Unusual way of feeding by the deutonymph of Neottialges evansi (Actinotrichida, Astigmata, Hypoderatidae), a subcutaneous parasite of cormorants, revealed by fine structural analyses.

The parasitic deutonymphs of hypoderatid mites live within the subcutaneous layer of their avian hosts, where they become greatly engorged despite not having functional mouthparts. The method by which they take up nutrients has been mysterious up to now. Here, we report on the morphology of hypoderatid deutonymphs using scanning and transmission electron microscopy and describe structures that may resolve the mystery. The deutonymph of Neottialges evansi (Hypoderatidae) from the cormorant Phalacrocorax carbo is a simply organized stage lacking both mouthparts and a functional foregut. The structure of midgut and hindgut indicate that they are not capable of processing food. The midgut consists of highly branching flat cells and rarely shows a lumen. Almost the entire space between integument, gut remnants and other organs (synganglion, developing gonads) is filled by huge cells containing protein and glycogen granules and numerous lipid inclusions. The anal opening is minute. The structure of the cuticle and epidermis suggests that nutrients are not absorbed through the general integument. Thus the two main existing hypotheses about feeding modes in hypoderatid deutonymphs, anal vs. integumentary food absorbtion, are not supported. We suggest instead that two pairs of genital papillae showing peculiar microanatomical features are actively involved in movement of liquid materials between host and mite and most probably are the nutrient-intake organs. J. Morphol. 277:1368-1389, 2016. © 2016 Wiley Periodicals, Inc.

Ultrastructural characterization of Acarispora falculifera n.gen., n.sp., a new microsporidium (Opisthokonta: Chytridiopsida) from the feather mite Falculifer rostratus (Astigmata: Pterolichoidea).

Only about 20 species of microsporidia have been described from mites. All except one species produce typical spores with a long polar filament and a polaroplast. This paper is the first study of an atypical microsporidium infection in a feather mite (Falculifer rostratus). The infection of the pigeon feather mite is restricted to the colon epithelium where it leads to hypertrophy of the concerned cells. During sporogony, a multinucleate plasmodial aggregate is formed within a sporont (endogenous sporogony resulting in a polysporophorous vesicle). The cisterns delimiting the single sporoblasts later form the spore walls. Sporogonial stages are in direct contact to the host cell cytoplasm. Merogonial stages were not present. Spores are tiny (3.6 µm × 2.6 µm), broad oval in form and monokaryotic. The spore wall of mature spores consists of a three-layered endospore and a thin, electron-dense, wavy exospore. The polar filament is anisofilar and completely coiled in 3-4 turns. In cross-sections, it has a star-like appearance because the electron-dense core forms rounded compartments of lucent material at its surface. In superficial sections, this results in a honeycomb-like pattern. A polaroplast is missing. The polar filament arises subapically at a polar sac that lacks an internal anchoring disk. These atypical spore structures clearly classify the species from the feather mite as a member of the order Chytridiopsida. It could not be clearly affiliated to one of the known genera, so we created a new genus, Acarispora, with the species A. falculifera.

Fine structure of the male genital system of the predatory mite Rhagidia halophila (Rhagidiidae, Prostigmata, Actinotrichida).

The male genital system of the actinotrichid mite Rhagidia halophila is described and compared with other mites and arachnids. The large testes are composed of germinal and glandular parts and produce numerous small sperm cells. The glandular parts are connected via a testicular bridge. Spermiogenesis occurs in cysts containing spermatids in equal stages of development. Cysts of spermatids are embedded in huge somatic cells. The nuclei of the spermatids loose their envelope. Mature sperm cells are simple exhibiting a ring-shaped chromatin body and lacking an acrosomal complex. They are most similar to the sperm cells of the related mite Linopodes motatorius. The spermatopositor contains the ejaculatory duct divided into a dorsal channel and a ventral channel that are connected via a narrow passage. At its distal end, the spermatopositor is divided into three eugenital lips. The function of the spermatopositor during deposition of the peculiar thread-like spermatophores is discussed. Details of the sensilla of the spermatopositor and the progenital lips are reported. The genital papillae located on the inner side of the progenital lips exhibit characteristics of cells performing transport of ions and/or water. The results confirm the overall similarity of actinotrichid genital systems, which is profoundly different from that of anactinotrichid mites. With reference to other Arachnida it is corroborated that testes and sperm structure of Actinotrichida are most similar to that of Solifugae. However, synapomorphies between sperm cells of Rhagidia and Solifugae that could suggest a closer relationship between these two taxa as was suggested in earlier studies were not recognizable. On the contrary, the sperm cells of Rh. halophila being devoid of an acrosomal complex appeared to be more apomorphic than those of many other actinotrichid mites as well as Solifugae.

