Musculature,nervous system and glands of metasomal abdominal segments of the male of Nomia melanderi Ckll. (Hymenoptera,Apoidea)

Each muscle of the third metasomal segment of the male of Nomia melanderi Ckll. is described in detail. The points of attachment of each muscle are compared with their homologs in other pregenital segments and with their homologs in the female. The function desgnated for each muscle describes its action alone or in conjunction with other muscle(s). New names are given to genital muscles by referring in the name to their points of attachment. Each intratergal muscle has homologous points of attachment in the pregenital segments of both sexes. The median tergo-dorsoplical muscle of the seventh segment and the oblique tergo-dorsoplical muscle of the eighth segment have changed their points of attachment. The intrasternal muscles are modified to suit the needs of courtship and mating, thus they are different from their homologs in the female. The spiracular muscles are well developed in all segments except the eighth, where the sterno-spiracular muscle is absent. The extrinsic genital muscles are derived from the intrasternal muscles of the eighth and ninth segments. The parameral and volsellar muscles are reduced in number. The aedeagal muscles, except the aedeago-phallic, have retained similar points of attachment to those found in primitive Hymenoptera. The topography of the metasomal nervous system is reported in detail by following each nerve and nervule to its termination. The study shows that (at least in Nomia) the criterion of nerve-concentration should not be used alone to indicate evolutionary levels. To accommodate the morphological changes in the terminal segments the Anterior and Posterior Lateral Nerves have migrated to new locations. The pattern of nerve topography (even at the nervule level) is homologous both in the different pregenital segments and between the sexes. The fact that homology does not exist between the external genitalia of the male and the modified ovipositor of the female supports the thesis that the male genital capsule is of phallic rather than prephallic origin. A pair of intersegmental membrane glands located between the seventh and eighth sterna is described. These glands may be the source of a pheromon responsible for gregariousness among “sleeping” males.     

The locust abdominal muscle receptor organ: response characteristics and its role in the control of segmental distance

A single mutipolar receptor cell is located at the dorsal edge of the lateral internal dorsal muscle in each abdominal segment of the locust (Locusta migratoria). Muscle and receptor cell form the abdominal muscle receptor organ. The receptor cell monitors length changes in the intersegmental muscle, and as a consequence also detects the length of an abdominal segment (cuticule and intersegmental membrane).The muscle receptor organ responds in a phasictonic fashion. The phasic component encodes the rate of change in the stimulus independent from the prevailing length of the muscle receptor organ. The tonic component monitors the absolute length of the muscle.Stimulation of a single muscle receptor organ leads to reflex effects on the ipsilateral longitudinal muscles in at least three adjacent segments. Muscles that shorten the abdomen are activated while their extending antagonists receive reduced activity.The reflex activation of the muscles is polysynaptic. Monosynaptic connections between the receptor and the motoneurones were not found.We identified an interneurone that receives monosynaptic input from the muscle receptor organs in at least three adjacent segments. The interneurone excites motorneurones to the longitudinal muscles of the next posterior segment.Abbreviations aMROII abdominal muscle receptor interneurone 1 - AS3 third abdominal segment - AS4 fourth abdominal segment - AS5 fifth abdominal segment - AS6 sixth abdominal segment - EPSP excitatory postsynaptic potential - MN median nerve - MR multipolar receptor cell - MRO muscle receptor organ - N1 tergal nerve - N2 sternal nerve     

Electrophoretical studies of proteins of the hypopharyngeal glands and of the larval food ofMelipona quadrifasciata anthidioides Lep. (Hymenoptera,Meliponinae)

Summary The electrophoretical protein patterns of hypopharyngeal glands, larval food ofMelipona, and royal jelly ofApis were compared.Since protein patterns of hypopharyngeal glands from newly emerged workers, brood cell provisioners and foragers are similar to freshly deposited larval food, the identical protein bands probably represent actual gland secretion. This suggests that, as inApis, the glands secrete proteins to the larval food, and maintain this ability throughout life, although at slightly different intensities, according to the activity of the bees.The similarity on the electrophoretic profiles of the major larval food protein inApis andMelipona is an interesting finding because of its probable evolutionary significance.     

