Morphological diversity of the metathoracic spiracle in the Lygaeoidea (Hemiptera: Heteroptera).

Spiracles are the openings in the exoskeleton of insects through which air enters into the respiratory system that is formed by a series of tubes called tracheae. They are primarily located on the abdomen, but can also occur on the thorax, including the metathorax. An insect metathoracic spiracle is usually composed of an external opening and a more internal filter apparatus. We propose new terminology for these structures, and we explore the value in their use in taxonomic and phylogenetic studies within the true bug infraorder Pentatomomorpha, with emphasis on the superfamily Lygaeoidea (Insecta: Hemiptera: Heteroptera). These structures were studied using scanning electron microscopy. Two types of metathoracic spiracle external openings were recognized: a narrow opening (type N), which is slit-like; and a wide opening (type W), with internal fine structures located between the mesothoracic and metathoracic margins of the interpleural suture clearly visible. The filter apparatus in the Pentatomomorpha consists of modified mushroom bodies of the metathoracic scent gland evaporatorium, for which the term mycoid filter processes is proposed. Eight different types of mycoid filter processes, and an unmodified microsculpture type (a type with usual cuticular microsculpture) and filter setae can be found on the anterior or posterior margins of the metathoracic spiracle. We believe the wide opening (type W) to be the plesiomorphic character state in the Pentatomomorpha, with multiple, independent transformations leading to the narrow opening in Lygaeoidea. Considerable variability in the structure of the spiracle opening (in Lygaeoidea), and in the structure of the mycoid filter processes (in Pentatomomorpha) was detected. Overall, we found the morphology of these structures to be of limited value concerning the taxonomy or for determining phylogenetic relationships of the higher taxa (families) of Pentatomomorpha, but they may be useful as additional evidence for taxonomic and phylogenetic studies at the generic and perhaps the tribal levels.

External morphology of the abdominal glands in Asopinae (Hemiptera: Heteroptera: Pentatomidae).

Heteropterans communicate chemically through thoracic or abdominal glandular complexes. The dorso-abdominal scent glands (DAGs) are externalized by cuticular specialized structures forming the external scent efferent system (SES). Different groups in Heteroptera present other glands, such as the Asopinae (Pentatomidae), whose males can have ventral abdominal pheromonal glands externalized by cuticular modifications, called glandular patches (GPs). Here we describe the SES of DAGs and, for the first time, the GPs of 22 Asopinae genera. The ostioles of SES 1 vary in shape. SES 2 and SES 3 were restricted to scars, indicating that only the SES 1 remains functional in adults. The GPs are found mostly in segments V and VI. We have not found any difference in the morphological pattern between the SES of species bearing GPs and those lacking it, meaning that the acquisition of GPs is likely related to some sexual behavior, thus not as a complex structure replacing any role of the DAGs. The cuticle of the GPs is microsculptured, bearing many pores surrounding the base of sensilla being the pathway for the secretions to reach the outside of the exoskeleton. The sensilla may be adaptations to increase the contact surface, assisting rapid evaporation of the volatile secretions from the pores.

Structures related to pheromone storage in alar androconia and the female abdominal scent gland of Heliconius erato phyllis, Heliconius ethilla narcaea, and Heliconius besckei (Lepidoptera: Nymphalidae: Heliconiinae).

We describe the morphology of alar androconia and the female abdominal scent gland of Heliconius erato phyllis, Heliconius ethilla narcaea, and Heliconius besckei. Androconial scales of Heliconius, which are arranged in overlapping wing bands, release pheromones during courtship, probably through vibratory movements of male wings over the female to induce her to mate. An antiaphrodisiac is produced by glands located in the valves of the male and is transferred during copulation to the yellow dorsal abdominal sac present in the virgin female, causing this sac to emit a scent that reduces the attractiveness of the female for courtship with other males. Stereomicroscopy, SEM, and TEM analyses were conducted to describe the morphology of the internal and external scales and the external abdominal scent sac. The findings revealed different sizes of external androconial scales and an internal group of porous structural vesicles that are probably related to the preservation of internal space, reception and storage of secretions, and elimination of volatiles when the male is actively involved in courtship. Translucent projections on the female abdominal scent sac create open reservoirs for the reception, storage, and emission of antiaphrodisiac volatiles along with stink clubs. Male valve denticles vary in form and probably attach securely to the female sac during mating, thus ensuring secretion transfer. These features are discussed in the context of a comparative analysis.