Fine structure of the urnulae of Balaustium mites (Actinotrichida: Erythraeidae) representing peculiar defense organs.

The urnulae, until now the enigmatic paired dorsal protrusions on idiosoma dorsum in active postlarval forms of Balaustium mites, were studied using electron microscopy. They consist of walls made of unmodified integument, which form a cylinder covered by a roof of thin cuticle. At the posterior border of the urnula, the roof has a crescent slit. On its inner surface, a rather large muscle inserts with several tendons. The roof forms a flap under which the modified columnar epidermal cells containing numerous lipid inclusions are located. These lipids are probably secreted through pore canals of the overlying cuticle. Materials mainly originating from an extensive vesicular tissue situated underneath the columnar cells of the urnula and under the adjacent unmodified epidermis are extruded through the mentioned slit. Our results support previous studies that have suggested a function of the urnulae as defensive organs. Our study further suggests that the agent that provides the repellent effect comes mainly from the vesicular tissue, whereas the columnar cells with their lipid secretions are likely to restore the external secretion layer of the epicuticle after its destruction during the repellent release. Further structural and functional details are discussed and compared with other putative defensive secretory organs.

Ultrastructure investigation of the secondary insemination system of the gamasid mite Hattena cometis domrow (Acari: Anactinotrichida: Ameroseiidae).

Many gamasid mites, mainly of the taxon Dermanyssina, possess a secondarily evolved insemination system that is generally described as occurring in two types, the laelapid and the phytoseiid-type, which are structurally considerably different. Considering that Dermanyssina represent the most recent and most diverse group of gamasid mites, it was expected that a greater diversity of insemination system than reflected by the two types could be present and could give an idea of its evolution within the taxon. Here, the authors present a description of the fine-structure of the female secondary insemination system in the dermanyssine mite Hattena cometis. The system consists of a pair of sperm induction pores (solenostomes) and short sperm access ducts (tubules) which end in a syncytium. The syncytial strands of both sides meet medially under the ovary s.str., where they form a spherical syncytial spermatheca. Mature sperm cells of a modified ribbon type were seen in the syncytial parts of the system. The insemination system of Hattena cometis is regarded as a modification of the laelapid type. However, it is much simpler than that of Varroa destructor, the only other gamasid mite with the laelapid type studied ultrastructurally until now, and shows also some structural differences (e.g., no presence of an unpaired sperm duct). Hence, the present study suggests that some intermediate types might be revealed in future ultrastructure studies representing steps in the evolution of the insemination system in the Dermanyssina.

Ultrastructure of the male genital tract, spermatogenesis and spermatozoa of Hattena cometis Domrow (Acari: Gamasida: Ameroseiidae).

The ameroseiid mite Hattena cometis has a male genital system that consists of an unpaired, u-shaped testis and paired deferent ducts leading into an unpaired accessory genital gland and ejaculatory duct. The genital opening is located anteriorly immediately in front of the sternal shield. Spermatogenesis is simple, probably due to the haploid nature of the male. Eight stages of spermatogenesis could be roughly distinguished. Mature spermatozoa as found in the deferent duct lumen are peculiar in having a bisected nucleus and numerous peripheral flat chambers, which were formed from indentations of the plasmalemma. In inseminated females, spermatozoa were observed in the syncytial tissue of the sperm access system and in the somatic cells of the ovary. These spermatozoa have achieved a new structure, i.e., an electron-dense plate dividing the cell into two unequal halves. The dense plate has an intricate substructure. Its function is unknown. These sperm cells are considered to represent capacitated spermatozoa. The peripheral chambers are reduced in number inside the female. Similar sperm cells, containing a dense plate, were seen in vacuoles within the epithelium of the deferent duct of one male. These cells are evidently under destruction, but before being completely dissolved had undergone a development leading beyond that of the mature sperm cells found in the deferent duct. Apparently, entering the cell of the deferent duct epithelium or the syncytium tissue triggers the production of the dense plate (or the capacitation process). Our observations are compared with results obtained from other anactinotrichid Acari, mainly Gamasida, and confirm and complete the interpretation of the correlated evolution of components of gamasid reproductive systems.

