Effects of Cephenemyia spp. (Diptera: Oestridae) on the nasopharynx of black-tailed deer (Odocoileus hemionus columbianus)

A study was conducted to determine gross and microscopic tissue changes in the nasopharynx of black-tailed deer (Odocoileus hemionus columbianus) infected with nasal bot fly larvae (Cephenemyia spp.). Paired retropharyngeal recesses were the preferred sites for the growing second and third stage larvae of two species of Cephenemyia (C. apicata and C. jellisoni). Retropharyngeal recesses distended into "pouches" that harbored up to 30 larvae. Pouches were oriented caudal-laterally toward the basisphenoid bone of the cranium. Lateral support of the pouch mass was provided by the stylohyoid bone. The laryngeal orifice was never occluded by the enlarged recesses. The distal pouch wall was relatively thin and remained uniform in thickness as expansion progressed. Occasionally, aberrant larvae were found protruding through the distal wall of the pouch. Disruption of the epithelium and submucosa by larval mouth hooks and integumentary spines were examined by scanning electron microscopy. Histological examination of infected recesses revealed substantial loss of epithelium and mucous glands. Enlargement of recesses into pouches resulted from fibrosis. Healing occurred after larvae egressed from the pouches. Degenerating mucous glands, epithelial metaplasia, epithelial desquamation, and intense inflammation were found near larvae. An eosinophilic exudate with a mixture of macrophages and erythrocytes was present in the lumen of the pouch. The presence of larvae within the pouch inhibited secondary bacterial infection and suppuration. Infection by larvae caused severe local trauma and intense tissue response.     

Metamorphic changes in the hindgut of Sarcophaga ruficornis with special reference to the development of rectal pads

In Sarcophaga ruficornis, during normal metamorphosis the larval hindgut epithelium degenerates and the imaginal hindgut epithelium is formed from the proliferation and differentiation of the cells of the posterior imaginal ring. The newly formed hindgut becomes slightly dilated posteriorly to form the rectal pouch. At four places in this pouch, the epithelium becomes thick and forms the primordia of the rectal pads. The cells in the primordia begin to divide and project into the lumen of the pouch and later form the cortex cells of the rectal pads. The cavity of the primordia becomes filled with blood cells and fibrous material, which give rise to the medulla of the rectal pads.     

Comparative studies on the structure of reproductive organs of four botryllid ascidians

Reproductive organs of four botryllid ascidians, Botryllus primigenus, Botryllus schlosseri, Botrylloides violaceus and Botrylloides leachi, were studied histologically. In every species, the egg follicle consisting of an egg and its inner and outer follicles, is attached to the follicle stalk, the vesicle being composed of a flat epithelium, which in its turn is connected to the atrial epithelium or to the brood pouch specialized from it. In B. schlosseri, the egg is ovulated into the atrial cavity and remains there held by the brood cup, of which the inner epithelium is derived from the follicle stalk and the outer one from the atrial epithelium. In B. primigenus, the brood pouch develops as a diverticulum of the atrial cavity, around the entrance of which a fold differentiates from the atrial epithelium and closes the pouch during embryogenesis. In both species of Botrylloides, the brood pouch is formed by the outgrowth of the thickened atrial epithelium into the blood space, the entrance of which is closed during embryogenesis. The discarded outer follicle completely disintegrates soon after ovulation in B. schlosseri, but part of it remains throughout embryogenesis in the blood space in B. primigenus or projecting into the interior of the brood pouch in Botrylloides. In primigenus, the testis, when it accompanies the egg follicle, is placed at the bottom of the brood pouch and the sperm is shed through the pouch prior to ovulation. In B. schlosseri and the Botrylloides species, the testis is located independently from the egg follicle and the sperm matures after ovulation.     

The basicoxal gland, a new exocrine structure in poneromorph ants (Hymenoptera, Formicidae)

The tegumental epithelium of the outer dorsolateral region in the proximal part of the coxae in the mid‐ and hindlegs of both workers and queens of the ants Odontomachus rixosus and O. simillimus is differentiated into a conspicuous and hitherto unknown exocrine gland. The glandular cells display a clear microvillar differentiation of their apical cell membrane, and are lined with the tegumental cuticle, which in this part contains crack‐like channels perpendicular to its surface, that carry the glandular secretions to the outside. Apical microvilli support the transport of substances, and contain an extension of tubular smooth endoplasmic reticulum in their centre. The function of the gland may be that of providing lubricant substances to the articulation region of the generally heavily sclerotized ponerine ant species. The gland is also found in several other ponerine and amblyoponine species, but not in the ectatommine species studied. The foreleg coxae lack a basicoxal gland in all species examined, which may be explained by the more limited articulation between the thorax and the coxae in the forelegs compared to the mid‐ and hindlegs.     