Anatomical, Histological, and Histochemical Analyses of the Scent Glands of the Scorpion Mud Turtle (Kinosternon scorpioides scorpioides).

Fossil evidence suggests that scent glands are basal features of Testudines. However, we know little about the structure of these glands in the Brazilian Kinosternidae. In this study, we described the macroscopic anatomy, histology, and histochemistry of the scent glands of three males and three females of Kinosternon scorpioides scorpioides from the Marajó mesoregion, Pará State, Brazil. In all of the specimens analyzed, regardless of sex, we found four scent glands, including two axillary and two inguinal glands that were structurally similar to each other. Each gland consisted of a single holocrine secretory lobule, a large lumen surrounded by relatively thin glandular secretory epithelium, an adjacent narrow layer of loose connective tissue, and a thick layer of skeletal striated muscle tissue surrounded by a serous tunic. The secretory epithelium produced a characteristic malodorous yellowish substance that was passed via a single duct through a bone channel in the bridge connecting the carapace to the plastron and excreted through an outer pore in the plate of each respective gland. Histologically, the secretory epithelium presented cells with two types of secretory vacuoles. Type 1 vacuoles stained red were the largest and most frequently found, and stained positively with Periodic acid-Schiff (PAS), suggesting they contained glycoproteic complexes. Type 2 vacuoles were translucent, smaller in size and fewer in number, and negative for PAS staining. Because they are very primitive structures, scent glands must play important roles in the lives of chelonians, but their real function remains unknown. Several hypotheses suggest that they can act as protection against ectoparasites, as a repellent of predators, in addition to attracting mates and eliciting other pheromonal responses. In this study, all animals reacted by exuding malodorous substances when handled, as a form of defense. However, these are just assumptions that need to be clarified with additional studies on animal behavior. Anat Rec, 303:1489-1500, 2020. © 2019 American Association for Anatomy.

Heterochrony of puberty in the European badger (Meles meles) can be explained by growth rate and group-size: Evidence for two endocrinological phenotypes.

Puberty is a key stage in mammalian ontogeny, involving endocrinological, physiological and behavioural changes, moderated by intrinsic and extrinsic factors. Thus, not all individuals within one population achieve sexual maturity simultaneously. Here, using the European badger (Meles meles) as a model, we describe male testosterone and female oestrone profiles (using Enzyme-immunoassays) from first capture (3 months, post-weaning) until 28 months (attaining sexual maturity and final body size), along with metrics of somatic growth, scent gland development and maturation of external reproductive organs as well as intra-specific competition. In both sexes, endocrinological puberty commenced at ca. 11 months. Thereafter, cub hormone levels followed adult seasonal hormone patterns but at lower levels, with the majority of cubs reaching sexual maturity during their second mating season (22-28 months). Interestingly, there was evidence for two endocrinological phenotypes among male cubs (less evident in females), with early developers reaching sexual maturity at 11 months (first mating season) and late developers reaching sexual maturity at 22-26 months (second mating season). Early developers also attained a greater proportion of their ultimate adult size by 11 months, exhibiting faster growth rates than late developers (despite having similar adult size). Male cubs born into larger social groups tended to follow the late developer phenotype. Our results support the hypothesis that a minimum body size is required to reach sexual maturity, which may be achieved at different ages, even within a single population, where early maturity can confer individual fitness advantages and enhance population growth rate.

