Ultrastructural investigation of the vesicular glands in Scutigera coleoptrata (Chilopoda,Notostigmophora)

In the notostigmophoran centipedes, two pairs of vesicular glands have evolved. These paired glands are situated in the first and second trunk segment and open via cuticular ducts in the upper part of the particular pleura. The vesicular glands of Scutigera coleoptrata were investigated using light and, for the first time, electron microscopical methods. The glands consist of wide sac‐like cavities that often appear vesicular. The epithelia of both glands are identically structured and consist of numerous glandular units. Each of these units consists of four different cells: a single secretory cell, a small intermediary cell, and one proximal and one distal canal cell. The intermediary cell forms a conducting canal and connects the secretory cell with the canal cells. Proximally, the intermediary cell bears microvilli, whereas the distal part is covered with a distinct cuticle. The cuticle is a continuation of the cuticle of the canal cells. This investigation shows that the ultrastructure of glandular units of the vesicular glands is comparable to that of the glandular units of other epidermal glands in Chilopoda and Diplopoda, although the glands look completely different in the light microscope. Thus, it is likely that the vesicular glands and epidermal glands share the same ground pattern. With regard to specific differences in the cuticular lining of the intermediary cells, a common origin of epidermal glands in Myriapoda and Hexapoda is not supported. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

Influence of the ventral nerve cord on the spermatogenetic cycle in Lithobius forficatus L. (Myriapoda,Chilopoda)

The role of the ventral nerve cord has been investigated in Lithobius forficatus L., using ganglionic destructions, ganglionic implantations and ganglionic electrostimulations. It seems that a stimulating principle is present in the sub-oesophageal ganglion. Its action is not so important as that of the pars intercerebralis.     

Ultrastructural investigation of a salivary gland in a centipede: structure and origin of the maxilla I-gland of Scutigera coleoptrata (Chilopoda, Notostigmophora)

The maxilla I-gland of Scutigera coleoptrata was investigated using light and electron microscopy methods. This is the first ultrastructural investigation of a salivary gland in Chilopoda. The paired gland opens via the hypopharynx into the foregut and extends up to the third trunk segment. The gland is of irregular shape and consists of numerous acini consisting of several gland units. The secretion is released into an arborescent duct system. Each acinus consists of multiple of glandular units. The units are composed of three cell types: secretory cells, a single intermediary cell, and canal cells. The pear-shaped secretory cell is invaginated distally, forming an extracellular reservoir lined with microvilli, into which the secretion is released. The intermediary cell forms a conducting canal and connects the secretory cell with the canal cell. Proximally, the intermediary cell bears microvilli, whereas the distal part is covered with a distinct cuticle. The cuticle is a continuation of the cuticle of the canal cells. This investigation shows that the structure of the glandular units of the salivary maxilla I-gland is comparable to that of the glandular units of epidermal glands. Thus, it is likely that in Chilopoda salivary glands and epidermal glands share the same ground pattern. It is likely that in compound acinar glands a multiplication of secretory and duct cells has taken place, whereas the number of intermediary cells remains constant. The increase in the number of salivary acini leads to a shifting of the secretory elements away from the epidermis, deep into the head. Comparative investigations of the different head glands provide important characters for the reconstruction of myriapod phylogeny and the relationships of Myriapoda and Hexapoda.     

The peristomatic structures of Lithobiomorpha (Myriapoda, Chilopoda): comparative morphology and phylogenetic significance

A comparative survey of the epipharynx and hypopharynx of lithobiomorph centipedes by light and scanning electron microscopy examines 18 species that sample the major groups of both families, the Lithobiidae and Henicopidae. Cladistic analysis of 11 characters of the peristomatic structures together with 29 additional morphological characters serves as a basis for interpreting the evolution of the lithobiomorph peristomatic structures. Scutigeromorpha is used for outgroup comparison in the framework of a homology scheme for the basic components of the epi- and hypopharynx. Compared to other chilopods, the monophyly of Lithobiomorpha is supported by a row of distinctive bottle-shaped gland openings at the border between the labral and clypeal parts of the epipharynx, as well as by a distinctive shape of the hypopharynx. Paired rows of elongate spines on the clypeal part of the epipharynx are an apomorphic character of Lithobiidae. The transformation of these spine rows into a few groups of branching spines is characteristic for the Monotarsobius group sensu Verhoeff. Similar groups of branching clypeal spines characterize the Anopsobiinae within Henicopidae, whereas Henicopinae possess a dense cluster of short, simple spines instead. The recently described genus Dzhungaria is resolved closer to Henicopinae than to Anopsobiinae, a hypothesis supported by a field of grooves on the medial labral part of the epipharynx. Monophyly of Henicopidae does not receive unique support from the peristomatic structures although two homoplastic characters contribute to this node; among these, the reduction of a median spine field between clypeal and labral parts of the epipharynx to a narrow transverse band also supports a close relationship between the Ezembius group and Hessebius within Lithobiidae. An Ezembius+Hessebius clade is additionally supported by the absence of a transverse bulge between the clypeal and labral parts of the epipharynx, a character otherwise present in all lithobiomorph species studied so far. Lithobius is resolved as polyphyletic, with different species being most closely related to such genera as Australobius, Hessebius and Pleurolithobius.     