Male specific natural products in the bug, Leptoglossus phyllopus: Chemistry and possible function

In the Coreoidea and some allied groups, male adults possess an abdominal gland opening through a midventral ostiole in the 7–8th abdominal intersegmental membrane. Using a GC-MS system, the following aromatic volatiles were identified in the abdominal gland secretion from males of the leaf footed bug, Leptoglossus phyllopus: guaiacol, benzyl alcohol, syringaldehyde, methyl p-hydroxybenzoate, acetosyringone, and vanillin. Males from which the glands were removed still mated and were competitive with control males for a limited number of females. The ventral abdominal gland secretion may act as a long-range attractant of females. The possibility that attraction of females by males in Heteroptera is an adaptation facilitating colonization of successional habitats is discussed.     

Abdominal stretch reception in Dipetalogaster maximus (Hemiptera: Reduviidae)

Each half abdominal segment in 5th-instar larvae of the giant bloodsucking reduviid, Dipetalogaster maximus, contains 3 stretch receptor neurones, one associated with the tergosternal muscles, one with the ventral intersegmental muscles and one with the dorsal intersegmental muscles. Each of the three receptors respond phasically to the onset of stretch in its respective muscle group, but none show persistent activity upon prolonged stretch. By contrast, stretch of the main abdominal nerves (which run between the thoracic ganglion and the ventral intersegmental muscles of each abdominal segment) is accompanied by a prolonged and sustained pattern of discharge by an as yet unidentified neurone, the rate of discharge being proportional to the degree of stretch. In life, the abdominal nerves become stretched to about 145% of their resting length when the larva takes a bloodmeal. Thus it appears that in Dipetalogaster stretch of the abdominal nerves themselves is the only mechanism for stretch reception after a blood meal.     

Comparative analysis of the male reproductive accessory glands of bats Noctilio albiventris (Noctilionidae) and Rhynchonycteris naso (Emballonuridae)

In eutherian mammals, the male reproductive accessory glands (RAGs) comprise the prostate, bulbourethral glands, ampullary glands, and the seminal vesicles. Their composition, anatomy and function vary widely between species. This study aimed to characterize histologically and compare the RAGs of bats. The RAGs of Noctilio albiventris (Noctilionidae) and Rhynchonycteris naso (Emballonuridae) were studied using anatomical and histological methods, and were reconstructed three dimensionally. The RAGs of N. albiventris and R. naso are composed of a compact glandular complex that surrounds the urethra and a pair of bulbourethral glands, which are extra‐abdominally located in the inguinal region. In both species, the glandular complex is composed of two well‐defined prostatic regions (ventral and dorsal). The ventral region showed an atypical epithelium (holocrine), where no obvious cellular limits were observed, and PAS‐positive secretion. The dorsal region had a pseudostratified cuboidal epithelium, with basal and secretory cells, and PAS‐negative secretion. Noctilio albiventris also had urethral glands (Littre glands) surrounding the urethra, however, R. naso had only muscles. Both species had bulbourethral glands, with simple columnar epithelium and PAS‐positive secretion. In conclusion, the RAGs of N. albiventris and R. naso comprised a pair of bulbourethral glands and an intra‐abdominal complex, composed of a prostate with two different regions (ventral and dorsal), while the ampullary glands and seminal vesicles were missing in both species. This morphology was more closely related between N. albiventris and R. naso, and to species of the family Phyllostomidae than to families Molossidae and Vespertilionidae. J. Morphol. 277:1459–1468, 2016. © 2016 Wiley Periodicals, Inc.     

Location,morphology and histology of sex pheromone glands of the female gypsy moth, Lymantria dispar (L.)

Scanning electron microscopy, histology and a male wing fanning bioassay were used in this study to locate the sex pheromone-producing glands of the female gypsy moth, Lymantria dispar. When exposed to female sex pheromone, adult males exhibit a strong wing fanning behaviour prior to take off. We found that adult males showed positive response to calling females and to tissue extract from both dorsal and ventral portions of the intersegmental membrane between 8th and 9th-abdominal segments. A typical male response usually starts with elevation of antennae, movement of head in different directions, walking, wing fanning and onset of search flight. Histological and scanning electron microscopic studies suggested that the sex pheromone glands are located on the dorsal and ventral aspects of the intersegmental membrane. The glands appear as two highly convoluted integumentary areas with hypertrophied glandular epidermal cells.     