The pregenital abdomen of Enicocephalomorpha and morphological evidence for different modes of communication at the dawn of heteropteran evolution.

The internal and external anatomy of the posterior metathoracic region, pregenital abdomen, and associated nervous system of the heteropteran infraorder Enicocephalomorpha are thoroughly described, using an array of state-of-the art techniques. Based on morphology, it is hypothesised which modes of communication these insects use. This study is based primarily on an undescribed species of Cocles Bergroth, 1905 (Enicocephalidae) and another undescribed species of Lomagostus Villiers, 1958 (Aenictopecheidae), but additional representatives of the infraorder are also examined. Our results are compared with the literature on other Heteroptera. The metathoracic scent gland system of Enicocephalomorpha uses the same muscles as that of more derived Heteroptera, although the efferent system is different. The presence of a tergal plate and well-developed longitudinal musculature in the families Enicocephalidae and Aenictopecheidae, as well as a sexually dimorphic set of sclerites and membranes that allow an as yet undetermined type of motion, may indicate the presence of vibrational signaling in the infraorder, although experimental confirmation is required. Our findings raise new research questions regarding heteropteran functional morphology and communication.

Musk gland seasonal development and musk secretion are regulated by the testis in muskrat (Ondatra zibethicus).

BACKGROUND: The muskrat is a seasonal breeder. Males secrete musk to attract females during the breeding season. The testosterone binding to the androgen receptor (AR) in musk glands of muskrat may play an important role conducting the musk secretion process. METHODS: The musk gland, testis and blood samples of musk rats are collected in both breeding and non-breeding seasons. Some part of the samples are kept in liquid nitrogen for transcriptome analysis and Western blotting test. Some part of the samples are kept in 70% alcohol for histology experiment, blood samples are kept at -20 °C for the serum testosterone measurement experiment. RESULTS: This study demonstrates that the quantity of secreted musk, the volume of the musk glands, the diameter of the gland cells and AR expression are all higher during the breeding season than at other times (p < 0.01). StAR, P450scc and 3ß-HSD expression in the Leydig cells of the testis were also higher during this season, as was serum testosterone. AR was also observed in the gland cells of two other musk-secreting animals, the musk deer and small Indian civet, in their musk glands. These results suggest that the testes and musk glands co-develop seasonally. CONCLUSION: The musk glands' seasonal development and musk secretion are regulated by the testes, and testosterone plays an important role in the seasonal development of musk glands.

Musk gland seasonal development and musk secretion are regulated by the testis in muskrat (Ondatra zibethicus)

BACKGROUND: The muskrat is a seasonal breeder. Males secrete musk to attract females during the breeding season. The testosterone binding to the androgen receptor (AR) in musk glands of muskrat may play an important role conducting the musk secretion process. METHODS: The musk gland, testis and blood samples of musk rats are collected in both breeding and non-breeding seasons. Some part of the samples are kept in liquid nitrogen for transcriptome analysis and Western blotting test. Some part of the samples are kept in 70% alcohol for histology experiment, blood samples are kept at -20 °C for the serum testosterone measurement experiment. RESULTS: This study demonstrates that the quantity of secreted musk, the volume of the musk glands, the diameter of the gland cells and AR expression are all higher during the breeding season than at other times (p < 0.01). StAR, P450scc and 3ß-HSD expression in the Leydig cells of the testis were also higher during this season, as was serum testosterone. AR was also observed in the gland cells of two other musk-secreting animals, the musk deer and small Indian civet, in their musk glands. These results suggest that the testes and musk glands co-develop seasonally. CONCLUSION: The musk glands' seasonal development and musk secretion are regulated by the testes, and testosterone plays an important role in the seasonal development of musk glands.

The Scent Glands of the Neotropical Harvestman Discocyrtus pectnifemur: Morphology, Behavior and Chemistry.

