Key to the world species of Ernogamasus Athias-Henriot, 1971 (Parasitiformes: Parasitidae).

A key to all globally known 33 species of Ernogamasus is presented.

A new species of mite in the genus Ernogamasus Athias-Henriot, 1971 (Parasitiformes: Parasitidae).

A new mite species Ernogamasus papilio sp. nov., originating from south-eastern Poland and north-eastern Slovakia, is described, as well as a supplementary description for E. aliitectatus (Athias-Henriot, 1967).

Five new species of mites in the genus Leptogamasus Trägårdh, 1936, and a new subgenus Medioperigamasus (Parasitiformes: Parasitidae).

Five new species of mites belonging to the genus Leptogamasus are described, in a new subgenus Leptogamasus (Medioperigamasus)-L. (M.) lamelligynus sp. nov., L. (M.) parcus sp. nov., L. (M.) ramosus sp. nov., L. (M.) sarcidanus sp. nov. and L. (M.) sasha sp. nov. A key to adults of all 12 globally known species of subgenera Breviperigamasus and Medioperigamasus is proposed.

Trachygamasus karuni sp. nov., a new mite species from Iran (Parasitiformes: Parasitidae).

A new mite species, Trachygamasus karuni sp. nov. is described from buffalo, sheep and horse manure at several locations north of Ahvaz city, Khuzestan, Iran. A key to 14 world species of Trachygamasus with described adults is also provided.

Key to the world species of Holoparasitus Oudemans, 1936 (Acari: Parasitiformes: Parasitidae).

Keys to the world species of the mite genus Holoparasitus (Parasitidae family) are provided separately for males and females. The genus includes 55 species, 42 of which are allocated to eight species groups. The Holoparasitus inornatus species group is newly defined.

Gonads and gametogenesis in astigmatic mites (Acariformes: Astigmata).

Astigmatans are a large group of mites living in nearly every environment and exhibiting very diverse reproductive strategies. In spite of an uniform anatomical organization of their reproductive systems, gametogenesis in each sex is highly variable, leading to gamete formation showing many peculiar features and emphasizing the distinct position of Astigmata. This review summarizes the contemporary knowledge on the structure of ovaries and testes in astigmatic mites, the peculiarities of oogenesis and spermatogenesis, as well as provides new data on several species not studied previously. New questions are discussed and approaches for future studies are proposed.

Gonads in Histiostoma mites (Acariformes: Astigmata): structure and development.

The development of male and female gonads in arrhenotokous and thelytokous species of Histiostoma was studied using transmission electron microscopy (TEM). All instars were examined: larvae, protonymphs, facultative heteromorphic deutonymphs (=hypopi), tritonymphs, and adults. In testis primordium, spermatogonia surrounding a testicular central cell (TCC) with a gradually enlarging, branched nucleus are present already at the larval stage. Spermatogonia and the TCC are connected via narrow, tubular intercellular bridges revealing that the TCC is a germline cell. Spermatocytes appear at the protonymphal stage. At the heteromorphic deutonymph stage, the testis primordium is similar to that of the protonymph, but in the tritonymph it is much larger and composed as in the adult: spermatids as well as sperm cells are present. The latter are congregated ventrally in the testis at the entrance of the deferent duct. In the larval ovary, an eccentrically located ovarian nutritive cell (ONC) is surrounded by oogonia which are connected with the ONC via tubular intercellular bridges. In later stages, the ovary grows and oocytes appear in the protonymph. Meiotic synaptonemal complexes in oocytes occur from the tritonymph stage. At about the time of the final molting, tubular intercellular bridges transform into peculiar diaphragm-crossed bridges known only in Histiostoma mites. In the adult female, growing oocytes at the end of previtellogenesis lose intercellular bridges and move ventro-laterally to the ovarian periphery towards the oviduct entrance. Vitellogenesis occurs in oviducts. Germinal cells in both the testis and ovary are embedded in a few somatic stroma cells which may be well discernible already in the larval ovary; in the testis, somatic stroma cells are evident not earlier than the end of the tritonymphal stage. The ovary has a thin wall of flat somatic cells, whereas the testis is covered by a basal lamina only. The obtained results suggest that gonads in Histiostoma and other Astigmata originate from two primordial cells only.

