Phylogenetic position of the mite family Myobiidae within the infraorder Eleutherengona (Acariformes) and origins of parasitism in eleutherengone mites

The position of the family Myobiidae in the infraorder Eleutherengona (Prostigmata) was analyzed with a cladistic parsimony approach for the first time. Species of the genera Anystis von Heyden (Anystidae), Pomerantzia Baker (Pomerantziidae), and Walytydeus Kuznetzov (Paratydeidae) were selected as outgroups. Among Eleutherengona, species of the following genera were selected as ingroup taxa: Hirstiella Berlese (Pterygosomatidae), Eucheyletia Baker, Cheyletiella Canestrini (Cheyletidae), Syringophilus Heller (Syringophilidae), Tarsocheylus Berlese (Tarsocheylidae), Heterocheylus Lombardini (Heterocheylidae), Pygmephorus Kramer (Pygmephoridae), Raphignathus Dugès (Raphignathidae), Neognathus Willmann (Caligonellidae), Storchia Oudemans (Stigmaeidae), and Tuckerella Womersley (Tuckerellidae). Three most parsimonious trees with similar topology were obtained. In all these trees, the family Myobiidae is situated outside of the clade joining the higher Raphignathae (Raphignathoidea and Cheyletoidea) and represents a branch within the earlier derivative Raphignathae. This result is based primarily on characters from leg setation and postembryonic development. The conclusion from this topology is that myobiid mites have developed some gnathosomal structures convergently with Cheyletoidea, including the stylet-like chelicerae and stylophore fused with the subcapitulum. The evolution of animal eleutherengone parasitism is discussed. Parasitism arisen independently in numerous phyletic lineages or superfamilies of Eleutherengona. The representatives of some phylogenetically distant eleutherengone lineages developed similar adaptations to predation and parasitism. However, in spite of some similarities in these adaptations, the evolutionary trends and pathways for switching to a parasitic mode of life are quite different in particular eleutherengone lineages.     

Phylogenetic analysis of Myobia musculi (Schranck, 1781) by using the 18S small ribosomal subunit sequence

We used high-fidelity PCR to amplify 2 overlapping regions of the ribosomal gene complex from the rodent fur mite Myobia musculi. The amplicons encompassed a large portion of the mite's ribosomal gene complex spanning 3128 nucleotides containing the entire 18S rRNA, internal transcribed spacer (ITS) 1,5.8S rRNA, ITS2, and a portion of the 5'-end of the 28S rRNA. M. musculi's 179-nucleotide 5.8S rRNA nucleotide sequence was not conserved, so this region was identified by conservation of rRNA secondary structure. Maximum likelihood and Bayesian inference phylogenetic analyses were performed by using multiple sequence alignment consisting of 1524 nucleotides of M. musculi 18S rRNA and homologous sequences from 42 prostigmatid mites and the tick Dermacentor andersoni. The phylograms produced by both methods were in agreement regarding terminal, secondary, and some tertiary phylogenetic relationships among mites. Bayesian inference discriminated most infraordinal relationships between Eleutherengona and Parasitengona mites in the suborder Anystina. Basal relationships between suborders Anystina and Eupodina historically determined by comparing differences in anatomic characteristics were less well-supported by our molecular analysis. Our results recapitulated similar 18S rRNA sequence analyses recently reported. Our study supports M. musculi as belonging to the suborder Anystina, infraorder Eleutherenona, and superfamily Cheyletoidea.     

A new mite superfamily Cloacaroidea and its position within the Prostigmata (Acariformes)

The external morphology of 2 closely related mite families, Cloacaridae and Epimyodicidae (Acari: Prostigmata), comprising highly specialized endoparasites of vertebrates, is analyzed. These mites exhibit strong regression of many structures ancestrally present in other Prostigmata as a consequence of their endoparasitic mode of life. The relationships of these 2 families with other taxa in the infraorder Eleutherengona are still not clear. Our reinterpretation of the chelicerae as unfused precludes inclusion of this lineage in the Cheyletoidea as proposed previously. A new superfamily, Cloacaroidea superfam, nov., incertae sedis, within the infraorder Eleutherengona is established for these 2 families, and their host-parasite relationships are briefly discussed. A new cloacarid species, Caminacarus dawsoni n. sp., from Graptemys pseudogeographica (Testudines: Emydidae) from the United States is also described.     

