Spermatozoa of an Old World Ricinulei (Ricinoides karschii, Ricinoidae) with notes about the relationships of Ricinulei within the Arachnida

The ultrastructure of spermatozoa is a valuable tool for phylogenetic and systematic studies. Ricinulei are enigmatic and poorly studied arachnids. So far, spermatozoa are only known from New World ricinuleids. The goals were to study, by means of light and transmission electron microcopy, the spermatozoa of an Old World species with regard to their phylogenetic implications, e.g., does the sperm structure contribute to the debated sister-group relationship of Acari and Ricinulei. The spermatozoa are coiled-flagellate and characterized by a cap-like acrosomal vacuole covered by electron-dense material, an elongated nucleus covered by a manchette of microtubules during spermiogenesis, an axoneme with a 9+2 microtubular pattern, a nuclear tube and axonemal basis which both originate underneath the acrosomal vacuole and cleistospermia as transfer form equipped with three intracellular plates. The data of the present study did not support a close relationship of Ricinulei and Acari which have aflagellate sperm with various synapomorphies as e.g., lacking nuclear envelopes/membranes in Actinotrichida (very similar to Solifugae) or vacuolated spermatozoa in Anactinotrichida. Affinities of Ricinulei are discussed in the light of the ultrastructure of arachnid spermatozoa.     

Cellular details of the midgut of Cryptocellus boneti (Arachnida: Ricinulei)

The midgut of Cryptocellus boneti was studied by light and electron microscopy. The epithelia of the diverticula and of the anterior part of the midgut tube are composed of two cell types: digestive and secretory. In contrast, the epithelia of posterior part of the midgut tube and of the stercoral pocket consist of one type of cells only. In some places, parts of the midgut system are connected by an intermediate tissue. Digestive cells are characterized by an apical system of tubules, nutritional vacuoles, and spherites; characteristic features of secretory cells are secretory granules and a prominent rough endoplasmic reticulum. Cells of the midgut tube appear not to be involved in the absorption of food. © 1994 Wiley-Liss, Inc.     

Ultrastructure of tarsal sensilla and other integument structures of two Pseudocellus species (Ricinulei, Arachnida)

Ricinuleids are one of the least investigated groups of Arachnida. In particular, knowledge of their ultrastructure is poor. Observations of the distal tarsomeres of ricinuleids show differences in their shape and equipment of surface structures. Legs I and II are used by the Ricinulei to explore their surroundings with tentative movements. The tarsomeres of these legs show similarities in shape and surface structures that distinguish them from those of legs III and IV. In this study, 11 different structures of the tarsomere surfaces of two cave-dwelling species, Pseudocellus pearsei and P. boneti from México, were investigated for the first time with scanning and transmission electron microscopy and discussed regarding their possible function: 1) a single treelike ramifying seta resembles a no pore single-walled (np-sw) sensillum; 2) setae occurring in a small number and possessing a bipartite shaft represent terminal pore single-walled (tp-sw) sensilla. The surface of the proximal half of the shaft shows small branches. The distal half has a smooth surface; 3) long setae with conspicuous longitudinal lamellae show characteristics of chemoreceptive wall pore single-walled (wp-sw) sensilla; 4) frequent small wp-sw sensilla with flat and irregular lamellae; 5) very short wp-sw sensilla occurring solitary or in groups; 6) a few short setae with smooth surface correspond to wp-sw sensilla; 7) a single short clubbed seta articulating in a flat pit is considered to be a np-sw sensillum; 8) common long setae with a pointed tip show characteristics of mechanoreceptive np-sw sensilla; 9) ventral setae with adhesive and mechanosensory function are accompanied by multicellular "class III" glands; 10) slit organs with mechanoreceptive function; and 11) dome-like tubercles with no indication of sensorial function. Several of these sensilla form a sensory field on the dorsofrontal surface which is particularly pronounced on the distal tarsomeres of legs I and II.     