Localization and density of phoretic deutonymphs of the mite Uropoda orbicularis (Parasitiformes: Mesostigmata) on Aphodius beetles (Aphodiidae) affect pedicel length.

The phoretic stage of Uropodina mites is a deutonymph with developed morphological adaptations for dispersal by insects. Phoretic deutonymphs are able to produce a pedicel, a stalk-like temporary attachment structure that connects the mite with the carrier. The aim of our study was to determine whether localization and density of phoretic deutonymphs on the carrier affect pedicel length. The study was conducted on a common phoretic mite--Uropoda orbicularis (Uropodina) and two aphodiid beetles--Aphodius prodromus and Aphodius distinctus. Our results show that pedicel length is influenced by the localization of deutonymphs on the body of the carrier. The longest pedicels are produced by deutonymphs attached to the upper part of elytra, whereas deutonymphs attached to femora and trochanters of the third pair of legs and the apex of elytra construct the shortest pedicels. In general, deutonymphs attached to more exposed parts of the carrier produce longer pedicels, whereas shorter pedicels are produced when deutonymphs are fixed to non-exposed parts of the carrier. A second factor influencing pedicel length is the density of attached deutonymphs. Mean pedicel length and deutonymph densities were highly correlated: higher deutonymph density leads to the formation of longer pedicels. The cause for this correlation is discussed, and we conclude that pedicel length variability can increase successful dispersal.

Morphological diversity of pedicels in phoretic deutonymphs of Uropodina mites (Acari: Mesostigmata).

The pedicel is a structure that connects the phoretic deutonymph of Uropodina mites with its carrier and enables dispersal. The shapes, lengths and diameters of pedicels formed by Uropoda orbicularis, Trichouropoda ovalis, Uroobovella pulchella and Uroobovella nova were studied by scanning and light microscopy. Pedicels of U. orbicularis and T. ovalis have the shape of a straight stalk. In U. pulchella, the pedicel is extremely short, irregularly shaped and composed of homogeneous material. The longest pedicel is found in U. nova and it may be helically coiled in this species. The length of the pedicel is positively correlated with deutonymph body size between species, but not within species. Pedicels of U. orbicularis and U. pulchella have the largest diameter. The pedicel diameter in U. orbicularis and T. ovalis is inversely proportional to its length, but not in U. nova and U. pulchella. The constituent of pedicel stems in U. pulchella is homogeneous, whereas in U. orbicularis and T. ovalis it contains a bundle of tightly packed fibres. In U. nova coiled pedicels are comprised of two layered materials of different electron density, one of which is electron lucid and located peripherally. Hypotheses on the origin of the pedicel are proposed.

Anatomy and fine structure of pedicellar glands in phoretic deutonymphs of uropodid mites (Acari: Mesostigmata).

Phoretic deutonymphs of uropodid mites are attached to their carrier via an anal pedicel which is formed by a secretion from the pedicellar gland. Since the ultrastructure of the pedicel and pedicellar gland has never been investigated, we studied these structures in three species, Uropoda orbicularis (Müller), Uroobovella marginata (Koch), and Trichouropoda ovalis (Koch) by light (LM) and electron microscopy (TEM, SEM). In addition, the pedicel in Uroobovella nova (Oudemans) was documented in SEM. The pedicellar gland is a distinct globular structure comprised of three types of secretory cells (A-, B-, and C-types) with apical parts directed towards a junctional zone of postcolon and anal atrium. Secretory cells of the A-type are located dorsally, whereas B-type cells are central, and C-type cells are distributed ventrally or both ventrally and dorsally. Protrusions of visceral muscle cells are distributed on the external surface of the gland. The cuticle-lined anal atrium is large and located between the gland and dorso-anal muscles. The pedicel is composed of a main stalk and two extended extremities: one adhering to the anal region of the deutonymph and a second connected to the cuticle of the carrier. In each case, the anal atria were empty, whereas the pedicellar material was located outside of the mite body.

Mite species inhabiting commercial bumblebee (Bombus terrestris) nests in Polish greenhouses.

