Molecular phylogeny and systematics of Dipodoidea: a test of morphology‐based hypotheses

Lebedev, V.S., Bannikova, A.A., Pagès, M., Pisano, J., Michaux, J.R. & Shenbrot, G.I. (2012). Molecular phylogeny and systematics of Dipodoidea: a test of morphology‐based hypotheses. —Zoologica Scripta, 42, 231–249. The superfamily Dipodoidea (Rodentia, Myomorpha) in its current interpretation contains a single family subdivided into six subfamilies. Four of them include morphologically specialized bipedal arid‐dwelling jerboas (Dipodinae – three‐toed jerboas, Allactaginae – five‐toed jerboas, Cardiocraniinae – pygmy jerboas and Euchoreutinae – long‐eared jerboas), the other two are represented by more generalized quadrupedal taxa (Zapodinae – jumping mice and Sminthinae – birch mice). Despite considerable effort from morphologists, the taxonomy as well as the phylogeny of the Dipodoidea remains controversial. Strikingly, molecular approach has never been envisaged to investigate these questions. In this study, the phylogenetic relationships among the main dipodoid lineages were reconstructed for the first time using DNA sequence data from four nuclear genes (IRBP, GHR, BRCA1, RAG1). No evidence of conflict among genes was revealed. The same robustly supported tree topology was inferred from the concatenated alignment whatever the phylogenetic methods used (maximum parsimony, maximum‐likelihood and Bayesian phylogenetic methods). Sminthinae branches basally within the dipodoids followed by Zapodinae. Monophyletic Cardiocraniinae is sister to all other jerboas. Within the latter, the monophyly of both Dipodinae and Allactaginae is highly supported. The relationships between Dipodinae, Allactaginae and Euchoreutinae should be regarded as unresolved trichotomy. Morphological hypotheses were confronted to findings based on the presented molecular data. As a result, previously proposed sister group relationships between Euchoreutes and Sicista, Paradipus and Cardiocraniinae as well as the monophyly of Cardiocaniinae + Dipodinae were rejected. However, the latter association is consistently supported by most morphological analyses. The basis of the obvious conflict between genes and morphology remains unclear. Suggested modifications to the taxonomy of Dipodoidea imply recognition of three families: Sminthidae, Zapodidae and Dipodidae, the latter including Cardiocraniinae, Euchoreutinae, Allactaginae and Dipodinae as subfamilies.     

Characterization and phylogenetic analysis of the complete mitogenome of Allactaga sibirica (Rodentia: Dipodidae)

Allactaga sibirica (Dipodidae) is widely distributed in the northwestern arid regions of China. The complete mitochondrial genome (mitogenome) of A. sibirica was 16,685 bp in length; included 13 protein-coding genes, 2 ribosome genes, 22 tRNA genes, and one control region; and had a structure that was typical of vertebrates. The base composition and codon usage of the mitogenome are described, and the structure of the non-coding sequence in the A. sibirica is reported for the first time. The putative origin of replication for the light strand of A. sibirica was approximately 45 bp long, and was highly conserved in the stem-loop and adjacent sequences. Phylogenetic analyses showed high resolution in each of the main divergent clades within Dipodoidea using mitogenomes data. The results indicated that the Zapodidae group was a representative of very basal taxon in Dipodoidea, and shared a common ancestor with Dipodidae species. Within Dipodidae clade, Allactaginae species was at basal position, and this result was in line with previous molecular systematic and morphological studies. Furthermore, Euchoreutes naso and A. sibirica had a close relationship could implicate a sister-group relationship between Euchoreutinae and Allactaginae. Meanwhile, this work also provided a set of useful data on phylogeny and molecular evolution in Dipodidae species.     

Limb myology and phylogenetic relationships in the superfamily Dipodoidea (birch mice, jumping mice, and jerboas)

Limb muscles were dissected in seven genera, representing all six superfamilies, of dipodoid rodents and myoloic characters were used to construct a phylogenetic hpothesis of relationships within this cfade. Mologic differences among genera suported tie monophyly of the superfamily Dipodoidea reLtive to the outrou taxon and reveaEd thatSicista is the sister group to all other zapodid and dipodid enera. Tkis picement of Sicista differs markedly from its position in previous classifications where it has been regarded merely as a primitive zapodid genus. The phlograrn based on rnyologic characters also indicated that Cardiocranius is not a rimitive dipodid genus; it is the sister group to the subfamily Dipodinae. Although myologic differences among taxa were not sufficient to warrant the continued separation of zaodids and dipodids into two families, a new classification that places Sicista in its own family, ficistidae, and places the remaining zaodids and dipodids in the family Dipodidae, is proposed. Differences in karyology, genitaP morholoy, and postcranial osteological characters among dipodoid rodents are discussed in light or this pjylogeny.     

