Expression patterns of leg genes in the mouthparts of the spider<Emphasis Type="Italic"> Cupiennius salei</Emphasis> (Chelicerata: Arachnida)

The leg genes extradenticle, homothorax, dachshund, and Distal-less define three antagonistic developmental domains in the legs, but not in the antenna, of Drosophila. Here we report the expression patterns of these leg genes in the prosomal appendages of the spider Cupiennius salei. The prosoma of the spider bears six pairs of appendages: a pair of cheliceres, a pair of pedipalps, and four pairs of walking legs. Three types of appendages thus can be distinguished in the spider. We show here that in the pedipalp, the leg-like second prosomal appendage, the patterns are very similar to those in the legs themselves, indicating the presence of three antagonistic developmental domains in both appendage types. In contrast, in the chelicera, the fang-like first prosomal appendage, the patterns are different and there is no evidence for antagonistic domains. Together with data from Drosophila this suggests that leg-shaped morphology of arthropod appendages requires an underlying set of antagonistic developmental domains, whereas other morphologies (e.g. antenna, chelicera) may result from the loss of such antagonistic domains.Edited by M. Akam     

Note on the morphology and systematic position of Alkenopterus burglahrensis (Chelicerata: Eurypterida: Eurypterina) from the Lower Devonian of Germany

The holotype of the eurypterid Alkenopterus burglahrensis from the Lower Devonian (Siegenian) of the Westerwald area (Rhineland-Palatinate, SW Germany) is re-examined. In contrast to a previous concept, this species has flattened distal podomeres seven and eight and a movable flattened spine on the seventh podomere of the sixth prosomal appendage (podomere VI-7a). Based on the presence of such a slender paddle, A. burglahrensis has to be regarded as a basal ‘swimming eurypterid’ (Eurypterina) and is formally transferred here to the family Onychopterellidae, extending the stratigraphical range of this family into the Lower Devonian.     

An unusual short flagellum inMallomonas guttata (Synurophyceae,chrysophyta)

A short flagellum bearing unusual appendages inMallomonas guttata is described. These appendages, ca. 0.2 μm in diameter and 10–20 μm long, 7–10 in number, originate from the distal end of the short flagellum. Their membrane is continuous with the flagellar membrane and no particular structure was observed in appendage lumen.     

Captopodus poschmanni gen. et sp. nov. a new stem-group arthropod from the Lower Devonian Hunsrück Slate (Germany)

A new arthropod from the Lower Devonian Hunsrück Slate is described on the basis of four specimens. The body of Captopodus poschmanni comprises a head, a trunk with an anal portion. The high number of trunk appendages (≥66 segments) is unusual. The function of one pair of cupola-like structures of the head shield is unclear. The presence of large grasping appendages in the head superficially resembles the ‘short great appendages’ of other euarthropods and grasping appendages of thylacocephalans. The phylogenetic position of the arthropod cannot be determined in detail, though several morphological aspects indicate a phylogenetic position as a stem lineage representative of the Euarthropoda, the morphology of the trunk appendages seem to indicate a more advanced phylogenetic position. This new taxon underlines the exceptional diversity of arthropods within the Hunsrück Slate in comparison to other Devonian fossil sites and highlights the significance of the Hunsrück Slate for the evolution of early arthropods.     

Weinbergina,a xiphosuran arthropod from the devonian hunsrück slate

Weinbergina is the first synziphosurid in which ventral appendages have been found. It is so far unique among chelicerates in having six pairs of prosomal legs in addition to the chelicerae. This indicates that the pregeni tal segment with the last pair of walking legs, corresponding to the chilaria ofLimulus, must be counted with the prosoma. The first interpretation of the opisthosomal appendages is given. They appear to be plate-like and to combine the presence of lamellar gills of limulid type with flattened spines (so called gill filaments) of trilobitomorph type.Weinbergina apparently was adapted for a life on soft substrate but not for burrowing.     