Harvestmen have a pair of scent glands that open through ozopores. The literature suggests a link between the morphology of the ozopore area and the emission of a defensive secretion. A previous study on a species that aggregates in open areas, where individuals are probably more easily spotted by predators, showed that this defensive secretion causes conspecifics to flee. However, it is unknown whether this behavior occurs in species that aggregate in sheltered areas, where prey are harder to find. Herein, we describe the morphology of the ozopore area, the mode of emission of the defensive secretion, and its chemical composition in the harvestman Discocyrtus pectinifemur. We also tested if the defensive secretion is used as an alarm pheromone. We found that D. pectinifemur releases the defensive secretion in different ways, one of them being as a jet. Emission as a jet contrasts with that known for all congeners previously studied, and is in accord with the expected morphology of the ozopore. We found that the defensive secretion of D. pectinifemur does not function as an alarm pheromone. The composition of the defensive secretion, a mixture of quinones, is congruent with those already described for the clade that includes Discocyrtus. Our results support the link between the morphology of the scent glands area and the emission behavior of the defensive secretion, and they suggest that the alarm pheromone function in harvestmen may be dependent on ecological factors.

Neotropical harvestmen (Arachnida, Opiliones) use sexually dimorphic glands to spread chemicals in the environment.

Sexually dimorphic glands have convergently appeared in animals and are often responsible for the production of pheromones. In the suborder Laniatores of the order Opiliones (Arachnida), glands of such type are widespread, but there is not a single paper on how they are used. Using Scanning Electron Microscopy and a behavioral approach, we describe glandular openings and how these glands are used, in the harvestmen Gryne perlata and Gryne coccinelloides (Cosmetidae). Males of these two species have glandular openings on the metatarsi of legs I and on the metatarsi IV. Males were shown rubbing the glands of the metatarsi I against their other legs, whereas glands on the metatarsi IV are gently touched on the substrate or rubbed either against other legs, or against the substrate. Not all behaviors were seen in both species.

Gender differences in the anatomy of the perineal glands in Guinea pigs and the effect of castration.

Perineal glands in guinea pigs are part of the sebaceous glandular complex. Their secretions are used for scent marking. This is important for social status and can be seen in both sexes and castrated males. Discrepancy exits about the existence of these glands in female guinea pigs and knowledge of the anatomical consequences of castration on the male perineal glands is sparse. To examine these uncertainties related to gender, perineal glands from 13 sexually mature pet guinea pigs were examined macro- and microscopically. Clear gender differences in the anatomy of perineal glands were found, and castrated males showed signs of atrophy and fatty infiltration in the glands. Females do have perineal glands, although smaller than the glands in the male. The glands are typically sebaceous with multiple excretory ducts. A macroscopic unique feature in the males was the clearly evident orifices of a large excretory duct on each side of the slightly everted perineal sac. However, the reason for this gender difference is not clear. In castrated males, the orifices were atrophied and difficult to see. In addition, the sebaceous glands of the hair follicles in the skin folds of the perineal opening were smaller and less abundant in females and castrated males. The changes in castrated males are presumably linked to the hormonal changes and decreased secretion after castration. The dense keratin layer in the perineal sac was thicker in males than in both castrated males and females and could contribute to the concrement formation seen mainly in males.

The metathoracic scent gland of the leaf-footed bug, Leptoglossus zonatus.

The metathoracic scent gland of 25-day-old adults of both sexes of the leaf-footed bug, Leptoglossus zonatus (Dallas) (Heteroptera: Coreidae), are described based on optical microscopy analysis. No sexual dimorphism was observed in the glandular composition of this species. The gland is located in the anteroventral corner of the metathoracic pleura between the middle and posterior coxal pits. The opening to the outside of the gland is very wide and permanently open as it lacks a protective membrane. In the internal part, there is a pair of metathoracic glands that consist of piles of intertwined and occasionally bifurcated cellular tubes or columns. These glands discharge their pheromonal contents into the reservoir through a narrow cuticular tube. The reservoir connects with the vestibule via two opposite and assembled cuticular folds that can separate muscularly in order to allow the flow of liquid away from the insect. The external part consists of an ostiole from which the pheromone is emitted. The ostiole is surrounded by a peritreme, a structure that aids optimum pheromone dispersion. The described gland is of the omphalien type.