A new species of Acontiophorus Brady, 1880 (Copepoda: Siphonostomatoida) from Ushuaia,Argentina

A new species of Acontiophorus belonging to the family Asterocheridae was found in Ushuaia, Patagonia, southern Argentina. The new species is distinctive in having 9-segmented antennule, five setae on terminal endopodal segment of leg 1, seven elements on terminal segment of leg 1 and leg 4 exopods, and characteristic setation on maxillule lobes. This combination of characteristic features does not occur in any other species of the genus. The genus Acontiophorus is recorded for the first time from the South Atlantic.     

Three new species of the subgenus Neoribates (Neoribates) (Acari,Oribatida, Parakalummidae) from Nepal

Three new parakalummid mites of the subgenus Neoribates (Neoribates), N. (N.) parabulanovae sp. n., N. (N.) paramacrosacculatus sp. n. and N. (N.) pararotundus sp. n., are described from Nepalese soils. Neoribates (Neoribates) parabulanovae sp. n. is morphologically most similar to N. (N.) bulanovae Grishina, 2009, N. (N.) rotundus Aoki, 1982 and N. (N.) setiger Balogh & Mahunka, 1978, however, it differs from N. (N.) bulanovae by the body length, body and leg integument, morphology of bothridial setae, absence of aggenital setae, length of interlamellar setae and location of adanal setae ad₃; from N. (N.) rotundus by the body size, body integument, morphology of bothridial setae and length of interlamellar setae; from N. (N.) setiger by the body size, number of genital setae and absence of aggenital setae. Neoribates (Neoribates) paramacrosacculatus sp. n. is morphologically most similar to N. (N.) macrosacculatus Aoki, 1966, however, it differs from the latter by the body size, body integument, length and morphology of bothridial setae, number of genital setae, absence of lamellar setae and length of interlamellar setae. Neoribates (Neoribates) pararotundus sp. n. is morphologically most similar to N. (N.) rotundus, however, it differs from the latter by the number of notogastral setal alveoli, body integument and length of interlamellar setae.     

New Quill Mites of the Family Syringophilidae (Acari: Cheyletoidea)

Five new species and two new genera belonging to the family Syringophilidae (Acari: Cheyletoidea) are described from birds that died in the Antwerp Zoo during their quarantine: Charadriaulobia vanelli n. g., n. sp. from Vanellus chilensis (Charadriiformes: Charadriidae) in Brazil; Fritschisyringophilus lonchurae n. g., n. sp. from Lonchura punctulata (Passeriformes: Estrildidae) in India; Mironovia coturnae n. sp. from Coturnix coturnix (Galliformes: Phasianidae) in Europe; Syringophiloidus daberti n. sp. from Passerina ciris (Passeriformes: Emberizidae) in Mexico; and S. serini n. sp. from Serinus mozambicus (Passeriformes: Fringillidae) in Central Africa. Charadriaulobia n. g. differs from the closely related Aulobia Kethley, 1970, in both sexes, by the divergent epimeres I; in females, by the absence of protuberances on the hypostomal apex and by the situation of the bases of setae l4 distinctly anterior the bases of setae d4. Fritschisyringophilus n. g. differs from the closely related Syringophiloidus Kethley, 1972, in both sexes, by the presence of setae vs ' on legs II, the absence of setae dT on legs III and IV; in females, by the presence of median hypostomal protuberances and by short setae l1, l2 and l3. The relationships between the Syringophilidae and their hosts are briefly discussed. A list of all known syringophilid genera and their distribution on bird families is provided.     

