Live for the moment—Adaptations in the male genital system of a sexually cannibalistic spider (Theridiidae, Araneae)

Monogyny in spiders culminates in extreme traits, like dramatic male self-sacrifice and emasculation of the male by the female during copulation. Here we show that monogynous males can be highly adapted for this fatal sexual behaviour. Dwarf males of the one-palped theridiid spider Tidarren argo, which are cannibalised immediately after the insertion of their single copulatory organ, stop spermiogenesis when reaching adulthood. Their testes atrophy, which might economise the energy expenditures of these males. We also found that the amount of seminal fluid produced is stored in an enlarged seminal vesicle until the single sperm induction takes place. The volume of the seminal vesicle is similar to the sperm droplet taken up into the copulatory organ (palpal organ). Sperm uptake takes much longer than in related species most likely due to the large amount of seminal fluid. As shown by histological observations males are able to fill one of the paired female sperm storage organs during copulation thereby presumably impeding subsequent charging by rival males.     

Synthesis of seminal ribonuclease in isolated lobules of bull seminal vesicles

A procedure is described for preparing and maintaining in culture isolated lobules of bovine seminal vesicles, consisting of glandular acini, surrounded by little connective tissue and with free access to the external medium, in which secreted material can be collected. After 48 h in culture, the isolated lobules appeared indistinguishable, by morphological and biochemical criteria, from freshly isolated lobules. After much longer culture times about one third of the glandular cells were still capable of effective protein synthesis. Studying the biosynthesis of seminal ribonuclease with preparations of isolated lobules we found that the enzyme was synthesized and secreted; only the fully amidated isoenzyme was synthesized and secreted, indicating that production of the selectively deamidated isoenzymic forms occurred after secretion, newly synthesized protein was rapidly exported, indicating that the high levels of enzyme previously reported for the seminal vesicle tissue were essentially due to its content of stored secretion.     

Male genital organs in the eulittoral meiofaunal polychaete <Emphasis Type="Italic">Stygocapitella subterranea</Emphasis> (Annelida,Parergodrilidae): ultrastructure,functional and phylogenetic significance

The marine interstitial polychaete Stygocapitella subterranea is characterized by aberrant morphological and biological traits resembling those of clitellates and Hrabeiella periglandulata, a terrestrial polychaete species. Although clearly related to the terrestrial Parergodrilus heideri, there are distinct differences in their morphology. An ultrastructural study of the male genital organs was undertaken to look for common apomorphic features in Parergodrilidae, to find structural evidence for clarifying their reproductive biology and mode of sperm transfer. Finally it should be elucidated whether a supposed sister-group relationship of Parergodrilidae and Orbiniidae based on molecular evidence can be supported by morphological characters as well. In S. subterranea the male organs consist of an unpaired seminal vesicle, a pair of sperm ducts and two large tube-like prostate glands. These glands constitute the distal parts of the gonoducts and open ventrally on a small genital papilla in chaetiger 9. True copulatory organs or organs for storage of mature sperm are lacking. The seminal vesicle is a coelomic cavity composed of two apposed coelomic linings supplied with blood spaces. The testes are found ventrally. The prostate glands are covered by a single layer of muscle fibres running in a longitudinal/spiral direction along the gland. There are no signs of spermatophore formation in any part of the male system. Since females always carry sperm, pseudocopulation can be excluded and the likelihood of either direct transfer of sperm or hypodermic injection is discussed. The structure of genital organs reveals similarities to those of P. heideri. Gonochorism, paired seminal vesicles and two pairs of male gonoducts opening in chaetigers 9 and 10 with a distal glandular part most likely belong to the ground pattern of Parergodrilidae. The observations confirm that consistencies with either clitellates or H. periglandulata are the result of convergent evolutionary events. On the other hand, the relationship of Parergodrilidae to an orbiniid/questid clade receives support from the present data. This paper is dedicated to our scientific teacher, Professor Wilfried Westheide, who made significant contributions to the reproductive biology of interstitial polychaetes, on the occasion of his 70th birthday.     