Tracing the Origin and Diversification of Dipodoidea (Order: Rodentia): Evidence from Fossil Record and Molecular Phylogeny

Dipodoidea are a diverse rodent group whose earliest known record is from the Middle Eocene. The evolution and diversification of this superfamily have been documented by fossils and comparative morphology, but have not yet been studied from the perspective of molecular phylogeny. This study is the first attempt to reconstruct an extensive phylogeny of Dipodidae and estimate divergence times based on a nuclear gene coding for interphotoreceptor retinoid-binding protein. We found that there is a wide measure of agreement with the fossil record. Each of the three ecological groups of the extant Dipodoidea (sicistines, zapodines, and jerboas) has its distinctive distribution; the distribution patterns have been shaped by the dispersal events. The key events of paleogeographic distribution coincided with major paleoenvironmental events in the Cenozoic. The first important diversification phase occurred during the period from the Oligocene to Early Miocene, when global climate underwent major changes beginning with the Eocene/Oligocene boundary. The second adaptive radiation occurred within jerboas and was associated with the expansion of open habitat starting with the late Middle Miocene. The diversification of jerboas can be correlated with habitat changes in response to global and regional climatic events.     

A new recent genus and species of three‐toed jerboas (Rodentia: Dipodinae) from China: A living fossil?

The new recent genus and species of three‐toed jerboas (Rodentia: Dipodinae), from southern Ningxia, China, is described. This form demonstrated a unique mixture of external, cranial, and dental characters that individually are typical for one or another of all known genera of Dipodinae. Based on morphological characters, it is recovered as the part of Dipodinae tree, distinct from all other members due to its unique combination of morphological characters, and appears to be a nearly ideal living ancestor of all other dipodines. In contrast to morphology, the molecular data indicate a relatively young age for this lineage and consistently place it as the sister group to Stylodipus. The results of the molecular clock analysis suggest that the separation of the two lineages dates back to the Early Pliocene or the Pliocene/Miocene boundary. The estimated geographic range of the new form seems extremely small. The conservational status of the new species remains to be determined; however, the available information suggests that it requires protection.     

中国蚌螨属支序分类分析(蜱螨亚纲:蚌螨科)

选择35个特征对我国蚌螨属15种的支序分析结果表明:蚌螨P-Ⅳ(胫节)端腹部的指状突是将它们聚集成两群的主要特征.今村蚌螨P-Ⅳ端腹部仅具有短矩,与外群最近,认为是我国的蚌螨属水螨中最原始的种类;雌螨前殖吸盘板的形态影响戊蚌螨亚属与多盘蚌螨亚属的种类聚类,由此推测可以将蚌螨P-Ⅳ端腹部的指状突与雌螨前殖吸盘板的形态也作为蚌螨亚属的区分特征.由于殖吸盘结构简单是祖征态之一,因此认为具有5对殖吸盘的邻近蚌螨比6对殖吸盘的厚蚌螨更原始.雌螨具有1对殖吸盘板的寄蚌螨亚属种类与沃蚌螨亚属种类在支序系统树中形成姊妹群,结果支持Vidrine 将沃蚌螨亚属从寄蚌螨亚属中分离的修订.     

Short Retroposons of the B2 Superfamily: Evolution and Application for the Study of Rodent Phylogeny

Short retroposons can be used as natural phylogenetic markers. By means of hybridization and PCR analysis, we demonstrate that B2 retroposon copies are present only in the three rodent families: Muridae, Cricetidae, and Spalacidae. This observation highlights the close phylogenetic relation between these families. Two novel B2-related retroposon families, named DIP and MEN elements, are described. DIP elements are found only in the genomes of jerboas (family Dipodidae) and birch mice (family Zapodidae), demonstrating the close relationship between these rodents. MEN element copies were isolated from the squirrel, Menetes berdmorei, but were not detected in three other species from the family Sciuridae. The MEN element has an unusual dimeric structure: the left and right monomers are B2- and B1-related sequences, respectively. Comparison of the B2, DIP, MEN, and 4.5S₁ RNA elements revealed an 80-bp core sequence located at the beginning of the B2 superfamily retroposons. This observation suggests that these retroposon families descended from a common progenitor. A likely candidate for this direct progenitor could be the ID retroposon. Received: 20 December 1996 / Accepted: 17 June 1997     

