camponotus japonicus. We used microscopy to study the histological and ultrastructural features of both glands in different castes. The number of secretory units in the prePG is considerably higher than in other ant species and shows a complex duct system which is made up by duct cells, secondary ducts and a main duct. These lead the secretions of hundreds to thousands of secretory cells into the prepharynx through a modified sieve plate at each side. The glove-shaped PG shows clear caste differences in tubule number. The ultrastructure of both the prePG and PG shows abundant mitochondria and secretion vesicles. Moreover, the prePG is loaded with rough endoplasmic reticulum (RER) which means its main secretions are proteinaceous compounds, while the PG is dominated by smooth endoplasmic reticulum (SER) which means the main secretions are lipids. The morphological differences like cell number of the prePG and tubule number of the PG indicate different secretory abilities of each caste. We for the first time introduce histology-based relative size to indicate secretory activity. The proportionally high development of the prePG in minor workers supports a role in trophallaxis.