Naupliar and Metanaupliar Development of Thysanoessa raschii (Malacostraca,Euphausiacea) from Godth?bsfjord,Greenland, with a Reinstatement of the Ancestral Status of the Free-Living Nauplius in Malacostracan Evolution

The presence of a characteristic crustacean larval type, the nauplius, in many crustacean taxa has often been considered one of the few uniting characters of the Crustacea. Within Malacostraca, the largest crustacean group, nauplii are only present in two taxa, Euphauciacea (krill) and Decapoda Dendrobranchiata. The presence of nauplii in these two taxa has traditionally been considered a retained primitive characteristic, but free-living nauplii have also been suggested to have reappeared a couple of times from direct developing ancestors during malacostracan evolution. Based on a re-study of Thysanoessa raschii (Euphausiacea) using preserved material collected in Greenland, we readdress this important controversy in crustacean evolution, and, in the process, redescribe the naupliar and metanaupliar development of T. raschii. In contrast to most previous studies of euphausiid development, we recognize three (not two) naupliar (= ortho-naupliar) stages (N1-N3) followed by a metanauplius (MN). While there are many morphological changes between nauplius 1 and 2 (e.g., appearance of long caudal setae), the changes between nauplius 2 and 3 are few but distinct. They involve the size of some caudal spines (largest in N3) and the setation of the antennal endopod (an extra seta in N3). A wider comparison between free-living nauplii of both Malacostraca and non-Malacostraca revealed similarities between nauplii in many taxa both at the general level (e.g., the gradual development and number of appendages) and at the more detailed level (e.g., unclear segmentation of naupliar appendages, caudal setation, presence of frontal filaments). We recognize these similarities as homologies and therefore suggest that free-living nauplii were part of the ancestral malacostracan type of development. The derived morphology (e.g., lack of feeding structures, no fully formed gut, high content of yolk) of both euphausiid and dendrobranchiate nauplii is evidently related to their non-feeding (lecithotrophic) status.     

A new species of the deep‐sea copepod genus Scutogerulus (Calanoida: Arietellidae) from the hyperbenthic waters of Okinawa,Japan

A new species of the rare arietellid genus Scutogerulus is described from deep hyperbenthic waters off Okinawa, southwestern Japan. This is the second species of the genus. Phylogenetically significant characteristics known only on the basis of the type species are confirmed by the discovery of the new congener, in particular: (1) the genital system of the female exhibits the most plesiomorphic condition of any arietellid genus; (2) the well developed setae on the endopodal segments of the maxillae and maxillipeds are modified into ‘shield‐like setae’ similar to the ‘button setae’ in another arietelloidean family, the Augaptilidae; (3) the outer distal spine on the first exopodal segment of leg 1 is absent.     

Observations on the honey bee tracheal mite <Emphasis Type="Italic">Acarapis woodi</Emphasis> (Acari: Tarsonemidae) using low-temperature scanning electron microscopy

Observations were made of cryo-preserved honey bee tracheal mites Acarapis woodi (Rennie) using scanning electron microscopy. We describe various new morphological attributes of A. woodi based on the ability of the cryo-technique to capture live mites in natural positions and observe the Low-Temperature Scanning Electron Microscopy (LT-SEM) photographs under a 3-D viewer. Most striking was the observation that each leg has the ability to independently twist its segments with the ambulacrum rotating a minimum of 180° during locomotion; this is a more sophisticated form of locomotion than has been proposed for the Acari. Adult daughter mites are known to be the dispersal instar moving from the tracheal tube to the thoracic hairs of the bee and then transferring to a new bee. We hypothesize that adult tarsal claws and setae on the legs play a role in attachment to hairs during dispersal. However, our evidence is that none of the life stases use their tarsal claws within the tracheal tubes. Larvae were observed to be freely moving within the tracheal system, their tarsal claws rendered inoperative due to an enlarged swollen pulvillar pad. The solenidia of leg I are now known to have striations and the famulus is bifurcated. The bifurcated famulus, solenidial striations, and segmentation of leg IV of females may have taxonomic implications in the family Tarsonemidae. The body and leg setae of adults appear to be used as a tactile tool to sense the amount of space within the tracheal tubes; most of the setae are oriented distally and may help the mite to measure the space or radius of the tracheal tubes. The modified caudal region of the male revealed remnants of the h ₁ and h ₂ setae and a smooth clean surface, void of a film, supporting that pharate nymphs are not attached in this species.This revised version was published online in May 2005 with a corrected cover date.     