The pheromonal gland of Lymantria dispar: Morphology and evidence for its innervation

The morphological features of the glandular epithelium that secretes pheromone in the polyphagous pest gypsy moth Lymantria dispar are described by light and electron microscopy. The monolayered gland cells are covered by the folded cuticle of the intersegmental membrane between the 8th and 9th abdominal segments showing neither sites of discontinuity nor distinct openings on its external surface. The cells bear a large, often irregularly shaped nucleus, and contain granules of variable amount and electron‐density. These granules are mostly located in the basal compartment of the cytoplasm, in a labyrinthine zone laying on a basement membrane. The apical membrane of the gland cells bear microvilli and cell–cell contact is established by different junctional structures. Nerve fibers enwrapped in glia are found beneath the basement membrane, in close contact with the secretory cells. This latter finding represents the first evidence of the innervation of the pheromonal gland in L. dispar. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

Neuromuscular coordination and proprioceptive control of rhythmical abdominal ventilation in intactLocusta migratoria migratorioides

Summary Within the fifth abdominal segment of intact locusts a group of dorso-ventral expiratory muscles and one inspiratory antagonist display alternating ventilatory patterns of three basic types. Accelerated movements in the dorso-ventral plane are supported by isometric activity of the intersegmental muscles which prevent extensions in the longitudinal axis.The intersegmental coupling of ventilatory motor patterns is strict during strong ventilation and loose and more metachronal with weaker pumping movements.In resting animals ventilatory rhythms are discontinuous and the long intervening pauses are interrupted by miniature inspirations only. Pumping series have a tendency to prolong the later ventilatory cycles, and interfering rhythms of different pumping types occur. Low concentrations of atmospheric CO₂ up to 3 % do not accelerate ventilatory rhythms.Afferent activity from proprioceptors could be related to ventilatory motor bursts and stimulation of the sensory nerve produces inspiratory bursts via the segmental ganglion.The neuronal mechanisms of synergistic and antagonistic muscle control as well as the segmental and intersegmental coordination and the effect of autonomous ganglionic oscillators in ventilation are discussed.     

New exocrine glands in the legionary ants of the genus Leptanilla (Hymenoptera,Formicidae, Leptanillinae)

Summary This paper reports the results of the first detailed study of the morphology of exocrine glands in two species of the ant subfamily Leptanillinae. Workers of Leptanilla escheri and L. japonica possess a large, unpaired sternal gland in the VIIth abdominal sternite, and an hypertrophied poison gland which is surrounded by a massive muscle layer. The sternal gland is absent in the queen of L. japonica, and the poison gland is highly degenerated. The queen is, however, endowed with a series of large, paired, intersegmental tergal and sternal glands, which occur between the IVth through the VIIth segments. The queen also posseses large spiracular plate glands.     

Fine structure of the intersegmental membrane glands of the sixth abdominal sternum of female Nomia melanderi (Hymenoptera,Apoidea)

The fine structure of the intersegmental glands of the sixth abdominal sternum in 1-week old females of Nomia melanderi is presented. The plasma membrane of the secretory cell is unfolded in many places and is covered by a basement membrane. The microvillous surface is invaginated to form a rather long sinuous cavity. The endoplasm is almost entirely filled by secretory granules. Many secretory granules are located close to the inner surface of the invaginated plasma membrane. The invagination contains a porous ductule, apparently of cuticulin origin, that is connected directly with the inner layer of the transport duct of the duct-forming cell. This type of arrangement allows the direct flow of the secretory substance to the outside in a continuous way. The cylindrical duct-forming cell, besides having typical cell organelles, contains a cuticular transport duct. This duct is composed of a thin cuticulin layer surrounded by a rather thick epicuticular one. The results suggest that the secretory cell has two secretory cycles. The first occurs while the gland is differentiating (at the pupal stage) and is involved in secretion of the cuticulin that forms the porous ductule. The second cycle, which starts by the beginning of nesting, is involved in the secretion of a substance that is carried to the outside via the transport duct of the duct-forming cell.     

The evolution of food-producing glands in eusocial bees (Apoidea,Hymenoptera)1

A food-producing role for cephalic exocrine glands has arisen independently in both taxa of highly eusocial bees, Apis and Meliponini. With several exceptions, there is little evidence that food is produced by glands of solitary bees or by most bees at lower levels of sociality. We suggest that this association with sociality is due to four adaptive features of these glands: (1) food from the glands allows feces from queens and larvae to have a small volume, (2) the queen's fecundity can be increased, (3) nutrient recovery via cannibalism can be facilitated, and (4) rearing of emergency replacement queens is accelerated. Acceleration of the rearing of other castes and of queens in the normal process of colony fission is not clearly an advantage ascribed to these glands. Trophic eggs produced by meliponine colony workers are analogous to the secretions from food-producing glands in Meliponini and Apis workers.     