Occurrence of certain cuticular structures confirms functionality of dorsal abdominal scent glands in Acanthosomatidae (Heteroptera: Pentatomoidea).

Elasmucha ferrugata (Fabricius, 1787) (Heteroptera: Acanthosomatidae) provides maternal care of eggs and larvae. Adults of both sexes have functional anterior dorsal abdominal scent glands (DAGs). Study of all internal and external cuticular structures of DAGs revealed that no DAGs are fully functional in the 1st larval instar, and very probably they are only slightly functional in the 2nd instar. Median and posterior DAGs are undoubtedly not functional in adults. There exists sexual dimorphism in the number of multicellular glandular units in anterior glands in adults. The occurrence of cuticular ductules of these units proves these to be functional glands. This is best considered in combination with the findings of a well-developed evaporatorium. Developed cuticular intima of the gland reservoir and/or the nearly closed ostiole or ostiolar scar bears no information about the functionality of the gland.

Morphology and volatile compounds of metathoracic scent gland in Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae).

Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae) is a serious insect pest of litchi and longan in South China. When disturbed, this insect could release large quantities of disagreeable odorous volatiles from its scent gland. Knowledge on the scent gland and its secretion is crucial for developing the semiochemical methods to manage this pest. Morphology and ultrastructure of the metathoracic scent glands (MTGs) were studied under stereo and scanning electron microscopy, and the volatile compounds of MTGs from both male and female T. papillosa were analyzed with coupled gas chromatography-mass spectrometry (GC-MS). The MTG complex is located between the metathorax and the first abdominal segment at the ventral surface of the insect, which has a well-developed single double valve cystic-shaped orange median reservoir, paired colorless lateral glands in both sides, and a long and wavy tubular accessory gland that inlays tightly into the ventral edge around the median reservoir. The MTG opens to the body surface through paired ostioles located between the meso- and metacoxae of the evaporatorium with mushroom bodies. The GC-MS analyses showed that female and male adults have nine major volatile components in common. Tridecane is the most abundant in both females and males, reaching up to 47.1% and 51.8% of relative amount, respectively. The minor component is benzophenone with only 0.28% and 0.14%. Furthermore, undecane, tetradecane, 3-methyl-tridecane, and cyclopentadecane were found only in males. The possible function of volatile compounds of MTG contents in T. papillosa is addressed.

Functional anatomy of scent glands in Paranemastoma quadripunctatum (Opiliones, Dyspnoi, Nemastomatidae).

The morphological organization and functional anatomy of prosomal defensive (scent) glands in Paranemastoma quadripunctatum, a representative of the dyspnoid harvestmen, was investigated by means of histological semithin sections, software-based 3D-reconstruction and scanning electron microscopy. Scent glands comprise large, hollow sacs on either side of the prosoma, each of these opening to the outside via one orifice (ozopore) immediately above coxa I. In contrast to the situation known from laniatorean, cyphophthalmid and some eupnoid Opiliones, ozopores are not exposed but hidden in a depression (atrium), formed by a dorsal integumental fold of the carapace and the dorsal parts of coxae I. Glandular sacs are connected to ozopores via a short duct which is equipped with a specific closing mechanism in its distal part: A layer of modified epidermal cells forms a kind of pad-like tissue, surrounding the duct like a valve. Several muscles attached to the anterior parts of the glandular reservoir and to the epithelial pad may be associated with ozopore-opening. The actual mechanism of secretion discharge seems to be highly unusual and may be hypothesized on the basis of corroborating data from behavioral observations, scent gland anatomy and secretion chemistry as follows: Enteric fluid is considered to be directed towards the ozopores via cuticular grooves in the surface of the coxapophyses of legs I. Then, the fluid is sucked into the anterior part of the scent gland reservoirs by the action of dorsal dilator muscles that widen the reservoir and produce a short-term negative pressure. After dilution/solution of the naphthoquinone-rich scent gland contents, a secretion-loaded fluid is thought to be discharged with the help of transversal compressor muscles. This is the first detailed study on the functional anatomy of scent glands and the mechanisms of secretion discharge in the Dyspnoi.