Morphology of Parasitengona mites (Acariformes: Parasitengona) and their possible evolutionary scenario

On the basis of the analysis of morphology and biology of representatives of the Parasitengona, mostly trombiculids, trombidiids and water mites, a new attempt is made to clarify probable evolutionary scenario in this group of the higher trombidiform mites (Actinedida). It is supposed that the very old ancestral group of terrestrial arachnids, having bite-sucking mouth-parts, poorly differentiated sac-like midgut and capability to extra-oral digestion, fed predatory on different small soil arthropods at all phases of the life cycle. They were small segmented orthotrichous homeomorphic arachnids at the rank of genus or family. The favorable feeding conditions of the adult phase have led to the small eggs rich in yolk and the small larva. The latter have led in turn to the necessity of intensive feeding at the larval stage to complete the ontogenesis. Further in evolution, this group gave rise at once to two or even more large paraphyletic branches. Most of them retained feeding on arthropods with transition of larvae to much more effective parasitic feeding provided with the additional specialization of the larval stage. This branch comprise divergently radiated paraphyletic terrestrial and secondary-water water mites each having long course of evolution resulted in the recent groups of Calyptostomatoidea, Erythraeoidea, Trombidioidea and several superfamilies of water mites. Another branch of the ancestral Parasitengona has followed the way of adaptation of larvae to feeding on vertebrates, which were being attacked by the larvae in the environment of pasture. The parasitism on vertebrates has lead to several radical specializations of these mites and their significant evolutionary progress. At the same time, the similar ontogenetic dynamics, as well as synchronous reduction of particular developmental stages in all parasitengones, inevitably indicate the monophyletic origin of the whole branch of Parasitengona with Pterygosomatidae as the most probable sister group.     

A new genus and species of oribatid mites <Emphasis Type="Italic">Scarabacarus longisensillus</Emphasis> gen. et sp. n. (Acariformes,Liacaridae) from the Caucasus

A new genus and species, Scarabacarus longisensillus gen. et sp. n., of the family Liacaridae are described from the Caucasus (Azerbaijan and Abkhazia): The new genus Scarabacarus is characterized by the highly convex body, straight anterior margin of the notogaster, narrow fusiform sensilla, the lamellae situated on the lateral margins of the prodorsum, the absence of interlamellar and notogastral setae (besides one posterior pair), presence of four pairs of genital setae, and tridactylous leg tarsi.     

Evolution of hyperflexible joints in sticky prey capture appendages of harvestmen (Arachnida,Opiliones)

The rigid leg segments of arthropods are flexibly connected by joints, which usually consist of two ball-and-bowl hinges, permitting a uniaxial pivoting up to 140°. Here, we report the occurrence of hyperflexible joints (range of movements?=?160–200°) in the pedipalps (second pair of appendages) of some harvestmen (Sabaconidae and Nemastomatidae), representing some of the most flexible leg joints among arthropods. Hyperflexion is achieved by a reduction of hinges and a strong constriction of the joint region. We demonstrate that hyperflexion occurs during prey capture and is used to clamp appendages of the prey, in addition to attachment by glue secreted by specialized setae. By means of high-speed video recordings, we found that in the Sabaconidae the tibiotarsal joint of the pedipalp can flex extremely rapidly (<5 ms), limiting prey escape. This is the fastest reported predatory strike in arachnids and caused both by leverage and a click mechanism. By comparative analysis of different related taxa, we retraced joint evolution and found that hyperflexion has independently evolved in Sabaconidae and Nemastomatidae, with totally different joint kinematics. We hypothesize that (rapid) hyperflexion evolved to enhance the efficiency of the pedipalp as a means of prey capture, because in springtails detachable scales limit the action of the sticky secretion of pedipalpal setae.     

'Vertition' of integumental organs in mites revisited: a case of fluctuating asymmetry.