Steroid glucuronides as male pheromones in the reproduction of the African catfish Clarias gariepinus—a brief review

After ovulation, female African catfish are strongly attracted by the odor of male conspecifics. This attraction depends on the presence of the seminal vesicle, a part of the male reproductive organs. Removal of the seminal vesicle illustrates this fact. A low dose of seminal vesicle fluid, added to the water, appears to be highly attractive for catfish which have ovulated. Fractionation of the fluid and testing of the different fractions shows that steroid glucuronides could be responsible for the attraction. These steroid glucuronides can be identified with gas chromatographic-mass spectrometric analysis. A mixture of glucuronides, prepared to resemble the composition of the seminal vesicle fluid, evokes a dose-dependent attraction. The most potent odorant, observed by measuring electrical responses from the olfactory epithelium and from the olfactory tract appears to be 3α,17α-dihydroxy-5β-pregnan-20-one-3α-glucuronide.     

The seminal vesicle of the African catfish,Clarias gariepinus

Summary The seminal vesicle of the African catfish, Clarias gariepinus, consists of 36–44 fingerlike lobes built up of tubules in which a fluid is secreted containing acid polysaccharides, acid-, neutral- and basic proteins, and phospholipids. In this fluid sperm cells are stored. The seminal vesicle fluid immobilizes the sperm cells. After ejaculation, it prolongs the period of sperm activity. The seminal vesicle fluid is secreted by the epithelium lining the tubules. The tubules in the proximal part of the lobes are predominantly lined by a simple cylindrical and those of the distal part by a simple squamous epithelium. These epithelial cells contain enzymes involved in energy-liberating processes, the enzyme activites being proportional to the height of the cells. Interstitial cells between the tubules have enzyme-histochemical and ultrastructural features indicative of steroid biosynthesis. Similar characteristics are found in testicular interstitial cells. The most rostral seminal vesicle lobes and the most caudal testicular efferent tubules form a network of tubules that opens at the point where the paired parts of the sperm ducts fuse with each other. The tubules of most seminal vesicle lobes, however, form a complex system that fuses with the unpaired part of the sperm duct.     

Ultrastructure of the male reproductive organs in the interstitial annelid Sphaerosyllis hermaphrodita (”Polychaeta”, Syllidae)

 The reproductive organs of the simultaneous hermaphrodite Sphaerosyllis hermaphrodita (Syllidae, Exogoninae) were examined by TEM and reconstructed from ultrathin serial sections. Oocytes are produced in the 11–13th chaetigerous segments and then attached to the outer body surface. The male organs comprise a seminal vesicle, testes, sperm ducts and copulatory chaetae. The unpaired seminal vesicle is an uncompartmented cavity above the gut and within the chaetigerous segments 8–10. Its interior is lined with a layer of gland cells that degenerate as spermatogenesis in the vesicle proceeds. The testes are situated ventrolaterally, close to the seminal vesicle in the 9th chaetigerous segment. They contain cells at early stages of spermatogenesis, which are connected to one another by zonulae collares. The testes and seminal vesicle are enclosed in epithelia. Paired sperm ducts run ventrally from about the midline of the body under the seminal vesicle and into the parapodia of the 9th chaetigerous segment. There they open, together with the protonephridia of this segment, to the outside next to the stout copulatory chaeta. Each sperm duct consists of six cells, the luminal surface of which bears microvilli but no cilia. Only in animals with fully differentiated sperm does the small opening of the proximal duct cell in each duct give access to the seminal vesicle. The mode of sperm transfer is discussed. Accepted: 9 December 1996     

Ultrastructure of the male reproductive system and spermatophore formation in Labidocera aestiva (Crust.: Copepoda)

Summary The male reproductive system of Labidocera aestiva produces a flask-shaped spermatophore connected to a chitin-like coupling apparatus. As immature spermatozoa leave the anterior region of the testis, they pass through the lumen of a long, sinuous duct composed of a ductus deferens and seminal vesicle. Ultrastructural examination of the ductus deferens reveals a highly glandular, columnar epithelium. The cells contain arrays of rough endoplasmic reticulum and abundant, well-developed Golgi complexes. This region produces and releases into the lumen, a flocculent substance and two granular secretions that constitute the seminal fluid. In its terminal part, the ductus deferens synthesizes another secretion that forms the spermatophore wall enclosing the spermatozoa and seminal fluid. Final synthesis of the spermatophore wall occurs within the thin-walled seminal vesicle, although this region functions primarily as a storage organ. Contiguous to the seminal vesicle is an elongate, highly glandular spermatophore sac. The chitin-like coupling apparatus, which functions to attach the spermatophore to the female, is formed in the anterior region of the sac by secretions from eight cell types. The posterior region of the sac stores the flask-shaped spermatophore and produces secretions that aid ejaculation of the entire spermatophore complex.Contribution No. 236, Harbor Branch Foundation, Inc.     