Ultrastructure of the male chelicerae of Hattena cometis Domrow (Acari: Gamasida: Ameroseiidae) functioning as gonopods.

The ultrastructure and functional adaptations of male chelicerae in Hattena cometis Domrow are discussed. In particular, as in many other gamasid mites, males of Hattena use the chelicerae, modified as gonopods, to transfer the sperm into the female. For such purpose, a slender process extending from the movable digit, the spermatodactyl, is mainly involved. The spermatodactyl is provided with a sperm transfer duct; in H.cometis, the dorsal surface bent and fused with the ventral surface forms this duct so that the spermatodactyl appears as a cuticular process, connected somehow with the movable digit, and folded on itself to delimit the sperm transfer tube. The organization and ultrastructure of the spermatodactyl are discussed and compared with other gamasid mites studied so far.

Fine structure of the gnathosoma of archegozetes longisetosus [corrected] aoki (acari: oribatida, trhypochthoniidae).

Oribatida are one of the main groups of Acari comprising mostly important decomposers in soils. Most species are particle feeders, an exceptional mode of nutrition in Arachnida. Hence, their feeding organs, the gnathosoma, are of special functional interest. We studied nearly all components using scanning and transmission electron microscopies as well as reconstructions based on synchrotron X-ray microtomography from the model oribatid Archegozetes longisetosus. Besides cuticular structures, we describe the full set of muscles and confirm the presence of a trochanter remnant at the base of the chelicera. Setae on the prodorsum and the anterior and posterior infracapitular setae are mechanoreceptors innervated by two dendrites ending with tubular bodies. Dendrites of adoral setae, anterior setae of the chelicerae, and the supracoxal setae show neither obvious tubular bodies nor wall or terminal pores. Thus their function remains obscure. For the first time, a muscular proprioreceptor has been found in Arachnida. It likely monitors the actions of muscles moving the movable digit of the chelicera. Glandular structures within and associated with the gnathosoma are described. Dermal glands represented by secretory porose areas are found within the infracapitulum. More complex associated glands comprise the podocephalic glands and the infracapitular glands, the ducts of which were traced completely for the first time. The components described are mostly fundamental for the gnathosoma of Actinotrichida (Acariformes), one of the two lineages of Acari, to which Oribatida belong. The gnathosoma is generally considered the most relevant putative synapomorphy of Acari. Since the monophyly of Acari has become more and more questionable during the last decades, a thorough reinvestigation of this body part is necessary for a comprehensive understanding of acarine and even arachnid phylogeny and evolution. This article provides a starting point of such a re-evaluation of the gnathosoma.

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.

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.

Effects of starvation on reproduction of the predacious mite Neoseiulus californicus (Acari: Phytoseiidae).

Effects of starvation on gravid females of Neoseiulus californicus were investigated at 20 degrees C and 85% RH. When females that had been reared with abundant prey were swapped, just after laying their first egg, to conditions without any prey and water, they laid 1.8 eggs and survived for 4.3 days. In the body of well-fed females, an egg with eggshell and/or two oocytes were observed in the ventral and dorsal regions, respectively. The larger oocyte had two roundish nuclei and abundant yolk granules, and was enveloped with a vitelline membrane. These two nuclei were not fused but were just close to each other. The smaller oocyte had a nucleus, but had not yet formed yolk granules and vitelline membrane. Females after 12 h starvation had an egg in the ventral region and an oocyte in the dorsal region of the body. After more than 24 h starvation females maintained an oocyte in the dorsal region of the body, but had no egg in the ventral region. The oocyte was filled with abundant yolk granules and contained two irregular nuclei when females were starved for 24 h, but when starved for more than 36 h it contained one irregular nucleus. These findings suggest that (1) gravid females maintained an oocyte in the dorsal region after laying two eggs during starvation, (2) the oocyte was not absorbed during starvation, (3) the oocyte advanced vitellogenesis and the fusion of two nuclei, and (4) the vitellogenic oocyte was not enveloped with an eggshell and had not started embryogenesis.