Postnatal development of vagina, clitoris and urethral glands of the golden hamster

We have made a histological study of the postnatal development of the clitoris, preputial glands, urethral glands and vagina of the golden hamster. The 'phallic groove' of the clitoris is closed at 10 days of life, then the urethra has a cuboidal stratified, a stratified squamous and a stratified keratinized epithelium. The preputial glands are composed of branched saccular glands. These glands develop, with few changes during their maturation period. Formation of the urethral glands begins at 5 days and the alveoli are fully developed at puberty. The hamster vagina has two origins; the upper part is Müllerian, the caudal part is sinusal. The wall of the Müllerian vagina has a cylindrical epithelium at birth, which becomes 'double epithelium' at puberty and thereafter changes cyclically in connection with the estrous cycle. The sinusal vagina is solid at birth, its lumen being formed in the first 10 days of life and its wall having a cuboidal stratified epithelium. At 15 days it becomes a stratified keratinized epithelium, which will later line the vaginal pouch. At the 5th day, an ectodermic invagination (stratified keratinized epithelium) is observed in the zone of the future introitus. At the time of vaginal opening this zone forms the distal segment.     

The characteristics of the initial link in the intestinal lesion by opportunistic Proteus mirabilis bacteria]

Interaction of Proteus mirabilis with the intestine epithelium in white mongrel mice has been studied by means of bacteriological, toxicological and electron microscopic methods. Introduction of the agar culture of bacteria to the intestine lumen has permitted finding its general toxic action on enterocytes and other cell elements of epithelium which was accompanied by a potentiation of the striated margin membrane degradation but caused no destruction of cells. A damaging factor of protei was connected with the thermostable large-molecular fraction of cultural fluid of bacteria and its action was traced at the first hours of the bacterial contact with the mucous surface of thin intestine. Preliminary weakening of protective barriers of the intestine mucosa permitted protei to penetrate deep into the tissue through the intercellular slots. Colonization of the intestine lumen wall zone by the proteus is a necessary condition of the above-mentioned changes.     

Functional morphology of the mantle of Nautilus pompilius (Mollusca, Cephalopoda)

This study presents histological and cytological findings on the structural differentiation of the mantle of Nautilus pompilius in order to characterize the cells that are responsible for shell formation. The lateral and front mantle edges split distally into three folds: an outer, middle, and inner fold. Within the upper part of the mantle the mantle edge is divided into two folds only; the inner fold disappears where the hood is attached to the mantle. At the base of the outer fold of the lateral and front mantle edge an endo-epithelial gland, the mantle edge gland, is localized. The gland cells are distinguished by a distinct rough endoplasmic reticulum and by numerous secretory vesicles. Furthermore, they show a strong accumulation of calcium compounds, indicating that the formation of the shell takes place in this region of the mantle. Numerous synaptic contacts between the gland cells and the axons of the nerve fibers reveal that the secretion in the area of the mantle edge gland is under nervous control. The whole mantle tissue is covered with a columnar epithelium having a microvillar border. The analyses of the outer epithelium show ultrastructural characteristics of a transport active epithelium, indicating that this region of the mantle is involved in the sclerotization of the shell. Ultrastructural findings concerning the epithelium between the outer and middle fold suggest that the periostracum is formed in this area of the mantle, as it is in other conchiferan molluscs.     