The female reproductive organ in podocopid ostracods is homologous to five appendages: histological evidence from Liocypris grandis (Crustacea,Ostracoda)

The giant ostracod, Liocypris grandis, found in South African temporary pools, has five pairs of additional appendage-like structures, seemingly associated with the Female Reproductive Organ (FRO). Original investigations of these structures by light and scanning electron microscopy could not unequivocally determine whether or not these structures are homologous to genuine appendages and if so, to how many. The present paper investigates morphology and attachment of these structures by frontal and transversal micro-sectioning (thickness = 1 μ m) of the caudal soft parts of mature females of L. grandis. It is found that all five appendages (R1–R5) have one individual attachment point where they enter into the FRO. Also R3(I) and R3(II) have a single attachment only, these two parts thus represent one single appendage. Whereas R1–R4 have clear lateral insertions, R5 is caudally positioned. The fusion of this latter appendage with the FRO occurs internally, at the posterior edge of the bulge of the FRO and is not externally visible. It is herewith thus accepted that R1–R5 are genuinely endogenous structures (and not, for example, exogenous parasites), that the five structures are homologous to five appendages, and not for example to parts of biramous appendages. It follows from this that the female reproductive organ is homologous to at least five, not to one segment. This would bring the number of (original) thoracic segments in podocopid ostracods to at least 8. An additional pair of triangular structures, inserted in between the two FRO is here described as structures ‘X’. Their homology and function remain unknown.     

Halosarpheia japonica sp. nov. (Halosphaeriales, Ascomycota) from marine habitats in Japan

A new Halosarpheia species, collected from driftwood from Hakkeijima beach, Yokohama, Japan, is described and illustrated and is compared with other species of the genus. The new fungus was growing together with its anamorph on a piece of decaying wood. SSU and LSU rDNA sequences for both morphs were 99% similar. Phylogenetic analyses of SSU and LSU rDNA sequences of the both morphs confirm their anamorph–teleomorph relationship and placed the new fungus in the Halosarpheia sensu stricto clade with high statistical support. Halosarpheia japonica is characterized by its polar appendage that is initially enclosed in a cellular sheath and dissolves in water, the appendage then swells to form a huge tree-like structure. The other three species currently included in Halosarpheia sensu stricto differ from H. japonica by having two polar appendages that uncoil in water to form long filaments.     

A New Carcinosomatid Eurypterid From The Upper Silurian Of Northern Vietnam

A new carcinosomatid eurypterid, Rhinocarcinosoma dosonensis sp. nov., and Hughmilleria sp., are described from the Dô Son Formation of the Dô Son Peninsula, northern Vietnam. R. dosonensis is characterized by podomere 7 of prosomal appendage VI produced into an anterodistal spine, a metastoma with a cordate posterior margin, and an indented opisthosomal differentiation (i.e. preabdomen–postabdomen margin is concave). The Dô Son Formation was originally interpreted as Late Devonian (Givetian/Frasnian) in age but the unit containing the eurypterid assemblage is now considered much older (Late Silurian). A deltaic palaeoenvironmental setting is interpreted from biotic associations and sedimentological evidence.     

Erysiphe wadae: a new species of Erysiphe sect. Uncinula on Japanese beech

 A new species of Erysiphe sect. Uncinula is described and illustrated from Japan. Erysiphe wadae sp. nov., found on Japanese beech (Fagus crenata, Fagaceae), is characterized by having two types of appendages, i.e., a long (true) appendage arising from the equatorial zone of the ascomata, and a short appendage arising from the upper part of the ascomata. This characteristic is shared by E. simulans, E. australiana, E. flexuosa, E. liquidambaris, E. prunastri, and E. togashiana. Erysiphe wadae differs from the latter five species in its brown-colored appendage. Erysiphe simulans is most similar to E. wadae, but differs in its loosely uncinate appendage and smaller number of ascospores. Identity of the nucleotide sequences of the rDNA ITS region is 92.3% between the two species. The significance of the two types of appendage in taxonomy and phylogeny of powdery mildews is discussed based on molecular phylogenetic analysis. Received: November 8, 2002 / Accepted: January 29, 2003 Acknowledgments We are grateful to Drs. Yukio Harada and Hideki Naito for help in collecting powdery mildew specimens; Dr. Uwe Braun for providing the specimen of E. flexuosa; and Mr. Tetsuya Hirata and Miss Sanae Matsuda for nucleotide sequences of E. togashiana and E. flexuosa. This work was partially supported by a Grant-in-Aid for Scientific Research (No. 13660047) from Ministry of Education, Culture, Sports, Science and Technology, Japan.     