The pedipalp of Pseudocellus pearsei (Ricinulei, Arachnida) - ultrastructure of a multifunctional organ

Ricinulei possess movable, slender pedipalps with small chelae. When ricinuleids walk, they occasionally touch the soil surface with the tips of their pedipalps. This behavior is similar to the exploration movements they perform with their elongated second legs. We studied the distal areas of the pedipalps of the cavernicolous Mexican species Pseudocellus pearsei with scanning and transmission electron microscopy. Five different surface structures are characteristic for the pedipalps: (1) slender sigmoidal setae with smooth shafts resembling gustatory terminal pore single-walled (tp-sw) sensilla; (2) conspicuous long, mechanoreceptive slit sensilla; (3) a single, short, clubbed seta inside a deep pit representing a no pore single walled (np-sw) sensillum; (4) a single pore organ containing one olfactory wall pore single-walled (wp-sw) sensillum; and (5) gustatory terminal pore sensilla in the fingers of the pedipalp chela. Additionally, the pedipalps bear sensilla which also occur on the other appendages. With this sensory equipment, the pedipalps are highly effective multimodal short range sensory organs which complement the long range sensory function of the second legs. In order to present the complete sensory equipment of all appendages of the investigated Pseudocellus a comparative overview is provided.     

Obsevations on the biology of Ricinulei (Arachnida) with descriptions of two new species of Cryptocellus

Two new species of Ricinulei belonging to the genus Cryptocellus are described from British Guiana. The habitat is described and figured and a method of keeping Rcinulei in captivity reported. Observations on feeding and mating are described and possible evolutionary implications of the mating behaviour are discussed     

Female genital morphology and mating behavior of Orchestina (Arachnida: Araneae: Oonopidae)

The unusual reproductive biology of many spider species makes them compelling targets for evolutionary investigations. Mating behavior studies combined with genital morphological investigations help to understand complex spider reproductive systems and explain their function in the context of sexual selection. Oonopidae are a diverse spider family comprising a variety of species with complex internal female genitalia. Data on oonopid phylogeny are preliminary and especially studies on their mating behavior are very rare. The present investigation reports on the copulatory behavior of an Orchestina species for the first time. The female genitalia are described by means of serial semi-thin sections and scanning electron microscopy. Females of Orchestina sp. mate with multiple males. On average, copulations last between 15.4 and 23.54 min. During copulation, the spiders are in a position taken by most theraphosids and certain members of the subfamily Oonopinae: the male pushes the female back and is situated under her facing the female's sternum. Males of Orchestina sp. possibly display post-copulatory mate-guarding behavior. The female genitalia are complex. The genital opening leads into the uterus externus from which a single receptaculum emerges. The dorsal wall of the receptaculum forms a sclerite serving as muscle attachment. A sclerotized plate with attached muscles lies in the posterior wall of the uterus externus. The plate might be used to lock the uterus during copulation. The present study gives no direct evidence for cryptic female choice in Orchestina sp. but suggests that sexual selection occurs in the form of sperm competition through sperm mixing.     

Sperm transfer during mating in the pharaoh's ant, Monomorium pharaonis

Abstract.  Sperm transfer in the pharaoh's ant Monomorium pharaonis (L.) is studied by making longitudinal sections through the gasters of mating pairs fixed in copula. Sperm is transferred inside a spermatophore similar to those found in two other ants, Diacamma sp. from Japan and Carebara vidua . Sharp teeth-ridges are present on the penis valves and, during copulation, these teeth make contact with a thick and soft cuticular layer covering the bursa copulatrix. This ensures an attachment long enough for the successful transfer of the spermatophore to the right position inside the female oviduct. The thick cuticle also protects the queen from serious damage by the male's sharp claspers. After a first successful copulation, sperm is still present inside the male's seminal vesicles, suggesting that males can mate multiply. Additional experiments, where single, initially virgin males are presented to several virgin females, confirm this.     

Gross morphology,histology, and ultrastructure of the alimentary system of Ricinulei (Arachnida) with emphasis on functional and phylogenetic implications

Ricinuleid functional mouthparts are the cucullus, the chelicerae, the pedipalps, and the labrum. These structures are movably jointed to the rest of the prosoma, most likely protruded upon hydrostatic hemolymph pressure and retracted by prosomal muscles. Seta‐like protrusions from the labrum and the pedipalpal coxae form a sieve‐like filter inside the preoral cavity and the mouth. Although the tip of the labrum can be elevated upon muscle constriction, ingestion of large, solid food particles is unlikely. The mouth has a crescent‐shaped cross section. The cuticle‐lined, also crescent‐shaped pharynx is equipped with a large dilator muscle but lacks antagonistic constrictor muscles. It represents a precerebral sucking pump. The triangular to Y‐shaped, cuticle‐lined esophagus is equipped with constrictor and dilator muscles. Its posterior part represents a postcerebral sucking pump. Four blind ending diverticula ramify from the anterior prosomal part of the entodermal midgut tube. Two of these diverticula remain inside the prosoma and form few short branches. The other two extend through the pedicel into the opisthosoma and ramify and coil there. A stercoral pocket protrudes ventrally out of the midgut tube. The most distal part of the midgut tube is modified into a contractile rectal gland. Its secretions may have defensive or physiological functions. A short anal atrium is formed by the cuticle‐lined ectodermal hindgut which opens at the end of the three‐segmented metasoma. The telescoping segments of the metasoma are protruded by hemolymph pressure and retracted by muscles. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Sperm investment and alternative mating tactics in bluegill sunfish (Lepomis macrochirus)