Fine structural organization of the rectum in the blowfly, Calliphora erythrocephala (Meig.) with special reference to connective tissue, tracheae and neurosecretory innervation in the rectal papillae

The rectal epithelium of Calliphora is made up of three quite distinct cell types: rectal, cortical and junctional cells. The thin wall of the rectal pouch is made up of rectal cells which are relatively simple and unspecialized; their general structure does not suggest any direct participation in ion transport. A function of ion and water transport can probably be ascribed to the cortical cells, which are arranged in the form of four cones which project into the rectal lumen. The cavity of each cone is filled up with tracheae, nerve and neurosecretory terminals, and connective tissue to form medulla. The medulla and cortex are separated from each other by deeply staining bridges or trabeculae to form an infundibular space. The most conspicuous feature of the cortex is the presence of an extensive intercellular sinus formed by complex infoldings of the lateral plasma-membranes. It is postulated that fluid, which is absorbed from the rectal lumen, is transported into the intercellular sinus and finally reaches the haemolymph via the infundibular space. The actual site of ion transport is probably the stacks of lateral plasma-membrane which are closely associated with mitochondria. The junctional cells, which are packed with microtubules, form a collar around the base of the papillae at the point of their insertion into the rectal wall. It is suggested that the neurosecretory terminals present in the medulla might release a hormone which controls rate of ion and water reabsorption by the papillae cells.     

Light and electron microscopic study of the hindgut of the ant (Formica nigricans, hymenoptera): II. Structure of the rectum

The rectum of the ant Formica nigricans is composed of six ovoid rectal papillae inserted into a rectal pouch. The wall of the rectal pouch is made up of a flat epithelium of simple rectal cells lined by cuticle, and surrounded by a circular muscle layer. Each rectal papilla is comprised by a simple columnar epithelium of principal cells facing the lumen, and a simple cuboid epithelium of secondary cells towards the hemolymph; a group of 20-25 slender junctional cells lies laterally between both epithelia enclosing an intrapapillar sinus. The muscle layer of the rectal wall also surrounds the base of the papillae. Principal cells do not exhibit extensive infoldings at the apical and basal plasma membranes. Lateral membranes, in contrast, develop highly folded mitochondria-scalariform junction complexes enclosing very narrow intercellular canaliculi between adjacent cells. These canaliculi open to wider intercellular sinuses that ultimately drain into the intrapapillar sinus at the sites of entry of tracheal cells. The lateral plasma membranes do not link to the apical or basal plasma membrane, thus originating a syncytium throughout the principal cells. The apical plasma membrane of secondary cells shows invaginations in relation with an apical tubulovacuolar system, bearing portasomes to the cytoplasmic side of the membrane. Secondary cells unite by convoluted septate junctions, and basolateral infoldings are also developed. These ultrastructural traits, some of them different from those found in other insects, are discussed and examined in relation to their role in water and solute absorption. A route for rectal transport in F. nigricans is proposed.     

Crypt architecture of tonsilla lingualis in the monkey,Macaca fascicularis

Summary Crypts of the lingual tonsil were investigated in 10 male and female Macaca fascicularis by use of correlated light and scanning-electron microscopy. Counting of crypt openings provided an estimate of the total number of respective crypto-lymphatic units, which were found to range from 20 to 39. Crypt openings appeared in three distinct morphological varieties, i.e. circular, oval or slit-like. Tonsillar units existed individually or were arranged in a rosary fashion below a slit-like mucosal fold serving as a common exit. Although the crypt epithelium was generally non-keratinized, individual cells showing a surface pattern similar to that of the keratinized cells could be encountered. The crypt epithelium was frequently fragmented and showed heavy mononuclear cell infiltration and surface discontinuities, with lymphoid cells coming in contact with luminal contents. The crypt lumen either appeared as a simple epithelial invagination or existed as a complex, cavernous pouch with many blind-ending diverticula. The lumen contained a mixture of exfoliated epithelial cells, leucocytes and bacteria. The secretory ducts of the posterior lingual glands opened occasionally at various levels into the crypt lumina or independently to the exterior.     

Ovoviviparity and the structure of the brood pouch in Melanoides tuberculata (Gastropoda: Prosobranchia: Thiaridae)

The freshwater gastropod Melanoides tuberculata broods its young in a pouch located in the anterodorsal region of the head-foot. The wall of the brood pouch is composed of smooth muscle surrounded by connective tissue. The lumen of the brood pouch is incompletely partitioned by trabeculae, formed by extensions or folds in the chamber wall that are composed of smooth muscle, connective tissue, nonciliated squamous epithelial cells, and some storage cells containing lipid and glycogen. The lumen of the chamber also contains a few cells with storage products. The general absence of secretory cells suggests that embryos derive little nutrition from the mother, and therefore embryonic development is probably ovoviviparous. Embryos in various stages of development were found within brood pouches, with later stage embryos varying in size. There was a negative relationship between embryo size and number of embryos in the brood pouch.     