The orbitofacial glands of bats: an investigation of the potential correlation of gland structure with social organization.

The facial glands of bats are modified skin glands, whereas there are up to three different orbital glands: Harderian, lacrimal, and Meibomian glands. Scattered studies have described the lacrimal and Meibomian glands in a handful of bat species, but there is as yet no description of a Harderian gland in bats. In this study we examined serial sections of orbitofacial glands in eight families of bats. Much variation amongst species was observed, with few phylogenetic patterns emerging. Enlarged facial glands, either sudoriparous (five genera) or sebaceous (vespertilionids only) were observed. Meibomian and lacrimal glands were present in most species examined (except Antrozous), though the relative level of development varied. Two types of anterior orbital glands were distinguished: the Harderian gland (tubulo-acinar: observed in Rousettus, Atribeus, Desmodus and Miniopterus) and caruncular (sebaceous: observed in Eptesicus and Dieamus). The relative development of the nasolacrimal duct and the vomeronasal organ did not appear to be correlated with the development of any of the exocrine glands examined. There does, however, appear to be a correlation between the presence of at least one well developed exocrine gland and the level of communality and known olfactory acuity, best documented in Artibeus, Desmodus, and Miniopterus.

Morphology of the femoral glands in the lizard Ameiva ameiva (teiidae) and their possible role in semiochemical dispersion.

Many lizards have epidermal glands in the cloacal or femoral region with semiochemical function related to sexual behavior and/or territorial demarcation. Externally, these glands are recognized as a row of pores, opening individually in the center of a modified scale. In many species the pores are used as systematic characters. They form a glandular cord or, in some species, a row of glandular beads below the dermis, and are connected to the exterior through the ducts, which continuously liberate a solid secretion. Dead cells, desquamated from the secretory epithelium, constitute the secretion, known as "a secretion plug." The present work focuses on the morphology of the femoral glands of the teiid lizard Ameiva ameiva, correlating it to the way in which the secretion is deposited in the environment. The results here obtained are compared to those available for other lizards and amphisbaenians. We observed that the diameter of the glandular pores did not show significant differences between males and females. The glands comprise germinative and secretory cells, which pass through at least three stages of differentiation, during which an accumulation of cytoplasmic granules, with a glycoprotein content, occurs. The cells eventually die and desquamate from the secretory epithelium, forming a secretory plug mostly constituted by juxtaposed nonfragmented secretory cells. Because of the arrangement of the rosette-like scales surrounding the femoral pores, we suggest that when the animal is in a resting position, with its femoral regions touching the ground, these scales may be involved in the breakage of their respective plugs, depositing tiny portions on the substrate. In this manner, it seems that the method for signal dispersion in this species involves specifically adapted structures and does not simply involve the chance breakage of the plug, as the gland secretes it. Signal dispersion must also be intimately associated with the animal's movement within its territory.

Putative chemical signals about sex, individuality, and genetic background in the preputial gland and urine of the house mouse (Mus musculus).