Ectoparasitic and phoretic arthropods of Virginia opossums (Didelphis virginiana) in central Tennessee

Thirteen species of ectoparasitic (12) or phoretic (1) arthropods were collected from 26 adult Virginia opossums, Didelphis virginiana, live-trapped from April through September 1987 in Davidson County, Tennessee. The cat flea Ctenocephalides felis and the American dog tick Dermacentor variabilis were the predominant species with respect to mean intensity and prevalence. The macronyssid mite Ornithonyssus wernecki and the atopomelid mite Didelphilichus serrifer, both specific parasites of this host, showed high intensities but low prevalences. Other fleas collected were Cediopsylla simplex, Ctenophthalmus pseudagyrtes, and Orchopeas howardi. The tick Amblyomma americanum, the myobiid mite Archemyobia inexpectatus, and the trombiculid (chigger) mites Eutrombicula splendens, Leptotrombidium peromysci (first record from this host) and Neotrombicula cavicola (first record from this host), were also recorded. One phoretic species, the glycyphagid mite Marsupialichus brasiliensis, was noted.     

记陕西蓝田晚中新世灞河组4种跳鼠(Dipodidae,Rodentia)化石

描述了陕西蓝田晚中新世早期灞河组的4属4种跳鼠:蓝田原跳鼠(新种)(Protalacta- ga lantianensis sp.nov.)、副跳鼠(未定种)(Paralactaga sp.)、原始三趾心颅跳鼠(新种)(Sal- pingotus primitivus sp.nov.)和小五趾心颅跳鼠(新种)(Cardiocranius pusillus sp.nov.)。蓝田原跳鼠兼有原跳鼠和副跳鼠的特征,可能是原跳鼠向副跳鼠进化过程中产生的一种过渡类型。副跳鼠和原始三趾心颅跳鼠及小五趾心颅跳鼠分别代表了副跳鼠属在中国的最早记录和心颅跳鼠亚科(Cardiocraniinae)在地史上的首次出现。4种跳鼠指示了蓝田地区当时的自然环境可能比现代更加干旱。     

Lung morphology in rodents (Mammalia, Rodentia) and its implications for systematics

A new nomenclature of the lung lobes and of the bronchial tree is presented, with which the lungs in 40 species of 11 rodent families are described. Whole, fixed lungs and silicone casts of the bronchial tree are tested for 23 characters, based on the distribution of lung lobes, the number and geometry of first order bronchi, the pulmonary blood supply, and lung symmetry. Ten lung morphotypes are recognized, seven of them representing one or more families: Castor type (Castoridae), Cryptomys type (Bathyergidae), Ctenodactylus type (Ctenodactylidae), Eliomys type (Gliridae), Myocastor type (Myocastoridae), Octodon type (Octodontidae and Echimyidae) and Rattus type (Sciuridae, Muridae pt. and Dipodidae). The Hydromys type is found only in Hydromys chrysogaster (Muridae), while Galea type A and B both appear in Galea musteloides (Caviidae). The data are phylogenetically analyzed by the program PAUP 4.0 using as outgroup Lagomorpha or Insectivora. On the species level, there are no well-resolved cladograms. On the family level, the cladograms do not contradict traditional rodent systematics with one exception: the Caviidae do not fall within Caviomorpha or even within the Hystricomorpha, but form a sister group to Dipodidae (Myomorpha). This appears to be a result of convergence. The lungs of Gliridae are more similar to those of Muridae than to those of Sciuridae. Included in the ingroup, Oryctolagus (Lagomorpha) forms a clade with Caviidae + Dipodidae. Thus, the "Glires hypothesis" is neither supported nor refuted.     