Ultrastructure of the digestive tract in Acarus siro (Acari: Acaridida).

The gut of the mite Acarus siro is characterized on the ultrastructural level. It consists of the foregut (pharynx, esophagus), midgut (ventriculus, caeca, colon, intercolon, postcolonic diverticula, postcolon), and hindgut (anal atrium). The gut wall is formed by a single-layered epithelium; only regenerative cells are located basally and these have no contact with the lumen. Eight cell types form the whole gut: (i) simple epithelial cells forming fore- and hindgut; (ii) cells that probably produce the peritrophic membrane; (iii) regenerative cells occurring in the ventriculus, caeca, colon, and intercolon; (iv) spherite cells and (v) digestive cells forming the ventriculus and caeca; (vi) colonic cells and (vii) intercolonic cells; and (viii) cells forming the walls of postcolonic diverticula and postcolon. Spherite and digestive cells change in structure during secretory cycles, which are described and discussed. The cycle of spherite, colonic, and intercolonic cells is terminated by apoptosis. Ingested food is packed into a food bolus surrounded by a single homogeneous peritrophic membrane formed by addition of lamellae that subsequently fuse together. The postcolonic diverticula serve as a shelter for filamentous bacteria, which also are abundant in the intercolon.

In situ observation of the Cardinium symbionts of Brevipalpus (Acari: Tenuipalpidae) by electron microscopy.

Brevipalpus (Acari: Tenuipalpidae) mites are important pests on a variety of host plant species. The mites damage their hosts directly by feeding and some species also serve as vectors of plant viruses. Among more than 200 described Brevipalpus species, three are recognized as vectors of plant viruses: B. phoenicis, B. californicus and B. obovatus. These species occur worldwide in subtropical and tropical regions. Brevipalpus mites reproduce mostly by thelytokous parthenogenesis and this condition was attributed to a bacterial endosymbiont, recently characterized as a member of the genus Cardinium. The same symbiont infects many other arthropods and is capable of manipulating their host reproduction in various ways. Generally the presence of Cardinium is determined by molecular, PCR based, techniques. In the current work we present visual evidence for the presence of these bacteria by transmission electron microscopy as a complement of previous detection by PCR. Cardinium is easily identified by the presence of a unique array of microtubule-like structures (ML) in the cell. Symbionts have been observed in several organs and eggs from different populations of all three Brevipalpus species known as vector of plant viruses. Cardinium cells were always immersed directly within the cytoplasm of infected cells. Bacteria were observed in all females of all instars, but were absent from all males examined. Females from some Brevipalpus populations were observed to be uninfected by Cardinium. This observation confirmed previous PCR-based results that these populations were aposymbiotic. The observed distribution of the bacteria suggests that these bacteria could have other functions in the mite biology beside feminization.

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.

Ultrastructural observations of spermatozoa of several tetragnathid spiders with phylogenetic implications (Araneae, Tetragnathidae).