Distal-less expression in embryos of Limulus polyphemus (Chelicerata, Xiphosura) and Lepisma saccharina (Insecta, Zygentoma) suggests a role in the development of mechanoreceptors, chemoreceptors, and the CNS

The homeobox gene Distal-less (Dll) is well known for its participation in the development of arthropod limbs and their derivatives. Dll activity has been described for all groups of arthropods, but also for molluscs, echinoderms and vertebrates. Generally, Dll participates in the establishment of the proximo-distal-axis and differentiation along this axis. During our investigation of the expression pattern in the silverfish Lepisma saccharina and the horseshoe crab Limulus polyphemus, we found several expressions in late stages which cannot be explained with the "normal" limb-specific function. The antenna, cerci and terminal filament of the silverfish show a striped expression; single cells on the labrum, mandibles, maxillary palps and anal valves are also strongly stained by the Dll antibody. In addition to cell groups in the developing ganglia of the CNS, in the coxal endites and several nerve cells in femur and the trochanter of the prosomal limbs, the whole prosomal shield of Limulus polyphemus is surrounded by Dll-positive cell clusters. Furthermore, the lateral processes of the opisthosoma and the edges of the opisthosomal appendages are Dll positive. To get an indication of the cell fate of these regions, we examined hatched larvae and juvenile stages of both species with the SEM. We found a striking correlation of these Dll-positive areas and different sense organs, especially mechanoreceptors. Since many sense organs in arthropods are situated on the limbs, interpretation of the Dll expression in limbs is problematical. This has critical implications for comparative analysis of Dll expression patterns between arthropods and for the claim of homology between limb-like structures. Furthermore, we discuss the possibility of convergent appendage evolution in various bilaterian groups based on the improvement of spatial sensory resolution.     

Functional stability of the aristaless gene in appendage tip formation during evolution

The appendages of an insect are subdivided into distinct segments or podomeres. Many genes responsible for the regionalization of the growing limb into subdomains have been isolated from Drosophila. So far, only one gene is known in the leg that is solely required for specifying the distal-most pattern element—the pretarsal claw. In Drosophila, the gene aristaless is expressed in the centre of the antennal and leg imaginal disc that represents the most distal position of appendages, and in a proximal region. When Drosophila aristaless function is impaired, antennae and legs develop without their distal-most structures—the arista and the claw. We describe here the analysis of aristaless in the beetle Tribolium—an insect that shows a different, more ancestral mode of appendage formation than Drosophila. In Tribolium, appendages grow out continuously during embryogenesis, and no imaginal discs are formed. Tribolium aristaless (Tc-al) expression starts midway during appendage elongation, and is seen in a distal and a proximal position of head and trunk appendages. At the end of embryogenesis, Tc-al is seen in four expression domains in the leg, in the dorsal epidermis, and ventrally in every segment in lateral groups of cells, presumably the histoblasts. Like in the Drosophila adult, Tc-al is required in the larva for the formation of the most distal structures of the leg and the antenna as revealed by RNAi experiments. We conclude that aristaless is evolutionarily robust, meaning that it has retained its expressional and functional characteristics, although a heterochronic change of the process of appendage elongation took place towards the evolution of the highly derived diptera.Edited by D. Tautz     

The functional morphology of mating in the Silurian eurypterid, Baltoeurypterus tetragonophthalmus (Fischer, 1839)

The functional morphology of the genital appendages, horn organs and modified prosomal limb III of the Silurian eurypterid, Baltoeurypterus tetragonophthalmus (Fischer, 1839) (Chelieerata: Eurypterida) is investigated. The longer type A genital appendage, interpreted as a female structure, is associated with a pair of horn organs, regarded as spermathecae. The shorter type B appendage, interpreted as male, is associated with scimitar lobes on prosomal limb III, regarded as clasping structures. A mechanism is proposed by which the type A appendage was lowered through flexure of the genital operculum acting along sutures in the cuticle. A model of mating is suggested, involving the male depositing an unstalked spermatophorc on the substrate and the female retrieving it for storage in the spermathecae. This model is more arachnid-like than mating in Limutus (an extant aquatic chclicerate), and suggests that sperm transfer using spcrmatophores and storage of sperm in spermathecae, allowed the timing of egg production in curyptcrids to be controlled.     

Phosphatocopine ostracodes with preserved appendages from the Upper Cambrian of Sweden