Phylogenetic systematics of the colorful, cyanide-producing millipedes of Appalachia (Polydesmida, Xystodesmidae, Apheloriini) using a total evidence Bayesian approach

Here, we provide an exemplar-approach phylogeny of the xystodesmid millipede tribe Apheloriini with a focus on genus-group relationships-particularly of the genus Brachoria. Exemplars for the phylogenetic analysis were chosen to represent the maximum breadth of morphological diversity within all nominal genera in the tribe Apheloriini, and to broadly sample the genus Brachoria. In addition, three closely related tribes were used (Rhysodesmini, Nannariini, and Pachydesmini). Morphological and DNA sequence data were scored for Bayesian inference of phylogeny. Phylogenetic analysis resulted in polyphyletic genera Brachoria and Sigmoria, a monophyletic Apheloriini, and a "southern clade" that contains most of the tribal species diversity. We used this phylogeny to track morphological character histories and reconstruct ancestral states using stochastic character mapping. Based on the findings from the character mapping study, the diagnostic feature of the genus Brachoria, the cingulum, evolved independently in two lineages. We compared our phylogeny against prior classifications using Bayes factor hypothesis-testing and found that our phylogenetic hypothesis is inconsistent with the previous hypotheses underlying the most recent classification. With our preferred total-evidence phylogeny as a framework for taxonomic modifications, we describe a new genus, Appalachioria; supply phylogenetic diagnoses of monophyletic taxa; and provide a phylogeny-based classification for the tribe Apheloriini.     

Phylogenetic systematics of the genus Echinococcus (Cestoda: Taeniidae)

Echinococcosis is a serious helminthic zoonosis in humans, livestock and wildlife. The pathogenic organisms are members of the genus Echinococcus (Cestoda: Taeniidae). Life cycles of Echinococcus spp. are consistently dependent on predator–prey association between two obligate mammalian hosts. Carnivores (canids and felids) serve as definitive hosts for adult tapeworms and their herbivore prey (ungulates, rodents and lagomorphs) as intermediate hosts for metacestode larvae. Humans are involved as an accidental host for metacestode infections. The metacestodes develop in various internal organs, particularly in liver and lungs. Each metacestode of Echinococcus spp. has an organotropism and a characteristic form known as an unilocular (cystic), alveolar or polycystic hydatid. Recent molecular phylogenetic studies have demonstrated that the type species, Echinococcus granulosus, causing cystic echinococcosis is a cryptic species complex. Therefore, the orthodox taxonomy of Echinococcus established from morphological criteria has been revised from the standpoint of phylogenetic systematics. Nine valid species including newly resurrected taxa are recognised as a result of the revision. This review summarises the recent advances in the phylogenetic systematics of Echinococcus, together with the historical backgrounds and molecular epidemiological aspects of each species. A new phylogenetic tree inferred from the mitochondrial genomes of all valid Echinococcus spp. is also presented. The taxonomic nomenclature for Echinococcus oligarthrus is shown to be incorrect and this name should be replaced with Echinococcus oligarthra.     

Ultrastructural organization of the anal organs in the anal capsule of Craterostigmus tasmanianus Pocock, 1902 (Chilopoda, Craterostigmomorpha)