Anatomy and ultrastructure of the male reproductive system in Pleioplana atomata (Platyhelminthes: Polycladida)

Abstract. The ultrastructure of the male reproductive system in the polyclad flatworm Pleioplana atomata is described. Numerous testes are scattered throughout the entire body but are heavily concentrated on the ventral side. All stages of differentiating sperm cells are present in all testes follicles. Intercellular bridges connect spermatocytes and spermatids derived from a single spermatogonium. In the distal part of spermatids, a zone of differentiation develops with a row of microtubules beneath the plasmalemma. Adjacent to these microtubules, an intercentriolar body is flanked by two basal bodies that give rise to two axonemes (each with a 9+“1” microtubular pattern) that face in opposite directions. The Golgi complex appears in the central portion of the spermatid and produces numerous small and large electron-dense bodies. The small bodies surround the nucleus, whereas the large bodies cluster along with the mitochondria in the central part of the spermatid. Development of the spermatid leads to cell elongation and formation of a filiform, biflagellate mature spermatozoon with cortical microtubules all along the sperm shaft. The male canal system consists of paired vasa deferentia that separately enter a single seminal vesicle. A single prostatic canal connects the seminal vesicle to the prostatic vesicle. Ultrastructurally, the seminal vesicle and prostatic canal are very similar, and along with the prostatic vesicle and stylet pocket, are lined by a ciliated epithelium. The ultrastructure of the prostatic vesicle indicates that it probably produces a large volume of seminal fluid that, along with spermatozoa, is transferred to the mating partner through a stylet. Some of the findings, particularly on sperm ultrastructure, may provide characters useful for phylogenetic analysis.     

Morphology of the male reproductive system of freshwater prawn Macrobrachium carcinus (Decapoda,Caridea): Functional and comparative aspects

Testes and vasa deferentia are parts of the male reproductive system of decapod crustaceans. Both organs show morphological differences among decapod species in terms of anatomical and histological patterns reflecting the diversity of this group. Describing these features may assist in systematics, phylogenetics, and studies of reproductive behavior, especially for species of commercial interest, such as Macrobrachium carcinus, a native American species that, unusually for this genus, has no precopulation courting behavior. This study aims to describe the reproductive morphology and spermatogenesis of the male freshwater prawn M. carcinus. The male reproductive system of this species consisted of lobed testes connected to the vasa deferentia. The testis of M. carcinus was divided into several lobules. Each lobule was formed by a cluster of germ cells surrounded by connective tissue and nurse cells. This microscopic anatomy and histology of the testicular histoarchitecture has been described for many species of Decapoda and may represent a derived design of the testes. Unlike that in other decapod species, spermatogenesis proceeds in short transitory phases that produce spermatozoa at high concentrations and frequencies, corroborating the uncommon male reproductive behavior of this species. In the spermatic pathway, the lobules develop and fuse before releasing spermatozoa from the testes; however, this process has not been observed in decapods, yet. The neutral compounds secreted by the vas deferens are important for sperm nutrition as females secrete a substance for spermatophore adhesion during reproduction. This study presents different features and dynamics of the spermatogenic process in the male reproductive system of M. carcinus that have not yet been presented in the literature for decapods.     

Steroid glucuronides as male pheromones in the reproduction of the African catfish Clarias gariepinus--a brief review

After ovulation, female African catfish are strongly attracted by the odor of male conspecifics. This attraction depends on the presence of the seminal vesicle, a part of the male reproductive organs. Removal of the seminal vesicle illustrates this fact. A low dose of seminal vesicle fluid, added to the water, appears to be highly attractive for catfish which have ovulated. Fractionation of the fluid and testing of the different fractions shows that steroid glucuronides could be responsible for the attraction. These steroid glucuronides can be identified with gas chromatographic-mass spectrometric analysis. A mixture of glucuronides, prepared to resemble the composition of the seminal vesicle fluid, evokes a dose-dependent attraction. The most potent odorant, observed by measuring electrical responses from the olfactory epithelium and from the olfactory tract appears to be 3,17-dihydroxy-5β-pregnan-20-one-3-glucuronide.     