The integument of the Queensland fruit fly,Ddacus tryoni (Frogg.)

Summary During the pupal stage of Dacus tryoni, the hypodermis of the larva is replaced by an imaginal generation of smaller cells. The hypodermal cells of the tergal glands on the fifth abdominal segment of the adult were examined with the electron microscope; they contain slender, membrane-limited bundles of hollow wax filaments that traverse the cuticle in branched pore canals. Outside the glandular areas, the pore canals are narrower. The cuticle of the adult undergoes its greatest increase in thickness soon after emergence; it becomes sclerotized gradually. No epicuticle was detected with either the light or electron microscopes.Early in adult development, bristles are formed over the general surface of the terga. Most of these are innervated by single, bipolar nerve cells, and have more or less enlarged trichogen cells that appear to secrete wax through pore-plates in the cuticle. The bristles in different regions of the abdomen range in function from pure sensory receptors to pure secretors. The sensory bristles on the tergal glands were examined with the electron microscope.For assistance with the electron microscopy, I thank Mr. Tony Webber and Miss Ann Miller of the Electron Microscopy Unit at Sydney University. — Supported by a C.S.I.R.O. Junior Post-Graduate Studentship.     

Histology and ultrastructure of the salivary glands and salivary pumps in the scorpionfly Panorpa obtusa (Mecoptera: Panorpidae)

Liu, S. and Hua, B. 2009. Histology and ultrastructure of the salivary glands and salivary pumps in the scorpionfly Panorpa obtusa (Mecoptera: Panorpidae). —Acta Zoologica (Stockholm) 91 : 457–465. The morphology, histology and ultrastructure of the salivary glands and salivary pumps in the scorpionfly Panorpa obtusa Cheng 1949 were investigated using light microscopy and scanning and transmission electron microscopy. The salivary glands display a distinct sexual dimorphism. The female has only two small sac‐like glands located in the prothorax, while the male possesses six long tubular glands extending into the sixth abdominal segment. The male salivary glands can be divided into five distinct regions. The apical long, thin secretory region possesses numerous secretory cells containing large secretory vesicles; the salivary reservoir expands in diameter, accumulating and temporarily storing the saliva in addition to secreting saliva; the constricted region contains prismatic cells with complex infolded plasma membrane; the sac has an internal brush border to absorb water and ions; the common salivary duct contains longitudinal muscles in the male, but not in the female. The salivary pump possesses independent strong dorsal muscles and abundant internal palm spines near its orifice. The anatomy and ultrastructure of the salivary glands and the salivary pump of scorpionflies as well as their possible functions are briefly discussed.     

Histology,histochemistry, and emptying mechanism of the venom glands of some elapid snakes

The venom glands of several species of elapid snakes are described. The main venom gland consists of many tubules which usually contain large amounts of secretion product. The accessory gland surrounds the entire venom duct and is usually composed of uniform mucous epithelium. The epithelium lining the tubules of the accessory gland of Naja naja is composed of two distinct types of cells. Histochemical tests indicate that the main venom gland reacts with mercury bromphenol blue and PAS but not with alcian blue. The accessory gland reacts with PAS and alcian blue, and not with mercury bromphenol blue. Treatment of sections with sialidase demonstrates the presence of a sialomucin in the accessory gland. Stimulation of the muscles associated with the venom gland offers an indication of the venom expulsion mechanism of Bungarus caeruleus. A comparison of the venom apparatus of elapid and viperid snakes emphasizes marked differences in the internal anatomy of the venom glands, muscles associated with the gland, and arrangement of glandular components. The morphological differences and dissimilar venom expulsion mechanisms support the recent view of the polyphyletic origin of venomous snakes.     