Nests of social insects are usually inhabited by various mite species that feed on pollen, other micro-arthropods or are parasitic. Well-known negative effects of worldwide economic importance are caused by mites parasitizing honeybee colonies. Lately, attention has focused on the endoparasitic mite Locustacarus buchneri that has been found in commercial bumblebees. However, little is known of other mites associated with commercial bumblebee nests. Transportation of commercial bumblebee colonies with unwanted residents may introduce foreign mite species to new localities. In this study, we assessed the prevalence and species composition of mites associated with commercial bumblebee nests and determined if the mites are foreign species for Poland and for Europe. The study was conducted on 37 commercial bumblebee nests from two companies (Dutch and Israeli), originating from two greenhouses in southern Poland, and on 20 commercial bumblebee colonies obtained directly from suppliers. The species composition and abundance of mites inhabiting commercial bumblebee nests were determined. Seven mite species from three families were found in nests after greenhouse exploitation. The predominant mite species was Tyrophagus putrescentiae (Acaridae) that was a 100-fold more numerous than representatives of the family Laelapidae (Hypoaspis marginepilosa, H. hyatti, H. bombicolens). Representatives of Parasitidae (Parasitellus fucorum, P. crinitus, P. ignotus) were least numerous. All identified mite species are common throughout Europe, foreign species were not found. Mites were not detected in nests obtained directly from suppliers. We conclude that probably bumblebee nests are invaded by local mite species during greenhouse exploitation.

Mitochondria-rich cells in the astigmatic mites, Diplaegidia columbae (Buchholz) (Analgidae) and Falculifer rostratus (Buchholz) (Falculiferidae) (Acari: Astigmata).

Mitochondria are well-characterized intracellular organelles usually concentrated in locations of high energy consumption. Light microscopic and transmission electron microscopic observations of the internal anatomy of the feather mites Diplaegidia columbae and Falculifer rostratus were conducted. In the anterior half of the bodies of the mites, we found several dozen of distinctive mitochondria-rich (MR) cells filled with abundant, large mitochondria. Mitochondria are placed individually or enclosed in small groups within an elaborated lamellar system forming a mitochondria-lamellae complex (MLC). The role of the MLC as well as the MR cells is not clear at present, but their involvement in heat generation is hypothesized and briefly discussed.

Molecular phylogeny of acariform mites (Acari, Arachnida): strong conflict between phylogenetic signal and long-branch attraction artifacts.

Acariformes (one of the two main lineages of Acari) represent an exceptionally diverse group of Arachnida. We performed first comprehensive phylogenetic analysis of Acariformes using sequence data from the nuclear small subunit rRNA gene (18S rDNA) and the mitochondrial cytochrome c oxidase subunit I (COI, amino acids). Our analyses confirm the monophyly of Acariformes and recognize two orders within Acariformes: Sarcoptiformes, consisting of Endeostigmata and Oribatida+Astigmata, and Trombidiformes. The data revealed the origin of Astigmata within Oribatida with the desmonomatan superfamily Crotonioidea as the source of astigmatan radiation and the sexual family Hermanniidae as the sister group, which generally supports previous morphological hypotheses. These results were found despite the strong conflict between long-branch attraction (LBA) artifacts and phylogenetic signal. It is likely that the conflict resulted from differences in the substitution rates among acariform lineages, especially comparing slowly evolving Oribatida with rapidly evolving Astigmata. The use of likelihood methods considered more resistant to LBA only slightly decreased the chance of falling into the LBA trap; the probability of recovering the origin of Astigmata within Desmonomata differs only by about 10% from that of having the long branched Astigmata and Trombidiformes either connected directly or shifted to deep parts of the tree due to outgroup attraction. Molecular dating using the rate-smoothing method PATHd8 shows that Acariformes originated c. 435 MYA and were probably among the earliest arthropods invading terrestrial habitats in late Silurian or the Lower Devonian, when the first vascular plants are thought to have arisen. Our analyses did not support the monophyly of Acari because we recovered clades Acariformes-Solifugae and Parasitiformes-Pseudoscorpionida. However, a formal revision of arachnid classification that would reflect these results must await future analyses.

Spermatogenesis and sperm structure in Carpoglyphus lactis (L.) (Acari: Astigmata).