The present study reports on the ultrastructure of spermatozoa and spermatogenesis of 12 tetragnathid spiders (10 Tetragnatha species [T. boydi, T. dearmata, T. extensa, T. montana, T. nigrita, T. obtusa, T. pinicola, T. reimoseri, T. shoshone, T. striata]; Pachygnatha listeri and Metellina segmentata). All species develop typical cleistospermia with a coiled nucleus in the center and a coiled axoneme in the periphery of the cell. Remarkable differences in the sperm ultrastructure of the investigated species comprise the shape of the main sperm cell components (nucleus, acrosomal complex, implantation fossa, and centriolar complex). Within the observed Tetragnatha species, three types of sperms were characterized: T. montana-type, T. boydi-type, and T. striata-type. The highly derivative T. montana-type is characterized by the following remarkable features: an extremely elongated nucleus, shaped like a corkscrew and twisted around the axoneme (before coiling); a deep implantation fossa; a corkscrew-shaped acrosomal vacuole; after the coiling process, the nucleus is coiled five to six times in the center of the spermatozoon and the axoneme is coiled five to six times peripheral to the nucleus. The T. boydi-type hardly differs from the T. montana-type, but is remarkable due to the triangular-shaped nucleus (in cross section). The T. striata-type differs especially by a peculiar acrosomal vacuole with a long, slightly curved process and a short appendix, as well as a nucleus that describes only three loose coils around the axoneme (before coiling). The spermatozoa of Pachygnatha listeri and especially Metellina segmentata differ strikingly from the described Tetragnatha-types and are similar to more primitive araneomorph spermatozoa, such as Hypochilus pococki. The described Tetragnatha-types completely correspond with Okuma's (1988a,b, J Fac Agr Kyushu U 32:165-181, 32:183-213) classification of Tetragnatha species. Furthermore, our results suggest an early derivative systematic position of Pachygnatha within Tetragnathinae and the position of Metellina within the Tetragnathidae.

Fine structure and functional morphology of the mouthparts of a male Veigaia sp. (Gamasida: Veigaiidae) with remarks on the spermatodactyl and related sensory structures.

Mites of the genus Veigaia are common gamasid inhabitants of forest litter. They engage in the peculiar reproductive strategy of podospermy which, along with other morphological and behavioral adaptations, involves modification of the chelicerae of the relatively rare males into gonopods. Each movable digit is provided with an appendage (spermatodactyl) that is involved in sperm transfer. We describe the gross anatomy, fine structure, and functional morphology of the mouthparts of a male Veigaia species and give ultrastructural details for the corniculi, laciniae, preoral cavity, labrum, pharynx, and movable and fixed digits. The fine structure of the spermatodactyl is illustrated here for the first time in detail. A semischematic reconstruction of the gnathosoma and spermatodactyl is provided. The spermatodactyl is totally fused with the movable digit and a sperm transfer duct runs along its entire length. This duct starts at the adaxial base of the movable digit, continues inside the digit into the tube of the spermatodactyl, and finally opens at the distal abaxial surface of the spermatodactyl. Several sensory structures associated with the spermatodactyl probably provide the male with mechanical and chemical clues.

Lead-induced hsp70 and hsp60 pattern transformation and leg malformation during postembryonic development in the oribatid mite, Archegozetes longisetosus Aoki.

The study aimed at analysing the impact of high lead concentrations on the morphological integrity and the stress protein hsp70 and hsp60 levels during postembryonic development of the oribatid mite, Archegozetes longisetosus. Independent of the treatment, the recorded hsp70 levels were far higher than the hsp60 levels in all investigated stages. There was a tendency towards lower hsp70 and hsp60 levels with proceeding development (deutonymph>tritonymph>adult) in untreated animals. Both the hsp70 and hsp60 levels in all investigated quiescent stages prior to moult were higher than in the corresponding active stages independent from lead exposure. Continuous lead treatment from the larval stage onwards caused malformation of the 4th pair of legs and, in parallel, a shift to elevated hsp70 (but not hsp60) levels in all subsequent stages, compared to controls. Neither effects occurred when continuous lead treatment started later in development. In this case, elevated hsp60 levels could particularly be found in those stages respectively following the initially exposed stage. The hsp70 response became obvious even quicker in tritonymphs and adults, where hsp70 level peaks could be observed right in those stages the lead exposure had started in.

Female genital system of the folding-trapdoor spider Antrodiaetus unicolor (Hentz, 1842) (Antrodiaetidae, Araneae): ultrastructural study of form and function with notes on reproductive biology of spiders.