Systematics and evolution of the genus Paraceroglyphus and related taxa (Acari: Acaridae) associated with fleas (Insecta: Siphonaptera)

Collections of fleas from terrestrial Sciuridae from New Mexico and Montana yielded 2 species of acarid mites: Acarus monopsyllus from Ceratophyllus ciliatus and Paraceroglyphus cynomydis n. sp. from 4 species of Oropsylla. The genera Acarus, Paraceroglyphus, and Trichopsyllopus form a clade distinct from other genera of flea-associated mites, with Paraceroglyphus the sister group of the other 2 genera. Paraceroglyphus cynomydis is the sister group of a clade comprising P. xenopsylla and possibly P. californicus, with P. meles as the nearest outgroup.     

The correlation of the size of the pinna and the auditory bulla in specialized desert rodents

Relation between size of pinna and auditory bulla in specialized desert rodents (families Allactagidae, Dipodidae, Gerbillidae, Heteromyidae, and genus Selevinia) is statistically significant and described by hyperbola. The empirical distribution of respective sizes can be interpreted as a total of two main trends. One is enlargement of the pinna, the bulla remains small (most of Allactagidae), another is enlargement of the bulla, the pinna remains small (Gerbillidae, Heteromyidae, Selevinia, most Dipodidae). A few species display moderate enlargement of both pinna and bulla, while in Euchoreutes they both are hypertrophied. Biomechanical preconditions are supposed to be following. Enlargement of the pinna is responsible for increasing of both sensitivity and locatory abilities of the hearing organ, while enlargement of the bulla is responsible for the first function only. So, the two main trends are results of selection of development of the hearing organ at the level of either pinna or bulla. It is shown by dispersion and correlation analyses that size of the parts of the hearing organ under investigation are correlated mostly with mobility and escaping mode. It is concluded therefore that the rodents characterized by active and speedy escaping from predation display the first of the above trends, while the rodents characterized by passive escaping from predation display the second trend.     

Phylogeny and infrageneric classification of Symplocos (Symplocaceae) inferred from DNA sequence data

Symplocos comprises ～300 species of woody flowering plants with a disjunct distribution between the warm-temperate to tropical regions of eastern Asia and the Americas. Phylogenetic analyses of 111 species of Symplocos based on the nuclear ribosomal internal transcribed spacer (ITS) region and the chloroplast genes rpl16, matK, and trnL-trnF yielded topologies in which only one of the four traditionally recognized subgenera (Epigenia; Neotropics) is monophyletic. Section Cordyloblaste (subgenus Symplocos; eastern Asia) is monophyletic and sister to a group comprising all other samples of Symplocos. Section Palura (subgenus Hopea; eastern Asia) is sister to a group comprising all other samples of Symplocos except those of section Cordyloblaste. Symplocos wikstroemiifolia (eastern Asia) and S. tinctoria (southeastern United States), both of subgenus Hopea, form a clade that groups with S. longipes (tropical North America) and the species of subgenus Epigenia. The remaining samples of subgenus Hopea (eastern Asia) form a clade. Section Neosymplocos (subgenus Microsymplocos; Neotropics) is well nested within a clade otherwise comprising the samples of section Symplocastrum (subgenus Symplocos; Neotropics). Section Urbaniocharis (subgenus Microsymplocos; Antilles) groups as sister to the clade comprising Symplocastrum and Neosymplocos. The data support the independent evolution of deciduousness among section Palura and S. tinctoria. The early initial divergence of sections Cordyloblaste and Palura from the main group warrants their recognition at taxonomic levels higher than those at which they are currently placed. An inferred eastern Asian origin for Symplocos with subsequent dispersal to the Americas is consistent with patterns from other phylogenetic studies of eastern Asian-American disjunct plant groups but contrary to a North American origin inferred from the earliest fossil occurrences of the genus.     

THE EVOLUTION OF BIPEDALISM IN JERBOAS (RODENTIA: DIPODOIDEA): ORIGIN IN HUMID AND FORESTED ENVIRONMENTS

Mammalian bipedalism has long been thought to have arisen in response to arid and open environments. Here, we tested whether bipedalism coevolved with environmental changes using molecular and paleontological data from the rodent superfamily Dipodoidea and statistical methods for reconstructing ancestral characteristics and past climates. Our results show that the post‐Late Miocene aridification exerted selective pressures on tooth shape, but not on leg length of bipedal jerboas. Cheek tooth crown height has increased since the Late Miocene, but the hind limb/head‐body length ratios remained stable and high despite the environmental change from humid and forested to arid and open conditions, rather than increasing from low to high as predicted by the arid‐bipedalism hypothesis. The decoupling of locomotor and dental character evolution indicates that bipedalism evolved under selective pressure different from that of dental hypsodonty in jerboas. We reconstructed the habitats of early jerboas using floral and faunal data, and the results show that the environments in which bipedalism evolved were forested. Our results suggest that bipedalism evolved as an adaptation to humid woodlands or forests for vertical jumping. Running at high speeds is likely a by‐product of selection for jumping, which became advantageous in open environments later on.     