The spermiogenesis and the early spermatozoa of <Emphasis Type="Italic">Armorloricus elegans</Emphasis> (Loricifera,Nanaloricidae)

The spermiogenesis consisting of five spermatid stages and the early spermatozoon has been investigated in Armorloricus elegans (Loricifera) with the use of transmission electron microscopy. The male reproductive system consists of three parts; testes, vasa deferentia and seminal vesicles. Caudally, the two seminal vesicles merge together in a ciliated duct and the excretory/gonadal—and digestive systems continue through the recto-urogenital canal, which opens via the lateral gonopores and the temporarily closed anal system. Spermiogenesis mainly occurs in the testes, whereas further maturation of the late spermatids and early spermatozoa occurs in the vasa deferentia and seminal vesicles. A maturation gradient (from spermatocytes to spermatozoa) is found from the posterior peripheral part of the testes to the anterior periphery and then centrally. During spermiogenesis the round nucleus becomes more osmiophilic and condensation of chromatin occurs. Later the nucleus elongates until it becomes rod-shaped in the early spermatozoa. In the second spermatid stage, a large vesicle is formed by saccules developed from the Golgi complex. This vesicle develops further and consists of three different osmiophilic parts with some crystal-like structures inside and is on the outside almost entirely surrounded by thick striated filaments. In the mid-piece the flagellum has a typical 9 × 2 + 2 axoneme and the two mitochondria are fused into a single sheet surrounding the flagellum. In the early spermatozoon stage an acrosomal-like cap structure with an acrosome filament appears proximal to the protruded rod-shaped nucleus. This cap is not formed by the Golgi complex and therefore might not be a true acrosome. Comparing the early spermatozoa of A. elegans with other cycloneuralians has shown some similarities with especially Kinorhyncha and Priapulida. These similarities are thought to be plesiomorphic.     

Ultrastructure and function of the seminal vesicle of Bittacidae (Insecta: Mecoptera)

The fine structure of the seminal vesicle and reproductive accessory glands was investigated in Bittacidae of Mecoptera using light and transmission electron microscopy. The male reproductive system of Bittacidae mainly consists of a pair of testes, a pair of vasa deferentia, and an ejaculatory sac. The vas deferens is greatly expanded for its middle and medio-posterior parts to form a well-developed seminal vesicle. The seminal vesicle is composed of layers of developed muscles and a mono-layered epithelium surrounding the small central lumen. The epithelium is rich in rough endoplasmic reticulum and mitochondria, and secretes vesicles and granules into the central lumen by merocrine mechanisms. A pair of elongate mesodermal accessory glands opens into the lateral side of the seminal vesicles. The accessory glands are similar to the seminal vesicle in structure, also consisting of layers of muscle fibres and a mono-layered elongated epithelium, the cells of which contain numerous cisterns of rough endoplasmic reticulum and mitochondria, and a few Golgi complexes. The epithelial cells of accessory glands extrude secretions via apocrine and merocrine processes. The seminal vesicles mainly serve the function of secretion rather than temporarily storing spermatozoa. The sperm instead are temporarily stored in the epididymis, the greatly coiled distal portion of the vas deferens.     

Comparative studies of testicular structure and sperm morphology among copulatory and non-copulatory sculpins (Cottidae: Scorpaeniformes: Teleostei)

Testicular structure of 9 species and sperm head morphology of 19 species of Cottidae were observed in order to clarify relationships between morphological characteristics of the male reproductive organ and reproductive mode (copulation or non-copulation). Morphological structure of the testis was divided into the following five types based on the sperm transfer and reservoir system: (1) a non-duct type in which the sperm duct is not a distinct exterior structure, but the tube for sperm transport traverses along the testis as an interior structure; (2) an anterior duct type with distinct anterior sperm ducts traversing along the testis; (3) a posterior duct type with distinct anterior sperm ducts traversing along the dorsal hilus of testis and posterior sperm ducts extending to the rear of the testis; (4) an anterior duct posterior vesicle type with distinct anterior sperm ducts traversing along the testis, and the right and left sperm ducts fusing in the rear of testis, forming the seminal vesicle; (5) a non-duct posterior vesicle type in which sperm ducts do not accompany the testis, and the testis and seminal vesicle are connected directly or through posterior sperm ducts. It is thought that in Cottidae the non-duct type of reproductive organ is primitive, and the anterior duct type is common to all non-copulating species. The testes and accompanying seminal vesicle were seen only in copulating species. Sperm head morphology was divided into three types according to the length/width ratio: oval type ≤2, intermediate type >2 and ≤3, and slender type >3. The type of sperm head corresponded closely to the reproductive mode; non-copulating species had oval sperm head, and copulating species had intermediate or slender ones. These results suggest that the structure of the testis and the morphology of the sperm head evolved from testes with anterior sperm ducts and oval sperm heads to testes with an associated seminal vesicle and slender sperm heads in association with the evolution from non-copulatory to copulatory reproduction in Cottidae.     