The genitalia of the female folding-trapdoor spider Antrodiaetus unicolor are characterized by two pairs of spermathecae that are arranged in a single row and connected to the roof of the bursa copulatrix. Each single spermatheca is divided into three main parts: stalk, bowl, and bulb, which are surrounded by the spermathecal gland. The epithelium of the spermathecal gland is underlain by a muscle meshwork and consists of different types of cells partly belonging to glandular cell units (Class 3 gland cells) that extend into pores in the cuticle of the stalk and bowl. Interestingly, the bulb lacks glandular pores and is characterized by a weakly sclerotized cuticle. This peculiarly structured bulb probably plays an important role in the discharge of the sperm mass. It is suggested that by contraction of the muscle layer the sperm mass may be squeezed out, when the bulb invaginates and expands into the spermathecal lumen, pushing the sperm to the uterus lumen. Each glandular unit consists of usually one or two central secretory cells that are for the most part surrounded by a connecting cell that again is surrounded by a canal cell. The canal cell, finally, is separated from the other epithelial cells (intercalary cells) located between the glandular units by several thin sheath cells that form the outer enveloping layer of the unit. The secretions are released through a cuticular duct that originates proximally between the apical part of the connecting cell and the apical microvilli of the secretory cells and runs into a pore of the spermathecal cuticle. The glandular products of the Class 3 gland cells likely contribute to the conditions allowing long-term storage of the spermatozoa in this species. Details regarding the ovary, the uterus internus, and the uterus externus are reported. Most of the secretion that composes the chorion of the egg is produced in the ovary. Glandular cell units observed in the uterus externus differ structurally from those in the spermathecae and likely play a different role. Finally, we briefly discuss our results on the female genitalia of A. unicolor in the light of knowledge about the reproductive biology of spiders.

Double spermatogenesis in Chelicerata.

Sperm dimorphism is a rare phenomenon in Chelicerata. Until now, it was known only from three species of the opilionid genus Siro (Sironidae, Cyphophthalmi). Fertilizing (eusperm) and nonfertilizing spermatozoa (parasperm) develop in the same cyst and are thus sister cells. The fine structure of the spermatozoa of two species has been examined and is compared here. In contrast to Siro rubens, S. duricorius spermatozoa lack an acrosomal complex. Both sperm types produce a transitional process, a more or less modified flagellum, which is later retracted. Hence, the spermatozoa are aflagellate. Eusperm and parasperm of all three species form highly ordered sperm balls that are stored in the deferent duct. Reviewing and adding new results about the sperm dimorphism in this arachnid taxon provides the basis for some considerations of another enigmatic morphological character found in Uropygi and Amblypygi, i.e., the tubular accessory genital glands that show holocrine extrusion. These glands are suggested to represent modified, infertile derivatives of the testis anlage. Their secretion is produced in a way reminiscent of a strongly degenerated spermatogenesis. Consequently, these products may be regarded as strongly degenerated germ cells representing a line of germ cell development, which has been separated very early in spermatogenesis from the usual line leading to fertilizing sperm cells. This further, although less evident, case of probable dichotomous germ cell development is discussed with respect to the controversial phylogenetic-systematic relationships between Uropygi (Thelyphonida and Schizomida), Amblypygi, and Araneae.

Fine structure and probable function of ring organs in the mite Histiostoma feroniarum (Acari: Actinotrichida: Acaridida: Histiostomatidae).

Histiostoma feroniarum, like other histiostomatid mites, possesses peculiar ring organs that are visible under the light microscope as ventrally located, characteristic rings of sclerotized cuticle. The ring organ is composed of three elements: a disc of modified cuticle, ring organ cells located underneath the disc, and an "empty" chamber frequently visible between the cuticular disc and the cells. The cuticle of the disc is not perforated and differs from the surrounding unmodified cuticle as revealed by special staining developed for light microscopy and by electron microscopy. The ring organ cells show a polarity, with a practically smooth apical surface and an extremely folded basal membrane. The basal invaginations reach the apical cell portion, where they form tubular canaliculi distributed beneath the apical cell membrane. The cytoplasm contains many mitochondria, which are usually in contact with the cell membrane invaginations. Structurally, the ring organ cells closely resemble the transport cells described in osmoregulatory organs both in water-inhabiting and terrestrial arthropods. Thus, our results support earlier suggestions of an osmoregulatory function performed by sclerotized rings (=ring organs), as an adaptation to aqueous environments. A possible homology with similar organs of other mites is discussed.