Neue Abdominaldrüsen bei Ameisen

Zusammenfassung Sechs Ponerinen aus dem Tribus Ponerini wurden auf Abdominaldrüsen untersucht:Leptogenys ocellifera (Roger),Leptogenys chinensis (Mayr),Diacamma sp.,Odontomachus haematodes (L.),Harpegnathus saltator (Jerdon) undPonera coarctata (Latreille). Eine große Vielfalt von verschiedenen Drüsenorganen konnte gefunden werden (Tabelle 1). Bei jeder untersuchten Art fanden wir dorsal zwischen dem 6. und 7. Abdominaltergit eine intersegmentale Komplexdrüse. Die Größe der Drüsen, die Anordnung ihrer Drüsenzellen und die Form der Reservoire sind z.T. sehr unterschiedlich ausgebildet. BeiLeptogenys wie auch beiH. saltator befindet sich intersegmental zwischen dem 6. und 7. Abdominalsternit eine Komplexdrüse.Leptogenys verfügt zusätzlich über eine Komplexdrüse zwischen dem 5. und 6. Abdominalsternit. An der Mündung der gut ausgebildeten Reservoire dieser Drüsen finden sich bei einigen Arten charakteristisch geformte Kutikulastrukturen, die als Oberflächenvergrößerung oder Sekretspeicher interpretiert werden.Tergo-sternal gelegene Bündel von Drüsenzellen finden sich beiDiacamma sp.,P. coarctata, H. saltator und beiO. haematodes. Einen weiteren Drüsentypus bilden Ansammlungen von Epitheldrüsenzellen. BeiL. ocellifera liegen diese Zellen dem 6. Sternit; beiL. chinensis dem 7. Sternit auf. Auch Stacheldrüsen sind in einer ähnlichen Vielfalt vorhanden. BeiDiacamma sp. befinden sich in den Stachelscheiden zwei verschiedene Typen von sezernierenden Zellen, Drüsenzellen mit einem Ausführkanal und Epitheldrüsenzellen. Bei den Vertreterinnen der GattungLeptogenys, O. haematodes undP. coarctata liegt nur der erstere Drüsentyp in ausgeprägter Form vor. Bei diesen Arten wie auch beiH. saltator sind die Epithelzellen der Stachelscheiden im Vergleich zu jenen Epitheldrüsenzellen beiDiacamma sp. geringfügig erhöht. Drüsenzellen mit einem Ausführkanal konnten beiH. saltator in den Stachelscheiden nicht gefunden werden.H. saltator undO. haematodes zeichnen sich an der Membran zwischen den oblongen Platten (Stachelscheidenbasis) durch einen großen paarigen Komplex von sezernierenden Zellen aus. An dieser Stelle finden sich auch beiDiacamma sp.,P. coarctata undL. chinensis einzelne Drüsenzellen. Zusätzlich ist beiDiacamma sp. hier eine Ansammlung von Epitheldrüsenzellen vorhanden, die angedeutet ebenfalls bei den anderen Arten vorliegt. Bei allen untersuchten Arten münden Kanäle von unterschiedlich großen, dorsolateral gelegenen Komplexen von Drüsenzellen in die membranose Verbindung des Stachelapparates mit dem letzten freien Segment. BeiDiacamma sp.,P. coarctata, O. haematodes undH. saltator befinden sich zudem lateral der Spirakularplatten kleine Ansammlungen von Drüsenzellen.Diacamma sp.,P. coarctata undH. saltator verfügen über latero-ventral befindliche Drüsenzellen, deren Kanäle in die Membran zwischen dem Stachelapparat und dem 7. Sternit münden.

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New abdominal glands in antsI. Ponerini (Formicidae: Ponerinae)