Vertition in mites is defined as a meristic variation for a bilateral integumental organ with a separate genetic control for each body side. A prominent hypothesis expressed by Grandjean is the role of vertition in the evolutionary trend towards a reduced number of hair-like organs (mechano- and/or chemo-receptors) known to have occurred in many mite groups. Observations on leg setae in the two-spotted spider mite Tetranychus urticae do not support this hypothesis. Meristic variation for leg setae rather conforms to the notion of fluctuating asymmetry: the difference between the number of leg setae on the right and left sides of the body had a unimodal distribution with a mean of zero. Moreover, lack of heritability for left/right absences of leg setae in an inbred laboratory strain suggests that vertition could be purely environmental. It is therefore argued that meristic variation for hair-like organs in mites is caused by random developmental accidents not corrected by homeostatic mechanisms normally resulting in a perfect bilateral symmetry.     

Host plant manipulation of natural enemies: leaf domatia protect beneficial mites from insect predators

Acarodomatia are small tufts of hair or invaginations in the leaf surface and are frequently inhabited by several taxa of non-plant-feeding mites. For many years, ecologists have hypothesized that these structures represent a mutualistic association between mites and plants where the mites benefit the plant by reducing densities of phytophagous arthropods and epiphytic microorganisms, and domatia benefit the mite by providing protection from stressful environmental conditions, other predaceous arthropods, or both. We tested these hypothesized benefits of domatia to domatia-inhabiting mites in laboratory and growth chamber experiments. In separate experiments we examined whether domatia on the wild grape, Vitis riparia, provided protection against drying humidity conditions or predaceous arthropods to two species of beneficial mite: the mycophagous species Orthotydeus lambi, and the predaceous species Amblyseius andersoni. For both taxa of beneficial mite, domatia significantly increased mite survivorship in the presence of the predatory bug, Orius insidiosus and the coccinellids Coccinella septempunctata and Harmonia varigata. There was no evidence for a protective effect of domatia with a third species of predatory arthropod, lacewing larvae Chrysoperla rufilabris. In contrast, there was no evidence for either species of beneficial mite that domatia provided any protection against low humidity. Thus in this system the primary mechanism by which domatia benefit beneficial mites is by protecting these organisms from other predatory arthropods on the leaf surface.     

External Anatomy of the Female Genital (Eighth) Limbs and the Setose Openings in Myodocopid Ostracodes (Cypridinidae)

The external anatomy of various cypridinid female genital (8th) limbs is described. Scanning electron microscopy shows that each female cypridinid genital limb has a medial depression and associated pore (= spermatophore pore); over all of which a male cements a spermatophore. The limb terminates laterally in a superficially smooth rounded knob bearing an additional small opening (= lateral pore). Dorsolateral to this genital limb there are grouped setae, previously called "brush organs", that are really paired, slit–like posterolateral openings, which are probably subdermally united to the female genital limbs. The "brush organs" of all myodocopid females also are likely to be the setose openings. The setose openings are probably homologous to the setose central lobe of male cypridinid copulatory limbs, but not homologous to the limb–like "brush organ" described for some podocopid males. We speculate that the eggs are fertilized via the spermatophore pore associated with the attached spermatophore, and that they are released from the slightly larger and more lateral setose openings into the brood chamber of the posterodorsal area of the bivalved carapace.     

How many species of Diacyclops? New taxonomic characters and species richness in a freshwater cyclopid genus (Copepoda,Cyclopoida)

The genus Diacyclops Kiefer, 1927 emend. Morton (1985) includes more than 100 species, widely distributed in all kinds of freshwater environments, and is the richest genus in the family Cyclopidae. Traditionally, Diacyclops species were defined according to differences in a few morphological characters; some characters (number of antennulary segments, segmentation pattern of swimming legs) are useful only in the separation of species groups, others (length of caudal rami and caudal setae) are highly variable even within the same population. During the study of the D. languidoides-group, minute morphological characters were used to differentiate between species (spinulation pattern and setation of antennary basis, setation and aesthetasc shape of male antennule, setation of mandibulary palp and maxilliped, shape of leg 4 basis) which allow to identify several valid species up to now concealed under the name `Diacyclops languidoides' (Lilljeborg, 1901). The coexistence of up to six congeneric species in the same sampling area: (a) supports the validity of the proposed taxonomic characters, (b) demonstrates that species richness may be highly underestimated in freshwater cyclopoid assemblages in absence of good taxonomic practice; and (c) requires an ecological explanation of species coexistence. The role of morphologically based taxonomy in order to solve general problems of distributional ecology and theoretical biology is explored.     