To explore whether preputial gland secretions and/or urine from the house mouse (Mus musculus) can be used for coding information about sex, individuality, and/or the genetic background of strain [ICR/albino, Kunming (KM), and C57BL/6], we compared the volatile compositions of mouse preputial glands and urine using a combination of dichloromethane extraction and gas chromatography coupled with mass spectrometry (GC-MS). Of the 40 identified compounds in preputial gland secretions, 31 were esters, 2 sesquiterpens, and 7 alcohols. We failed to find any compound unique to a specific sex, individual, or strain. However, many low molecular weight compounds between the sexes, most compounds among individuals, and several compounds among the 3 strains varied significantly in relative ratios. These quantitative differences in preputial gland volatiles (analog coding) are likely to convey information about sex, individual, and the genetic background of mouse strain. We identified 2 new main and male-elevated compounds, 1-hexadecanol (Z=3.676, P=0.000, N=19 in ICR; Z=3.576, P=0.000, N=18) and 1-hexadecanol acetate (Z=3.429, P=0.000, N=19 in ICR; Z=3.225, P=0.001, N=18), which were eluted in GC chromatogram after the 2 sesquiterpens. They might also be potential male pheromones, in addition to the well-known E-beta-farnesene and E,E-alpha-farnesene. Additionally, a few compounds including 1-hexadecanol also varied with strains and might also code for genetic information. Of the 9 identified volatile compounds in male urine, (s)-2-sec-butyl-4,5-dihydrothiazole and R,R-3,4-dehydro-exo-brevicomin are known urine-originated male pheromones from previous studies. We also detected 6-hydroxy-6-methyl-3-heptanone, a male urinary pheromonal compound, which had not been directly detected by GC-MS previously. Chemical analysis shows that the genetically more closely related ICR and KM strains had a higher similarity in the volatile compositions of preputial glands and urine than that between ICR or KM and C57BL/6. R,R-3,4-dehydro-exo-brevicomin, in particular, was sensitive to genetic shifts and differed in relative abundance among the 3 strains, whereas (s)-2-sec-butyl-4,5-dihydrothiazole differed between ICR or Km and C57BL/6. Hence, these 2 compounds might code for information about their genetic background.

Morphology of caprine skin glands involved in buck odour production.

The distribution and morphology of the cornual, sub-caudal, mental and preputial glands were studied macro- and microscopically in four Toggenburg and eight miniature male goats. Although the cornual and sub-caudal glands could be readily located macroscopically, the mental glands in the inter-mandibular region and the preputial glands at the preputial orifice were not visible macroscopically. On histological section, all glands were found to be composed of lobulated sebaceous tissue combining both normal and modified holocrine secretory units. Over a period of 18 months, five consecutive glandular swabs for scent tests were taken to assess the influence of age and season on buck odour production. Buck odour was most apparent in the cornual gland area, less distinct at the mental gland region, and faint or absent in the other glandular areas. Surgical removal of the cornual glands caused a decrease in buck odour and persisting scent was ascribed to smaller skin glands dispersed in the cranial body half. Complete absence of buck odour was only observed in castrated bucks.

A stingless bee (Melipona seminigra) marks food sources with a pheromone from its claw retractor tendons.

By depositing scent marks on flowers, bees reduce both the search time and the time spent with the handling of nonrewarding flowers. They thereby improve the efficiency of foraging. Whereas in honey bees the source of these scent marks is unknown, it is assumed to be the tarsal glands in bumble bees. According to histological studies, however, the tarsal glands lack any openings to the outside. Foragers of the stingless bee Melipona seminigra have previously been shown to deposit an attractant pheromone at sugar solution feeders, which is secreted at the tips of their tarsi. Here we show that the claw retractor tendons have specialized glandular epithelia within the femur and tibia of all legs that produce this pheromone. The secretion accumulates within the hollow tendon, which also serves as the duct to the outside, and is released from an opening at the base of the unguitractor plate. In choice experiments, M. seminigra was attracted by feeders baited with pentane extracts of the claw retractor tendons in the same way as it was attracted by feeders previously scent marked by foragers. Our results resolve the seeming contradiction between the importance of foot print secretions and the lack of openings of the tarsal glands.