Summary The abdominal glands of six ponerine ants belonging to the tribe Ponerini were analysed:Leptogenys ocellifera (Roger),Leptogenys chinensis (Mayr),Diacamma sp.,Ponera coarctata (Latreille),Odontomachus haematodes (L.) andHarpegnathus saltator (Jerdon). A great variety of glands was found. An intersegmental complex gland is located between the sixth and seventh abdominal tergite in each species investigated. But size, arrangement of gland cells and shape of reservoir differ. In addition, representatives of the genusLeptogenys andH. saltator both have sternal intersegmental complex glands. InL. ocellifera andL. chinensis these glands are located between the fifth and sixth and also between the sixth and seventh abdominal sternite. InH. saltator one sternal gland is situated between the sixth and seventh abdominal sternite. In some species we found characteristical sculptures on the cuticle at the orifice of the well developed reservoirs of the glands. These sculptures could be interpreted as an enlarging of the surface of the cuticle or as a reservoir.Another type of gland cells are epithelial glandular cells. They form distinct layers on the seventh sternite inL. chinensis and on the sixth sternite inL. ocellifera.Tergo-sternal bunches of secretory cells were observed inDiacamma sp.,P. coarctata, H. saltator and inO. haematodes. A similar variety of glands was found associated with the sting apparatus in the gonostyli, at the membrane between the two oblong plates and at the membranous connections between the sting apparatus and the last abdominal segment. InDiacamma sp. two distinct glandular cells are located in the gonostylar sclerites, i.e. secretory cells, each drained by a cuticular ductule and epithelial glandular cells. In the two representatives of the genusLeptogenys, inO. haematodes and inP. coarctata only the first type of gland cell was found. In these species as well as inH. saltator the epidermal cells of the gonostylar sclerites form different states of transition from degenerated epithelial cells to glandular epithelial cells. InH. saltator there are no secretory cells with a cuticular ductule in the gonostyli.Likewise, large paired complexes of gland cells were found at the base of two genostylar sclerites inH. saltator and inO. haernatodes. Though less developed, gland cells of the latter type are inDiacamma sp., P. coarctata and inL. chinensis also located at the base of the gonostyli. InDiacamma sp. the epidermal cells of the membrane connecting the two gonostyli do have secretory function. In the other species investigated they are less developed, and a secretory function cannot be considered certain. In each species investigated ductules of gland cells also open dorsolaterally into the sting chamber. Furthermore inDiacamma sp.P. coarctata, O. haematodes and inH. saltator, ductules of gland cells also open laterally and, except inO. haematodes, latero-ventrally into the membranous connection between the sting apparatus and the last abdominal segment.

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Mit Unterstützung der Deutschen Forschungsgemeinschaft. Herrn Dipl.-Biol. R. Klinger danken wir für stete Diskussionsbereitschaft sowie Herrn Dr. W. Gnatzy für wertvolle Hinweise.     

Compartments in the abdomen of Drosophila and the role of the engrailed locus

A clonal analysis has shown that the dorsal surface of the first abdominal segment of Drosophila melanogaster is subdivided into anterior and posterior compartments. Cells of the posterior compartment grow up to but not beyond the anterior-posterior compartment border within the first abdominal segment and the intersegmental border that defines the boundary between the first and second abdominal segments. Growing within these boundaries, a narrow band of tissue clonally isolated from the adjoining tissue is formed. When these posterior cells are deficient for the engrailed locus, however, neither the compartment nor the segment border is maintained. The implications, that compartmentalization is essential for segmentation, and that all insect segments are subdivided by anterior and posterior compartments, are discussed.     

Morphological analysis of the male reproductive accessory glands of the bat Artibeus lituratus (Phyllostomidae: Chiroptera)

Bats are distributed worldwide from tropical to temperate regions. Despite their wide geographical radiation and advances in studies using evolutionary approaches, aspects related to the reproduction of these animals remain poorly explored, especially those related to the male reproductive accessory glands (RAGs). Thus, the aim of this study was to analyze the morphophysiology of the male RAGs in the bat Artibeus lituratus. The RAGs in A. lituratus are composed of a compact intra‐abdominal glandular complex, consisting of the prostate with two prostatic regions (ventral and dorsal), plus Littre glands and a pair of extra‐abdominal bulbourethral glands. The ventral region of the prostate has an epithelium with variable morphology, due to its holocrine type of secretion. In contrast, the dorsal region has a typical cubic‐to‐columnar pseudostratified epithelium. Both regions contain two cell types, basal and secretory cells. Similar to the epithelial morphology, the secretion also varies, with the ventral region containing numerous PAS‐positive globular vesicles, whereas the dorsal region has a more fluid, hyaline and PAS‐negative secretion. Littre glands are dispersed in the connective tissue of the urethra, while the bulbourethral glands are located in the penile root, both glands with cubic‐to‐columnar pseudostratified epithelium and globular PAS‐positive secretion. The results demonstrate that the RAGs of A. lituratus are composed of two prostatic regions, ventral and dorsal, and urethral and bulbourethral glands, with no seminal vesicles. Each prostatic region has unique and distinctive characteristics, with the ventral region presenting an exclusive holocrine nature and the dorsal region having similarities to the ventral prostate of rodents.