We describe the ultrastructural organization of the anal organs of Craterostigmus tasmanianus, which are located on the ventral side of the bivalvular anal capsule. Each part of the capsule bears four pore fields with several anal pores. The pores lead into a pore canal, which is surrounded by the single-layered epithelium of the anal organs. Each anal organ is composed of four different cell types: transporting cells of the main epithelium, junctional cells, isolated epidermal glands, and the cells forming the pore canal. The transporting cells exhibit infoldings of the outer cell membranes, forming a basal labyrinth and a poorly developed apical complex. The cells are covered by a specialized cuticle with a widened subcuticular layer. Only the cuticle of the main epithelium is covered by a mucous layer, secreted by the epidermal glands. The ultrastructural organization of the anal organ is comparable to the coxal and anal organs of other pleurostigmophoran Chilopoda. It is likely that the coxal and anal organs of the Pleurostigmophora are homologous, due to their identical ultrastructural organization. Differences concerning the location on the trunk of Pleurostigmophora are not sufficient to reject a hypothesis of homology. Anal organs are found not only in Craterostigmomorpha, but also in most adult Geophilomorpha, and in larvae and most adults of Lithobiomorpha. The anal organs of C. tasmanianus are thought to play an important role in the uptake of atmospheric water. J. Morphol.     

Ultrastructure of the epidermal maxilla II-gland of Scutigera coleoptrata (Chilopoda, Notostigmophora) and the ground pattern of epidermal gland organs in Myriapoda

The epidermal maxilla II-gland of Scutigera coleoptrata was investigated using light and electron microscopy. The glandular epithelium surrounds a spacious integumental cavity at the base of the maxilla II. The gland is formed as a compound gland organ that is composed of thousands of epidermal gland units. Each of them consists of four different cell types: a secretory cell, an accessory or intermediary cell, and a proximal and distal canal cell. The intermediary and the two canal cells form a conducting canal. Only in the most distal part of the intermediary cell is the canal lined by a cuticle. In the area of the two canal cells, the conducting canal is completely covered by a cuticle. The canal passes through the cuticle and opens into the spacious integumental cavity, which serves as a secretion reservoir. The structural organization of the epidermal maxilla II-gland was compared to that of other compound epidermal gland organs in Chilopoda and Diplopoda. All these glandular organs in Myriapoda share the same ground pattern.     

Detecting taxonomic signal in an under-utilised character system: geometric morphometrics of the forcipular coxae of Scutigeromorpha (Chilopoda)

To date, the forcipules have played almost no role in determining the systematics of scutigeromorph centipedes though in his 1974 review of taxonomic characters Markus Würmli suggested some potentially informative variation might be found in these structures. Geometric morphometric analyses were used to evaluate Würmli's suggestion, specifically to determine whether the shape of the forcipular coxa contains information useful for diagnosing species. The geometry of the coxae of eight species from the genera Sphendononema, Scutigera, Dendrothereua, Thereuonema, Thereuopoda, Thereuopodina, Allothereua and Parascutigera was characterised using a combination of landmark- and semi-landmark-based sampling methods to summarize group-specific morphological variation. Canonical variates analysis of shape data characterizing the forcipular coxae indicates that these structures differ significantly between taxa at various systematic levels. Models calculated for the canonical variates space facilitate identification of the main shape differences between genera, including overall length/width, curvature of the external coxal margin, and the extent to which the coxofemoral condyle projects laterally. Jackknifed discriminant function analysis demonstrates that forcipular coxal training-set specimens were assigned to correct species in 61% of cases on average, the most accurate assignments being those of Parascutigera (Parascutigera guttata) and Thereuonema (Thereuonema microstoma). The geographically widespread species Thereuopoda longicornis, Sphendononema guildingii, Scutigera coleoptrata, and Dendrothereua linceci exhibit the least diagnostic coxae in our dataset. Thereuopoda longicornis populations sampled from different parts of East and Southeast Asia were significantly discriminated from each other, suggesting that, in this case, extensive synonymy may be obscuring diagnosable inter-species coxal shape differences.     

On the systematics of some marine haploporids (Trematoda) with the description of a new species of Megasolena Linton, 1910