The structured diversity of specialized gut symbionts of the New World army ants

Symbiotic bacteria play important roles in the biology of their arthropod hosts. Yet the microbiota of many diverse and influential groups remain understudied, resulting in a paucity of information on the fidelities and histories of these associations. Motivated by prior findings from a smaller scale, 16S rRNA‐based study, we conducted a broad phylogenetic and geographic survey of microbial communities in the ecologically dominant New World army ants (Formicidae: Dorylinae). Amplicon sequencing of the 16S rRNA gene across 28 species spanning the five New World genera showed that the microbial communities of army ants consist of very few common and abundant bacterial species. The two most abundant microbes, referred to as Unclassified Firmicutes and Unclassified Entomoplasmatales, appear to be specialized army ant associates that dominate microbial communities in the gut lumen of three host genera, Eciton, Labidus and Nomamyrmex. Both are present in other army ant genera, including those from the Old World, suggesting that army ant symbioses date back to the Cretaceous. Extensive sequencing of bacterial protein‐coding genes revealed multiple strains of these symbionts coexisting within colonies, but seldom within the same individual ant. Bacterial strains formed multiple host species‐specific lineages on phylogenies, which often grouped strains from distant geographic locations. These patterns deviate from those seen in other social insects and raise intriguing questions about the influence of army ant colony swarm‐founding and within‐colony genetic diversity on strain coexistence, and the effects of hosting a diverse suite of symbiont strains on colony ecology.     

Evolution of specialized spermatheca morphology in ant queens: Insight from comparative developmental biology between ants and polistine wasps

In many ant species, the queens can keep spermatozoa alive in their spermatheca for several years, which goes along with unique morphological characteristics of the queen's spermatheca. The relative spermatheca size in ant queens is prominently larger than that in social wasps. Furthermore, the epithelium lining the spermatheca reservoir of ants consists of columnar cells in the hilar region and squamous cells in the distal region, whereas it is formed by columnar cells only in social wasps. To study the evolution of the unique spermatheca morphology in ant queens, we compared the various processes during spermatheca development between two ponerine ant species of the genus Pachycondyla (= Brachyponera) and three polistine wasp species of the genus Polistes. From histological observations, we can define four developmental events in the ant queens: (1) invagination of the spermatheca primordium, (2) the reservoir wall thickness becomes unequal, (3) the reservoir diameter doubles as the lining epithelial cells become flattened except for the hilar region, and (4) the increase in thickness of the reservoir epithelium is limited to the hilar region which doubles in thickness. In polistine wasps, the second and the third developmental events are absent and the entire epithelium of the spermatheca wall becomes thick in the final step. We therefore conclude that for ant queens the second and third steps are crucial for the enlargement of the spermatheca size, and that the second to the fourth steps are crucial for the specialization of the reservoir wall structure.     

Disentangling the assembly mechanisms of ant cuticular bacterial communities of two Amazonian ant species sharing a common arboreal nest

Bacteria living on the cuticle of ants are generally studied for their protective role against pathogens, especially in the clade of fungus‐growing ants. However, little is known regarding the diversity of cuticular bacteria in other ant host species, as well as the mechanisms leading to the composition of these communities. Here, we used 16S rRNA gene amplicon sequencing to study the influence of host species, species interactions and the pool of bacteria from the environment on the assembly of cuticular bacterial communities on two phylogenetically distant Amazonian ant species that frequently nest together inside the roots system of epiphytic plants, Camponotus femoratus and Crematogaster levior. Our results show that (a) the vast majority of the bacterial community on the cuticle is shared with the nest, suggesting that most bacteria on the cuticle are acquired through environmental acquisition, (b) 5.2% and 2.0% of operational taxonomic units (OTUs) are respectively specific to Ca. femoratus and Cr. levior, probably representing their respective core cuticular bacterial community, and (c) 3.6% of OTUs are shared between the two ant species. Additionally, mass spectrometry metabolomics analysis of metabolites on the cuticle of ants, which excludes the detection of cuticular hydrocarbons produced by the host, were conducted to evaluate correlations among bacterial OTUs and m/z ion mass. Although some positive and negative correlations are found, the cuticular chemical composition was weakly species‐specific, suggesting that cuticular bacterial communities are prominently environmentally acquired. Overall, our results suggest the environment is the dominant source of bacteria found on the cuticle of ants.     