Phylogenetic relationships among Bactrocera species (Diptera: Tephritidae) inferred from mitochondrial DNA sequences

Several members of the dipteran family Tephritdae are serious pests because females lay eggs in ripening fruit. The genus Bactrocera is one of the largest within the family with over 500 described species arranged in 28 subgenera. The phylogenetic relationships among the various species and subgenera, and the monophyly of specific groups have not been examined using a rigorous phylogenetic analysis. Therefore, phylogenetic relationships among 24 Bactrocera species belonging to 9 subgenera were inferred from DNA sequence of portions of the mitochondrial 16S rRNA, cytochrome oxidase II, tRNA(Lys), and tRNA(Asp) genes. Two morphological characters that traditionally have been used to define the four groups within the subgenus Bactrocera were evaluated in a phylogenetic context by mapping the character states onto the parsimony tree. In addition, the evolutionary trend in male-lure response was evaluated in a phylogenetic context. Maximum parsimony analyses suggested the following relationships: (1) the genus Bactrocera is monophyletic, (2) the subgenus B. (Zeugodacus) is paraphyletic, (3) the subgenus B. (Daculus) is a sister group to subgenus B. (Bactrocera), and (4) the subgenus B. (Bactrocera) is monophyletic. The mapping analyses suggested that the morphological characters exhibit a simple evolutionary transition from one character state to another. Male-lure response was identified as being a labile behavior that has been lost on multiple occasions. Cue-lure response was plesiomorphic to methyl-eugenol response, and the latter has evolved independently within the Bactrocera and Zeugodacus groups of subgenera. The implications of our results for devising a coherent, consolidated classification for Bactrocera is discussed.     

Phylogeny of Oriental voles (Rodentia: Muridae: Arvicolinae): molecular and morphological evidence

The systematics of Oriental voles remains controversial despite numerous previous studies. In this study, we explore the systematics of all species of Oriental voles, except Eothenomys wardi, using a combination of DNA sequences and morphological data. Our molecular phylogeny, based on two mitochondrial genes (COI and cyt b), resolves the Oriental voles as a monophyletic group with strong support. Four distinct lineages are resolved: Eothenomys, Anteliomys, Caryomys, and the new subgenus Ermites. Based on morphology, we consider Caryomys and Eothenomys to be valid genera. Eothenomys, Anteliomys, and Ermites are subgenera of Eothenomys. The molecular phylogeny resolves subgenera Anteliomys and Ermites as sister taxa. Subgenus Eothenomys is sister to the clade Anteliomys + Ermites. Caryomys is the sister group to genus Eothenomys. Further, the subspecies E. custos hintoni and E. chinensis tarquinius do not cluster with E. custos custos and E. chinensis chinensis, respectively, and the former two taxa are elevated to species level and assigned to the new subgenus Ermites.     

Molecular phylogeny of the snubnose darters, subgenus Ulocentra (genus Etheostoma, family Percidae)

Snubnose darters comprise one of the largest subgenera of the percid genus Etheostoma. Many species are described based on differences in male breeding coloration. Few morphological synapomorphies have been proposed for the subgenus and their relatives, making it difficult to delineate monophyletic clades. The phylogenetic relationships of the 20 snubnose darter species of the subgenus Ulocentra and 11 members of its proposed sister subgenus Etheostoma were investigated with partial mitochondrial DNA sequences including 1033 bp encompassing the entire mitochondrial control region, the tRNA-Phe gene, and part of the 12S rRNA gene. Two hypotheses on the relationship and monophyly of the two subgenera were evaluated. Both maximum-parsimony and neighbor-joining analyses supported monophyly of the subgenus Ulocentra and resolved some species-level relationships. The banded darter, E. zonale, and its sister taxon, E. lynceum, were not closely related to the snubnose darters and appear to be diverged from the other members of the subgenus Etheostoma, fitting their former distinction as the recognized subgenus Nanostoma. The sister group to Ulocentra appears to be a restricted species assemblage within the subgenus Etheostoma containing E. blennioides, E. rupestre, E. blennius, and the E. thalassinum species group. The placement of the harlequin darter, E. histrio, is problematic, and it may represent a basal member of Ulocentra or of the restricted subgenus Etheostoma. Despite recent estimates of divergence times between nominal Ulocentra taxa, each species exhibits its own unique set of mtDNA haplotypes, providing no direct evidence for current genetic exchange between species. The nominal taxa of snubnose darters thus appear to be evolving independently from each other and therefore constitute valid species under the Phylogenetic Species Concept.     