Megasolena mikra sp. nov. is described from the queen angelfish, Holacanthus ciliaris (Linnaeus), off Florida, USA. The new species can be differentiated from all other species of Megasolena Linton, 1910 except Megasolena littoralis Muñoz, George-Nascimento, and Bray, 2017 in possessing testes that are smaller in diameter than the ovary. The new species can be differentiated from M. littoralis in lacking tegumental spines and possessing oral sucker papillae. Molecular data are provided for two species each of Cadenatella Dollfus, 1946, Hapladena Linton, 1910, and Megasolena Linton, 1910. Bayesian inference analysis of concatenated internal transcribed spacer region-2 (ITS2) and partial 28S rDNA sequences of 50 haploporoids revealed 1) a monophyletic Atractotrematidae Yamaguti, 1939 sister to the rest of the haploporoids tested; 2) a paraphyletic Megasoleninae Manter, 1935 – if Hapladena is included; and 3) a monophyletic Cadenatellinae Gibson and Bray, 1982 sister to the ‘mugilid’ haploporids. The ‘mugilid’ haploporids formed a monophyletic clade consisting of the subfamilies Chalcinotrematinae Overstreet and Curran, 2005, Forticulcitinae Blasco-Costa, Balbuena, Kostadinova, and Olson, 2009, Haploporinae Nicoll, 1914, and Waretrematinae Srivastava, 1937. Based on our analysis we restrict the Megasoleninae to include Megasolena, Vitellibaculum Montgomery, 1957, and Metamegasolena Yamaguti, 1970, all of which have species with two testes. To accommodate the former megasolenine taxa with a single testis, we erect the Hapladeninae subf. nov. for species in Hapladena and tentatively, Myodera Montgomery, 1957. Our results further support that haploporoids had a common marine ancestor with two testes, and that members of the Haploporoidea Nicoll, 1914 underwent diversification following a shift from a primarily marine life history with eupercarian hosts to a more euryhaline one with diadromous hosts (namely mullet).     

Notes on Pauropoda (Myriapoda) from USA, with descriptions of two new species

Two new species of Pauropoda are described from USA, Kionopauropus alyeskaensissp. n. (Pauropodidae), and Eurypauropus arcuatussp. n. (Eurypauropodidae). The genus Kionopauropus is reported from the Western Hemisphere for the first time.     

Phylogenetic analysis of the Brachyura (Crustacea, Decapoda) based on characters of the foregut with establishment of a new taxon

The Brachyura, within the decapod crustaceans, is one of the most species-rich taxa with up to 10 000 species. However, its phylogenetic history, evolution and fossil record remain subjects of controversy. In our study, we examined the phylogenetic relationships of the Brachyura based on morphological characters of the foregut. The cladistic analysis supports a monophyletic Brachyura including the Dromiidae and Raninidae. A clade comprising Dromiidae and Dynomenidae forms the most basal assemblage within the Brachyura, followed by the Homolidae and Latreilliidae. As a result, neither Podotremata nor Archaeobrachyura form a clade. In contrast, foregut data suggest that the classical taxon Oxystomata, comprising Calappidae, Parthenopidae, Dorippidae, Leucosiidae, Cymonomidae and Raninidae, is monophyletic. This makes the Heterotremata paraphyletic or polyphyletic. A newly established taxon, Neobrachyura, embraces some representatives of the Heterotremata and the monophyletic Thoracotremata.     

Molecular systematics of Eumolpinae and the relationships with Spilopyrinae (Coleoptera, Chrysomelidae)

The 3400 species of Eumolpinae constitute one of the largest subfamilies of leaf beetles (Chrysomelidae). Their systematics is still largely based on late 19th century monographs and remains highly unsatisfactory. Only recently, some plesiomorphic lineages have been split out as separate subfamilies, including the southern hemisphere Spilopyrinae and the ambiguously placed Synetinae. Here we provide insight into the internal systematics of the Eumolpinae based on molecular phylogenetic analyses of three ribosomal genes, including partial mitochondrial 16S and nuclear 28S and complete nuclear 18S rRNA gene sequences. Sixteen morphological characters considered important in the higher-level systematics of Eumolpinae were also included in a combined analysis with the molecular characters. All phylogenetic analyses were performed using parsimony by optimizing length variation directly on the tree, as implemented in the POY software. The data support the monophyly of the Spilopyrinae outside the clade including all sampled Eumolpinae, corroborating their treatment as a separate subfamily within the Chrysomelidae. The systematic placement of the Synetinae remains ambiguous but consistent with considering it a different subfamily as well, since the phylogenetic analyses using all the available evidence show the representative sequence of the subfamily also unrelated to the Eumolpinae. The Megascelini, traditionally considered a separate subfamily, falls within the Eumolpinae. Several recognized taxonomic groupings within Eumolpinae, including the tribes Adoxini or Typophorini, are not confirmed by molecular data; others like Eumolpini seem well supported. Among the morphological characters analyzed, the presence of a characteristic groove on the pygidium (a synapomorphy of the Eumolpini) and the shape of tarsal claws (simple, appendiculate or bifid) stand out as potentially useful characters for taxonomic classification in the Eumolpinae.     