3种蛾类雄性内生殖系统形态学研究

为了探究蛾类昆虫新的有效分类特征,本研究对直脉青尺蛾Geometra valida、白雪灯蛾Chionarctia niveus和多斑豹蠹蛾Zeuzera multistrigata雄性内生殖系统进行了解剖观察和形态学比较。结果表明：3种蛾类雄性内生殖系统均由精巢、贮精囊（前贮精囊和后贮精囊）、输精管、复射精管、单射精管（单射精管原节和单射精管表皮节）和附腺组成;种间形态差异表现在贮精囊的形状及连接方式、复射精管的形状及长短、单射精管和附腺的长度等方面,这些鉴别特征可为蛾类分类及系统发育研究提供形态学借鉴。     

Positive effects of testosterone and immunochallenge on energy allocation to reproductive organs

A number of studies have suggested the incompatibility of simultaneous increases in immune and reproductive functions. Other research has indicated that immune responses may be modulated depending on the relative benefits of increased survival and prospects for current and future reproduction. We tested the hypothesis that energy allocation to reproductive and other organ systems is not affected by testosterone level and energy expenditure on immune functions. Adult male white-footed mice (Peromyscus leucopus) with or without elevated testosterone levels and with or without immunochallenges were tested. Testosterone treatment was associated with reduced humoral immune response indicating immunosuppressive effects, reduced masses of gastrointestinal organs, reduced corticosterone level, increased kidney and seminal vesicle masses, and increased hematocrit. Immunochallenge was associated with increased resting metabolic rate and testes and seminal vesicle masses. Reproductive organ masses were greatest in immunochallenged mice with exogenous testosterone. Simultaneous increases in energy allocation to immune and reproductive structures may be an adaptive response that would enhance survival and current prospects for reproduction.     

Development of the reproductive system of Echinostoma paraensei in Mesocricetus auratus analyzed by light and confocal scanning laser microscopy

This study was performed to gain insight into the maturation of the reproductive system of Echinostoma paraensei worms grown in an early infection of Mesocricetus auratus. Hamsters were infected with 100 metacercariae and necropsied on days 3, 5, 7, 10 and 14 post infection (dpi). Recovered flukes stained with hydrochloric carmine were preserved as whole mounts and analyzed by light and confocal scanning laser microscopy. The average worm recovery was 43.7 per host. Images of the male and female reproductive systems were taken. The ovary and anterior and posterior testis were evidenced on day 3, while the ootype and cirrus sac were present on day 5. Confocal imaging showed primordium testis and ovary as a cluster of primordial cells from day 3 onward. The testes, ovary, cirrus sac and uterus organs were already present during the first week of life. The two testes were seen as individual structures on 7 dpi while the cirrus sac and vitelline glands were in development. The ovary was connected to the uterus while the ootype was adjacent to it. Both testes were larger than the ovary, showing cells at different stages of development, but with few bundles of functional spermatozoa in 10 day-old worms. On day 14, eggs and spermatozoa were seen in the uterus and seminal vesicle, respectively, while oocytes appeared in the ootype as fertilized eggs. We conclude that the reproductive system of E. paraensei was functional on 14 dpi in the hamsters.     