Structure of the ovary and mode of oogenesis in a freshwater crab Potamon dehaani

Structure of the adult ovary and oogenetic mode were examined in the freshwater crab Potamon dehaani. An H‐shaped ovary consisting of a pair of long ovarian sacs connected by a narrow bridge tube is located in the cephalothorax on the dorsal side of the stomach. A short oviduct with a seminal receptacle is connected with the posterior end of each ovarian sac, and a genital pore opens on the sternum of the sixth thoracic segment. The ovarian wall consists of a layer of ovarian epithelium that infolds to form a number of oogenetic pouches of various sizes. These are present mainly in the anterior regions of the ovarian sacs, are scarce in the posterior regions of the ovarian sacs, and are absent from the bridge tube. Each oogenetic pouch contains an egg or a relative large oocyte in its lumen. Germaria containing oogonia, very early previtellogenic oocytes, and somatic interstitial cells are located in the ovarian epithelium near the necks of the oogenetic pouches in the anterior regions of the ovarian sacs and are randomly scattered throughout the ovarian epithelium in the posterior regions of the ovarian sacs. In cross section, the germaria appear to be concentrated into a central germarial cluster in the ovarian sac. In the posterior regions of the ovarian sacs, however, the germaria are randomly scattered throughout the ovarian epithelium. An early previtellogenic oocyte leaves its germarium and raises the ovarian epithelium infolds to form a new oogenetic pouch in which it grows to maturity. Mature eggs are ovulated from the oogenetic pouches into the ovarian lumen, transferred from the ovarian lumen into the oviducts, fertilized there by sperm stored in the seminal receptacles, and then oviposited through the genital pores. The female reproductive system is surrounded wholly and tightly by a thin, cellular, membranous sheath, which has often been mistaken as the ovarian epithelium in some decapod crustaceans. J. Morphol. 239:107–114, 1999. © 1999 Wiley‐Liss, Inc.     

Investigation of chicken intestinal bacterial communities by 16S rRNA targeted fluorescence in situ hybridization

The aim of the investigation was to quantify selected dominant bacterial groups in the chicken intestinal tract. Conventional production was used as model and the effect of the supplement with Salinomycin was evaluated. Hybridization conditions were optimized for published probes with respect to a panel of reference bacteria. In chicken intestinal samples bacteria were quantified by fluorescence in situ hybridization with 16S rRNA oligonucleotides directed towards bacteria related to Lactobacillus, Bacillus, Enterococcus-Streptococcus-Lactococcus, Enterobacteriaceae, Bacteroides, Clostridium and the domain Bacteria in lumen of ileum and cecum as well as on the intestinal wall including mucus of four individuals. Salinomycin in feed reduced counts of the Lactobacillus-, Enterobacteriaceae- and Clostridium-like bacteria in lumen of ileum compared to the conventional control. Increased or decreased bacterial counts were registered by Salinomycin in the ceca compared to the control. Relatively higher counts of Bacteroides- and Clostridium-like bacteria were found on the intestinal wall including mucus compared to lumen. The increase in numbers of some bacterial groups as well as the expected reduction by Salinomycin and the observed difference in the relative distribution of bacteria between lumen and intestinal wall are new observations that will need further investigation.     

Ultrastructure of sperm storage and male genital ducts in a male holocephalan, the elephant fish, Callorhynchus milii.