Origin and higher-level relationships of psoroptidian mites (Acari: Astigmata: Psoroptidia): evidence from three nuclear genes

Phylogenic relationships of the Psoroptidia, a group of primarily parasitic mites of vertebrates, were investigated based on sequences from three nuclear genes (4.2 kb aligned) sampled from 126 taxa. Several morphological classification schemes and a recent molecular analysis, suggesting that the group may not be monophyletic were statistically rejected by newly generated molecular data, and the results are robust under a range of analytical and partition strategies. Six families Psoroptidae, Lobalgidae (mammalian parasites), Pyroglyphidae (house dust mites and parasites inside feather calamus), Turbinoptidae (upper respiratory track parasites of birds), Psoroptoididae (downy feather mites), and Epidermoptidae (skin parasites of birds) form a well-supported monophyletic group (the epidermoptid-psoroptid complex). These relationships, recovered by combined and separate analyses of all gene partitions, were previously suspected based on some morphological evidence, but evidence has been dismissed as resulting from convergence based on similar parasitic ecologies. The existence of the epidermoptid-psoroptid complex and the statistical rejection of Sarcoptoidea (the morphology-based group joining all mammal-associated mites) indicate that current classification criteria, influenced as they are by host preferences, need to be reassessed for non-pterolichoid superfamilies. However, two of our findings remain sensitive to analytical methods and assumptions: (i) the families Heterocoptidae and Hypoderatidae as the first and second closest outgroups of Psoroptidia, respectively, and (ii) the superfamily Pterolichoidea (including Freyanoidea) forming a sister clade to the remaining psoroptidian superfamilies. Our findings suggest that (i) house dust mites (Pyroglyphidae: Dermatophagoidinae) originated from a parasitic ancestor within the core of Psoroptidia, violating a basic principle of evolution that it is virtually impossible for a permanent parasite to become free-living, and (ii) there were at least two shifts from presumably avian to mammalian hosts.     

A new species of the genus Paracryptops from India, with remarks on the generic and specific characters (Chilopoda: Scolopendromorpha: Cryptopidae)

Paracryptops spinosus , a new centipede species from Delhi, is described. The species is unique in the genus in that its anal legs have their femora and tibiae beset distally with extraordinary spines. It is pointed out that the exceedingly short poison claws supposed to be incapable of meeting in the middle cannot be depended upon to identify the genus Paracryptops . This character is apparent, not real and should be discarded. It is also pointed out that the number of tibial teeth used as a key character at species level, being subject to age variation, can be of limited taxonomic value and should be used with caution.     

中国森林田鼠族系统分类研究进展(啮齿目:仓鼠科:䶄亚科)

森林田鼠族（Myodini）是䶄亚科（Arvicolinae）的重要类群,广泛分布于整个全北区及东洋区部分区域,该族目前存在的分类学问题是缺乏化石标本、短时间的物种分化、有限分子样本和部分类群采样困难等综合因素影响的结果。近年中国森林田鼠族的系统分类研究成果主要有：（1）绒鼠属（Eothenomys）的采样和系统发育研究基本涵盖了全部类群,发现1新亚属和4新种,分别为川西绒鼠亚属（Ermites）和石棉绒鼠（Eothenomys shimianensis）、金阳绒鼠（E.jinyangensis）、美姑绒鼠（E.meiguensis）、螺髻山绒鼠（E.luojishanensis）,原来被普遍接受的黑腹绒鼠福建亚种（E.melanogaster colurnus）、中华绒鼠康定亚种（E.chinensis tarquinius）、西南绒鼠康定亚种（E.custos hintoni）被证实为3个独立有效种,滇绒鼠（E.eleusis）、丽江绒鼠（E.fidelis）的种级分类地位进一步得到确认;大绒鼠贡山亚种（E.miletus confinii）和黑腹绒鼠滇西亚种（E.melanogaster libonotus）实为克钦绒鼠（E.cachinus）的同物异名。（2）基于形态和分子系统发育的研究支持恢复东亚䶄属（Craseomys）的属级分类地位,同时解决了䶄属（Myodes）的非单系起源问题;研究进一步证实山西䶄（Craseomys shanseius）应为棕背䶄的山西亚种（Craseomys rufocanus shanseius）。（3）高山䶄属（Alticola）的平颅高山䶄亚属（Platycranius）分类地位受到分子系统发育研究结果质疑,该亚属的唯一物种平颅高山䶄（Alticola strelzowi）与高山䶄属指名亚属物种聚在同一枝;库蒙高山䶄（A.stracheyi）实为斯氏高山䶄（A.stoliczkanus）的同物异名。基于近年系统分类研究结果,目前中国森林田鼠族分布有5属共30种。