Testes, spermatogenesis and spermatozoa are described in the mite Carpoglyphus lactis (L.), the first representative of the Hemisarcoptoidea superfamily studied ultrastructurally. Paired testes are located posteriorly in the idiosoma, with germaria situated dorsolaterally. The germarium consists of a compact group of spermatogonia; no testicular central cell was found. The remainder of the gonad is occupied by germ cells in different stages of spermatogenesis, distributed separately rather than in cysts, and embedded in a few large somatic cells filling the remaining space. Spermatocytes are covered by a spongy layer, a product of the Golgi apparatus. Spermatids are anucleate. Their chromatin condenses into granular and then tubular threads. As spermiogenesis progresses, the spongy layer assembles at a single site and forms a structure termed the spongy body; mitochondria become electron dense, elongate and gather forming a bundle; a narrow ER cistern, promptly transforming into a dense lamella, appears between the mitochondria and chromatin. Mature spermatozoa are small, highly electron-dense cells interdigitating with others via superficial protrusions. They possess chromatin threads, electron-dense lamella and mitochondria, but do not have an acrosome. Our results support the monophyly of Astigmata, but do not explain the phylogenetic affinities of Hemisarcoptoidea to other superfamilies of astigmatic mites.

Sperm structure and phylogeny of Astigmata.

The Astigmata, a large and variable group, is still a subject of taxonomic dispute. Particularly, their origin from ancestors of the lower oribatid mites (e.g., Malaconothroidea) seems well documented by many lines of evidence. The structure of spermatozoa has been successfully applied to phylogenetic investigations in many animal groups. The aim of our study was to provide new data on spermatozoon structure in Astigmata and to consider its appropriateness in phylogenetic studies. The study reveals information on spermatozoa in 17 species of Astigmata (11 species studied for the first time) extending our knowledge to 18 species (one species known only from the literature) representing 12 families and 7 superfamilies. Spermatozoa have the same basic structure in all species: cells are multiform and the chromatin forms thin threads embedded directly in the cytoplasm; the acrosome is absent. The cytoplasm in most species contains electron-dense lamellae, varying in both number and arrangement within the cell. In Sarcoptoidea, electron-dense tubules in contact with lamellae margins were also observed in Psoroptidae (Psoroptes equi), whereas in two representatives of Sarcoptidae (Notoedres cati and Sarcoptes scabiei), only electron-dense tubules were found. In two species, Canestrinia sellnicki (Canestrinioidea: Canestriniidae) and Scutulanyssus obscurus (Analgoidea: Pteronyssidae), neither lamellae nor tubules were present. The mitochondria in a spermatozoon are usually gathered at the cell periphery and their structure is usually modified to form so-called mitochondrial derivatives. The chromatin threads are an autapomorphy strongly supporting the monophyly of Astigmata. As spermatozoa vary considerably between species in Astigmata, we deduce that sperm structure may be useful for phylogenetic analyses within the group. Several conclusions concerning the affinities within Astigmata are presented. Spermatology seems to be unhelpful, however, in questions on the origin of Astigmata (particularly for Astigmata-Oribatida relationships), since their sperm do not possess synapomorphies with sperm of the remaining groups of Acariformes, i.e., Endeostigmata, Prostigmata, and Oribatida.

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.

Fine structure of the genital system in the females of Pergamasus mites (Acari: Gamasida: Pergamasidae).

The female reproductive system in Pergamasus mites consists of an unpaired vagina, vaginal duct, uterus, and ovary. Additionally, there are paired vaginal glands, as well as unpaired ventral and paired lateromedial glandular complexes. The vagina and vaginal duct are cuticle-lined. In the dorsal wall of the vagina, this lining forms the endogynium which possesses a "sac" and two conspicuous "spherules" and is armed with "stipula" and other cuticular protrusions. The endogynium functions as a spermatheca, being a storing site for the spermatophore. The spherule procuticle is perforated by microvilli of underlying cells that are structurally very unusual. The lining of the vaginal duct forms numerous cuticular fibers directed toward the vagina. There is an external layer of muscles, supposedly functioning as a sphincter. The uterus is an organ in which the fertilized egg is stored for some time and starts embryonic development. Its wall is composed of glandular epithelial cells. The ovary consists of inner and outer parts. The former part is formed by a nutritive syncytium, whereas the latter contains growing oocytes. Two groups of glands connect with the genital tract. Paired vaginal glands are composed of glandular and secretion-storing parts and open into the vagina. Paired lateromedial and unpaired ventral glandular complexes empty into the genital tract between the vaginal duct and uterus. The structure of the female genital system is discussed in terms of its function and phylogeny. J. Morphol. 240:195-223, 1999. © 1999 Wiley-Liss, Inc.