Although alternative mating tactics are found in males of manyspecies, little is known about tactic-specific adaptations tosperm competition and the mechanism by which fertilization successis obtained. We now report on the sperm investment patternsof males that use alternative mating tactics in bluegill sunfish(Lepomis macrochirus). Bluegill males are characterized by threealternative mating tactics: parentals, sneakers, and satellites.Parentals delay maturation and adopt a courting and guardingtactic, whereas sneakers and satellites mature precociouslyand, respectively, use sneaking and female mimicry to stealfertilizations from parentals. We found differences among thetactics in testes weight, sperm longevity, and ejaculate spermdensity and competitiveness. Parentals had the largest testes,but the smallest relative to their body weight. Satellites hadintermediate-sized testes, and sneakers had the smallest testes,but the largest relative to their body weight. Within each tactic,there was a positive relationship between testes weight andbody weight, but the exact relationship differed among the tacticsand could not be attributed solely to allometry. Parentals hadlonger-lived sperm compared with that of both sneakers and satellites.Ejaculate sperm density was greatest in sneakers, satelliteswere intermediate, and parentals had the lowest. Competitionexperiments involving equal volumes of ejaculate showed thatfertilization success increased linearly with sperm number,consistent with a rafflelike process. However, independent ofsperm number, parental sperm were superior at fertilizing eggs.We interpret these sperm investment patterns in relation todifferences in sperm competition risk, number of mating opportunities,and alternative investment options.     

Die Calyptopis- und Furcilia-Stadien vonEuphausia hanseni (Crustacea: Euphausiacea)

The larval development is discussed and calyptopis stages I–III as well as furcilia stages I–VII are described. The larval development is similar to that ofE. spinifera, E. longirostris andE. triacantha (group d of genusEuphausia). Antennule, antenna, mandible, maxillule, maxilla and spines on the telson are unsuitable for identifying the larvae of different species of the genusEuphausia. The following characteristics can be used to distinguish the different larvae: form and marginal spines of the carapace, form of the rostrum, spines on the segments of the abdomen, and number and form of the pleopods. The mandible of larval Euphausiacea produces an asymmetric lacinia mobilis and a pair of processus incisivus accessorius (spine row). These two appendages are lacking in adult Eucarida.

---

---

Mitglied der Taxonomischen Arbeitsgruppe an der Biologischen Anstalt Helgoland     

Sperm morphology of the neotropical harvestman Iporangaia pustulosa (Arachnida: Opiliones): comparative morphology and functional aspects

We describe herein the sperm morphology of the harvestman Iporangaia pustulosa. Adult males were dissected, the reproductive tract was schematized and the seminal vesicle was processed by light, transmission and scanning electron microscopy. The male reproductive tract is composed of a tubular testis, two deferent ducts, a seminal vesicle, a propulsive organ and a penis, similar to that observed in other Opiliones. The spermatozoa from the seminal vesicle are oval, aflagellate and immotile, presenting a nucleus surrounding an invagination of the cytoplasm, as well as a complex acrosome and projections on the cell surface. In the testis, spermatozoa are devoid of projections. In the seminal vesicle, they gradually acquire the projections with tufts adhering to it. Consequently, spermatozoa in various distinct stages of projection development can be found in the seminal vesicle. We believe that these projections (1) could help transport sperm along the male and perhaps female reproductive tracts; (2) are used to anchor the spermatozoa inside the female spermatheca in order to avoid mechanical displacement by the genitalia of other males and (3) may play a role in oocyte recognition. We propose that the evolution of aflagellarity in Opiliones is related to the unique morphology of the female reproductive tract. Since eggs are fertilized on the tip of the ovipositor just prior to being laid, there is no advantage favoring sperm mobility. Additionally, female sperm receptacles are small and males that produced small spermatozoa would have a higher chance of fertilizing more eggs.     