The anatomy of the adult uropodid Fuscouropoda agitans (Arachnida; Acari), with comparative observations on other Acari

A thin, compressible, lateral suture and ventral plate overlap permit limited movement of the thick and rigid dorsal and ventral plates of Fuscouropoda agitans. Seven pairs of large dermal glands debouch onto the surface. Trochanteral rotation permits defensive leg adpresion and an insectan type of ambulation. The complex hypopharynx-pedipalpal-coxae has a buccal and cheliceral cavity separated by an atriculated epipharynx. The pharynx is Y-shaped in cross section. Extensive paired salivary glands lie above the very long and dexterous 3-segmented chelicerae, and a large pair of coxal glands debouch on coxae 1. From four blunt-ended tracheae, bundles of unbranching tracheoles extend in specific tracts to all organs. The ventriculus is small with three pairs of large caeca; a tightly packed single layer of digestive cells individually enlarged to absorb-phagocytize and digest the food. A typical mesostigmatid excretory tube is present. A typical acarine synganglion is present; mixed nerves have a basal swelling. A postulated neurosecretory organ arises from the pedipalpal nerve. The oocytes enlarge within funicular stalks from the walls of the small median ovary. A large spermatophore is stored in the seminal vesicle; fertilization occurs during oviposition. A tension hinge partially opens both male and female genital plates; closure effected by muscles acting on very long genital plate apodemes. Within sequentially produced spermatogonial cysts of the testes, meiosis is completely synchronous. A large, multilobed male accessory gland produces a large volume of seminal fluid; a mixture of at least four secretions. The origins and msertions of the body wall, genital organ, digestive tract, mouthpart and leg muscles are listed and illustrated. A comparison of anactinotrichid and actinotrichid mites indicates fundamental and consistent morphological differences in aspects of the cuticle, leg articulations, digestive system, excretory system, reproductive system and coxal glands.     

Ultrastructure and histochemistry of the male reproductive system of the genus Callinectes Stimpson, 1860 (Brachyura: Portunidae)

We described the ultrastructure and histochemistry of the reproductive system of five Callinectes species, and evaluate the seasonal variation in weight of the reproductive system and hepatopancreas by comparing annual changes of somatic indices. The somatic indices changed little throughout the year. In Callinectes, spermatogenesis occurs inside the lobular testes and, within each lobule, the cells are at the same developmental stage. Spermatogenesis and spermiogenesis follow the same development pattern in all Callinectes studied. Mature spermatozoa are released into the seminiferous ducts through the collecting ducts. Cells of the vas deferens are secretory as evidenced by rough endoplasmic reticulum, Golgi complex, and secretory vesicles that produce the seminal fluid. The anterior vas deferens shows two portions: proximal and distal. In proximal portion (AVDp), spermatozoa are clustered and embedded in an electron-dense, basophilic glycoproteinaceous secretion Type I. In the distal portion (AVDd), the spermatophore wall is formed by incorporation of a less electron-dense glycoproteinaceous secretion Type II. The secretion Type I change to an acid polysaccharide-rich matrix that separates the spermatophores from each other. The median vas deferens (MVD) stores the spermatophores and produces the granular glycoproteinaceous seminal fluid. The posterior vas deferens (PVD) has few spermatophores. Its epithelium has many mitochondria and the PVD seminal fluid changes into a liquid and homogeneous glycoprotein. Many outpocketings in the PVD and MVD help to increase the fluid production. Overall, the reproductive pattern of Callinectes is similar to other species that produce sperm plugs. The secretions of AVD, MVD, and PVD are responsible for the polymerization that forms the solid, waxy plug in the seminal receptacle. The traits identified here are common to all Portunidae species studied so far.     

Male traits associated with alternative reproductive tactics in Gobius niger

Male black goby Gobius niger , adopting parental or sneaking tactics, differed in secondary sex traits (elongation of the 4th ray of the first dorsal fin and black nuptial colouration) thus allowing the classification of wild-caught males. Parental males were larger and older than sneaker males, suggesting that the mating tactic is an expression of an ontogenetic gradient. Males adopting alternative tactics differed also in primary sex traits, including their testes and their two pairs of accessory structures: the seminal vesicles and mesorchial glands. Sneaker males had a higher investment in testes, while parental males showed larger seminal vesicles and more developed mesorchial glands. Histological analyses also showed that seminal vesicles from parental males presented some functional differences from those of sneakers. In the former these organs were devoted solely to mucin secretion, while in the latter they stored sperm and had a lower activity of secretion. Seminal vesicle features influenced ejaculate (sperm trail) characteristics and performance. Parental male trails were richer in mucins, but poorer in sperm than trails deposited by sneakers. As a consequence, while sneakers produced trails that released a large amount of immediately active sperm, trails laid by parental males released less sperm more constantly over a long time.