In chondrichthyes, the process of spermatogenesis produces a spermatocyst composed of Sertoli cells and their cohort of associated spermatozoa linearly arrayed and embedded in the apical end of the Sertoli cell. The extratesticular ducts consist of paired epididymis, ductus deferens, isthmus, and seminal vesicles. In transit through the ducts, spermatozoa undergo modification by secretions of the extratesticular ducts and associated glands, i.e., Leydig gland. In mature animals, the anterior portion of the mesonephros is specialized as the Leydig gland that connects to both the epididymis and ductus deferens and elaborates seminal fluid and matrix that contribute to the spermatophore or spermatozeugmata, depending on the species. Leydig gland epithelium is simple columnar with secretory and ciliated cells. Secretory cells have periodic acid-Schiff positive (PAS+) apical secretory granules. In the holocephalan elephant fish, Callorhynchus milii, sperm and Sertoli cell fragments enter the first major extratesticular duct, the epididymis. In the epididymis, spermatozoa are initially present as individual sperm but soon begin to laterally associate so that they are aligned head-to-head. The epididymis is a highly convoluted tubule with a small bore lumen and an epithelium consisting of scant ciliated and relatively more secretory cells. Secretory activity of both the Leydig gland and epididymis contribute to the nascent spermatophores, which begin as gel-like aggregations of secretory product in which sperm are embedded. Fully formed spermatophores occur in the ductus. The simple columnar epithelium has both ciliated and secretory cells. The spermatophore is regionalized into a PAS+ and Alcian-blue-positive (AB+) cortex and a distinctively PAS+, and less AB+ medulla. Laterally aligned sperm occupy the medulla and are surrounded by a clear zone separate from the spermatophore matrix. Grossly, the seminal vesicles are characterized by spiral partitions of the epithelium that project into the lumen, much like a spiral staircase. Each partition is staggered with respect to adjacent partitions while the aperture is eccentric. The generally nonsecretory epithelium of the seminal vesicle is simple columnar with both microvillar and ciliated cells.     

Mechanism of chlorpromazine binding by Gram-positive and Gram-negative bacteria

Chlorpromazine forms charge-transfer complexes with xanthene dyes in bacteria. These complexes permit the differentiation of Gram-positive and Gram-negative bacteria in both light and polarization microscopy. The birefringence induced by the charge-transfer complex might explain the molecular basis of bacterial staining.The charge-transfer complexes formed between chorpromazine and xanthene dyes accumulate in the bacterial cell, mainly inside the bacterial cell wall. The complexes give the cells a color, which depends on the chemical composition of the staining structure, and in particular the polysaccharides of the cell wall in bacteria.Metachromatic granules were seen inside Gram-positive bacteria after chlorpromazine and rose bengal staining. Although the nature of these granules remains unclear, this type of binding may have a role in the inhibition of biochemical processes in the bacterial cells.     

Pathogenesis of Helicosporidium sp. (Chlorophyta: Trebouxiophyceae) in susceptible noctuid larvae

Helicosporidium sp. is a unique, achlorophyllous green alga that has been reported to infect various insect orders, including Lepidoptera, Diptera, and Coleoptera. The infectious cyst stage is ingested by the host, ruptures in the midgut lumen, and releases a filamentous cell. Histopathological examinations using larvae of a susceptible noctuid host, Spodoptera exigua, showed both cysts and filamentous cells affiliated with the microvillar lining of the midgut epithelium. A considerable proportion of the ingested cysts (22-39%) were recovered in feces collected 24 h after ingestion. A small number of filamentous cells passed the midgut epithelium and entered the hemocoel within 4-24 h after cyst ingestion. After 48 h, vegetative cell stages were detected in the hemolymph, followed by a 4- to 5-day period of increasing multiplication. Cyst differentiation in the colonized hemolymph began 6-7 days after the treatment.     

Phylogenetic diversity of cellular organization in the cardia of muscoid flies (Diptera: Schizophora)

In each of 30 dipteran species, representing 13 acalyptrate and 7 calyptrate families, the cardia is formed from specialized cells at the junction between foregut and midgut. Foregut epithelium forms the stomodeal valve; midgut epithelium envelops the valve to form the cardia's outer wall. Cytological characteristics within these epithelia differ from region to region and from species to species. Since the cardia secretes the peritrophic membrane, cardias with diverse patterns of cellular differentiation may be expected to produce peritrophic membranes with similarly diverse properties. Close relatives often share more details of cardia structure than do distantly related taxa. Within the monophyletic Calyptratae, a common pattern of cellular differentiation includes three distinct zones of columnar midgut cells enclosing a flanged stomodeal valve. Among species in the paraphyletic Acalyptratae, midgut typically includes a single zone of tall columnar cells, while the valve may be spheroidal, cylindrical, conical, or flanged. The correlation of phylogenetic distance with divergence in cardia organization implies a strong influence of ancestry upon current structure, regardless of current diet. However, at least some of the observed diversity in cardia structure is associated with dietary divergence. Calyptrate flies with derived blood-feeding behavior display cellular differentiation that is simplified from that seen in calyptrate relatives with less specialized feeding habits. This evolutionary modification suggests that cardia organization and hence peritrophic membrane structure can adapt to dietary changes, with possible significance for the spatial organization of digestive processes and interactions with ingested microorganisms.