Sperm transfer is affected by mating history in the simultaneously hermaphroditic snail Lymnaea stagnalis

Abstract. Males are predicted to strategically allocate sperm across mating partners in order to maximize their chances of paternity. This requires that males have the ability to detect aspects of their partner's mating history or the number of potential mates. We investigated whether simultaneous hermaphrodites mating in the male role strategically adjust sperm transfer depending on rearing conditions. The pond snail Lymnaea stagnalis (Basommatophora) is known to donate sperm repeatedly to different partners during a breeding season and store received sperm for >3 months. The rearing conditions of the donor as well as the recipient affect the amount of sperm transferred. Sperm donors raised in isolation transfer more sperm than those raised in groups. Furthermore, isolated sperm donors transfer less sperm to partners that were raised in groups than to those raised in isolation, i.e., virgins. These findings suggest that snails raised in isolation shift their sex allocation toward the male function and indicate that they can somehow assess the mating status of their partner.     

Parthenogenesis in Mites and Ticks (Arachnida: Acari)

SYNOPSIS. Parthenogenesis is discussed according to type (arrhenotoky,thelytoky, deuteroLoky, artificial parthenogenesis, gynogenesis)and presence in various acarine taxa. Evidence for parthenogenesisis based on rearing and/or cytological studies. Arrhenotokyoccurs in Mesostigmata, Prostigmata and Astigmata and usuallydoes not occur sporadically throughout these taxa; when presentit operates in many closely related species as the major typeof reproduction in certain genera, subfamilies or families.Thelytoky is present in Mesostigmata, Metastigmata, Prostigmata,Astigmata and Cryptostigmata, but in no case is the major modeof reproduction in higher taxa, i.e. subfamilies, families.Its sporadic occurrence may or may not be accompanied by polyploidy.Deuterotoky occurs in one acarine group (Listrophoridae) andartificial parthenogenesis is possible in ticks (Metastigmata).Gynogenesis is present in several species of haplo-diploid Mesostigmata(some Dermanyssidae, Phytoseiidae), although it is usually consideredin connection with thelytoky. Current studies of the effectsof ploidy level on morphological variation indicate greatercharacter variability in diploid females than in haploid males.In some species of ticks thelytoky is expressed differentiallyeven on an intraspecific level. One geographic strain of a speciesmay have the ability to reproduce thelytokously, but if matedproduces males and females.     

Sperm transfer and reproductive biology in species of hermaphroditic bivalves (Galeommatoidea: Montacutidae)

The histology of the reproductive organs is studied in the protandric hermaphroditic Tellimya ferruginosa. In NW Europe the species reproduces from May through August. Sperm transfer takes place when mature testis follicles are transplanted to the gills or walls of the mantle cavity in recipient hermaphroditic or female bivalves. Transplantation is accompanied by histological changes and sperm cells are released when transplants perish with age. Details are given on the reproduction in Montacuta percompressa which takes place from March through October in North Carolina, USA. All shelled bivalves are females and it is postulated that spermatogenic bodies attached to gills or other surfaces in the female's mantle cavity and previously considered to be dwarf males arise from transplanted larval gonads. The ultrastructure of the euspermatozoa and/or the anucleate paraspermatozoa is described in T. ferruginosa, T. tenella, and M. percompressa. The sperm of the first two species share a number of significant apomorphies with those of another montacutid, Brachiomya stigmatica. In the simultaneous hermaphroditic M. substriata the nucleate paraspermatozoa associate with the euspermatozoa to form spherical spermatozeugmata that are stored in the testis.     

Sperm transfer in the antCarebara vidua F. Smith (Hymenoptera: Formicidae)

Summary Sperm transfer in the myrmicine antCarebara vidua was studied by making histological sections of newly mated queens. Queens are multiply inseminated, and longitudinal sections through the sperm- filled vagina show that the bundle of sperm from each male is enclosed in a layer of accessory gland secretion terminating in a plug of secretion in the female's vaginal orifice. This spermatophore breaks down soon after mating. The huge quantity of sperm packed into the spermatheca (ca. 16 mm³) suggests that the queen lays a large number of eggs in her lifetime, confirmed by the average of 1672 ovarioles in the two ovaries.