The idiosomal chaetotaxy of astigmatid mites

Within the Astigmata, setal homologies with the general chaetotaxic systems developed for acariform mites by F. Grandjean have never been convincingly established. This study deals with all body regions, exclusive of legs and gnathosoma, mostly utilizing as models astigmatid species from three different families. Six hypotheses which attempt to explain the ontogeny of segments, setae and cupules in the caudal bend region are compared using three-dimensional views obtained by scanning electron microscopy. The hypotheses are evaluated by in-group comparisons testing their consistency with presumed segmental boundaries, and by out-group comparisons with oribatid mites. The strongest, most parsimonious hypothesis suggests that larval astigmatid mites possess segment F, but not its setae, and the two pairs posterior to segment F are h _l and h ₂, whilst the five setae added in the protonymph are h ₃, f ₂ and ps _1–3. Six pairs of structures on the deutonymphal anal plate are identified as modified setae, and evidence supporting their homologies with setae of other instars is discussed for the first time. A comparison of the principal chaetotaxic systems used for the dorsal and anal regions of the Astigmata is presented. Application of Grandjean's setal signatures to these regions, as well as to the coxisternal and genital regions, is discussed and illustrated by examples.     

Late postnaupliar development of Monstrilla sp. (Copepoda: Monstrilloida), a protelean endoparasite of benthic polychaetes

Polychaete specimens from Hawaii were infected by the copepod Monstrilla. The development of these protelean parasites has remained unstudied for more than a century. Three postnaupliar endoparasitic stages were obtained: copepodids CIII, CIV, and CV, the latter stage found previous to and during emergence. Copepodid development, including the body and appendages (antennules, legs 1–4, caudal rami), is described and analyzed. The feeding tubes and the exiting from the host are also described. In light of the recently proposed inclusion of monstrilloids among caligiform copepods, it was found that monstrilloid copepodid development diverges from caligiforms and other copepod groups in: (1) the segmentation of the urosome at CIII, (2) the early formation of a genital complex, (3) early completion of swimming legs setation, at CIII; (4) delayed segmentation of rami of leg 3 at CIII (vs. the usual two-segmented pattern), (5) loss of one exopodal seta of leg 1 at CIV, (6) full development of leg 1 endopod vs. usually vestigial condition in caligiforms; (7) earlier segmentation of leg 4 rami, and (8) stable interstage (CIII–CV) setation pattern of legs 3 and 4. Overall, monstrilloid development appears to have unique characters and their phylogenetic relations deserve further study.     

A systematic revision of the genus Neopterygosoma Fajfer, 2019 (Acariformes: Pterygosomatidae) with the description of a new species

Systematic Parasitology - A systematic revision of the scale mites of the genus Neopterygosoma Fajfer, 2019 (Acariformes: Pterygosomatidae) formerly placed in the genus Pterygosoma Peters, 1849, is...     

New Species of Oribatid Mites of the Superfamily Oripodoidea Jacot, 1925 (Acari,Oribatida) from Trinidad and Tobago

Two new species of oribatid mites of the superfamily Oripodoidea (Acari, Oribatida) are described from Trinidad and Tobago. Scheloribates (Scheloribates) tobagoensis sp. n. (Scheloribatidae) differs fromS. (Scheloribates) papillaris Tseng, 1984 in the structure of the rostral and lamellar setae and leg solenidia, in the length of the prolamellae, the absence of aggenital setae, and in reduction of setae on tarsi III. Areozetes ryabinini sp. n. (Haplozetidae) differs from A. altimontanus Hammer, 1961 in the structure of the bothridial and notogastral setae and rostrum and in the number of claws on the tarsi.