The bushlike radiation of muroid rodents is exemplified by the molecular phylogeny of the LCAT nuclear gene

Phylogenetic relationships among 40 extant species of rodents, with an emphasis on the taxonomic sampling of Muridae and Dipodidae, were studied using sequences of the nuclear protein-coding gene LCAT (lecithin cholesterol acyl transferase). Analysis of 804 bp from the exonic regions of LCAT confirmed many traditional groupings in and around Muridae. A strong support was found for the families Muridae (represented by 29 species) and Dipodidae (5 species). Compared with Sciuridae, Gliridae, and Caviomorpha, the Dipodidae family appeared the closest relative of Muridae, confirming the suprafamilial Myodonta concept. Within the speciose family Muridae, the first branching leads to the fossorial Spalacinae and semifossorial Rhyzomyinae. The remaining components of Muridae appear as a polytomy from which are issued Sigmodontinae, Calomyscinae, Arvicolinae, Cricetinae, Mystromyinae, Nesomyinae, and some Dendromurinae (Steatomys and Dendromus). This phylogeny is interpreted as the result of a bushlike radiation at the end of the early Miocene, leading to emergence of most living subfamilies. The separation between three additional taxa, Murinae, Gerbillinae, and "Acomyinae" (which comprises the genera Acomys, Deomys, Uranomys, and Lophuromys), has occurred more recently from a common ancestor issued from the main basal radiation. As previously shown by other molecular studies, the vlei rats, Otomyinae, are nested within Old World Murinae. In the same way, the zokors, Myospalacinae, appear strongly nested within the hamsters, Cricetinae. Finally, we propose a sister group relationship between Malagasy Nesomyinae and south African Mystromyinae.     

Mites of the family Cheyletidae (Acari: Prostigmata): phylogeny, distribution, evolution and analysis of parasite-host relationship

A modern system, phylogeny, distribution and host parasite relationships of cheyletid mites (Acari: Prostigmatal Cheyletidae) is shortly discussed. According to the phylogenetic hypothesis proposed by Bochkov and Fain (2001), the family Cheyletidae includes now 15 tribes: Acaropsellini, Bakini, Cheletogenini, Cheletosomatini, Chelonotini, Cheyletiini, Cheyletiellini, Cheyletini, Cheletomorphini, Criokerontini, Metacheyletiini, Niheliini, Ornithocheyletiini, Teinocheylini and one unnamed tribe including the genera Caudacheles and Alliea. The parasitic Cheyletidae were primarily free-living predators, frequently associated with nests of vertebrates. These mites, being predators, have numerous preadaptations to the parasitic mode of life and they possess high ecological plasticity. Therefore it was quite easy for these mites to adapt to parasitism on the vertebrates. According to our phylogenetical hypothesis, the parasitism on vertebrates has arisen independently in several phylogenetic lines of the cheyletids associated with nests of vertebrates. Such transition from nest predation to true parasitism probably occurred repeatedly and at different times. The cheyletid mites are more widely represented on birds than on mammals. Possibly, it is in relation with a more early origin of parasitism in the cheyletids associated with bird nests than in the cheyletids associated with mammal nests. An independent origin of the parasitism in many different cheyletid phyletic lines, arisen significantly later than the origin of such a parasitic group as myobiid mites, is probably the main reason, which could explain the recent mosaic distribution of the Cheyletidae among the mammalian taxa. Parasitic associations between cheyletids and vertebrates are more common than the associations between these mites and the invertebrates. In the invertebrates, these associations are generally restricted to a phoresy. The zoogeographical analysis showed that this family as whole is characterised by the extremely low endemisms. The most part of the free-living cheyletid mites are associated with Holarctic region (87%) and, therefore, this family, probably, originated there.