Mülkr. Klaus 1. 1979 01 IS: Phosphatocopine ostracodes with preserved appendages from the Upper Cambrian of Sweden. Lethaia . Vol. 12. pp. 1–27. Oslo. ISSN 0024–1164.
More than 400 specimens of phosphatocopine ostracodes. representing different moult stages of Vestrogothia. Falites and Hesslandona have been recovered, with body and appendage structures preserved in minute detail. from anthraconites in the Upper Cambrian of Sweden and drift boulders deriving from that area. The secondary phosphatization of these structures, leading to preservation. is extensively discussed. It comprises even the inner lamella, but the abdomen is not preserved. Hypostome and lower lip are well developed. The large antennula is composed of a basipodite with probably two podomeres and a long exopodite with up to 18 podomeres bearing long spines, forming a natatory organ, and a broad endopodite of two to three podomeres. The biramous organization of the antennula is a unique, most primitive feature of phylogenetic significance. The antennae and up lo four additional pairs of appendages are also biramous and similar to the antennulae, not yet differentiated for specialized functions. Only the last pair is uniramous, but a second branch may have developed in subsequent moult stages not represented in the material. The movement of food into the mouth was achieved by the endites of antennulae and antennae, a function transferred backwards to other appendages during the course of ostracode phylogeny and reflected in ontogenetic trends among Recent ostracodes. The phosphatocopines were nectobenthic, filtering plankton feeders.     

Structure and membrane affinity of new amphiphilic siderophores produced by Ochrobactrum sp. SP18

The coastal α-proteobacterium Ochrobactrum sp. SP18 produces a suite of three citrate-derived, cell-associated amphiphilic siderophores, ochrobactins A–C. The ochrobactins are composed of a citric acid backbone amide-linked to two lysine residues. Each ε-amine of lysine is hydroxylated and acylated forming two hydroxamic acid moieties. One of the acylated appendages of each ochrobactin is (E)-2-decenoic acid. The other acylated appendages for ochrobactins A–C are (E)-2-octenoic acid, octanoic acid and (E)-2-decenoic acid, respectively. The ferric ochrobactin complexes are photoreactive in UV light, producing an oxidized ligand with loss of 46 mass units that can still coordinate Fe(III). The relative partitioning of the apo-ochrobactins, Fe(III) ochrobactins and Fe(III) photoproducts into 1,2-dimyristoyl-sn-glycero-3-phosphocholine vesicles is presented. The ochrobactins are the first example of aerobactin-based siderophores with two fatty acid appendages produced in a suite with varying acyl appendage lengths.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

The arthropod Offacolus kingi (Chelicerata) from the Silurian of Herefordshire,England: computer based morphological reconstructions and phylogenetic affinities

The small, non-biomineralized, three-dimensionally preserved arthropod Offacolus kingi Orr et al. from the Wenlock Series (Silurian) of Herefordshire, England, is re-evaluated, and the new family Offacolidae erected. This new study is based on specimens which have been serially ground, reconstructed by computer and rendered in the round as coloured models. Offacolus possesses a prosomal appendage array similar to that of Limulus, but also bears robust and setose exopods on appendages II-V which are unlike those found in any other arthropods. Opisthosomal appendages are similar in number and morphology to the book-gills of Limulus. Cladistic analysis places Offacolus basally within the Chelicerata, as a sister taxon to the eurypterids and extant chelicerates, but more derived than the Devonian Weinbergina.     

Annelids from the Lower Devonian Hunsrück Slate (Lower Emsian, Rhenish Massif, Germany)

Abstract:  Four new genera and species of annelid, Hunsrueckochaeta hohensteini , Ewaldips feyi , Crocancistrius lutzi and Scopyrites magnus , are described from the Hunsrück Slate (Lower Emsian) of Germany, as well as new material of Bundenbachochaeta eschenbachensis   Bartels and Blind, 1995 . The specimens preserve details of the appendages and other aspects of the morphology as a result of pyritization. A phylogenetic analysis using the morphological data matrix of Rouse and Fauchald places four of the five genera basal to the Aciculata; the fifth Ewaldips falls within the Scolecida. The diversity of annelids represented in the Hunsrück Slate is similar to that in the other major Palaeozoic Konservat-Lagerstätten that yield them, including the Cambrian Burgess Shale and the Late Carboniferous Mazon Creek biota.     

Double-stranded RNA interference in the spider Cupiennius salei: the role of Distal-less is evolutionarily conserved in arthropod appendage formation

Chelicerates represent a basal arthropod group, which makes them an excellent system for the study of evolutionary processes in arthropods. To enable functional studies in chelicerates, we developed a double-stranded RNA-interference (RNAi) protocol for spiders while studying the function of the Distal-less gene. We isolated the Distal-less gene from the spider Cupiennius salei. Cs-Dll gene expression is first seen in cells of the prosomal segments before the outgrowth of the appendages. After the appendages have formed, Cs-Dll is expressed in the distal portion of the prosomal appendages, and in addition, in the labrum, in two pairs of opisthosmal (abdominal) limb buds, in the head region, and at the posterior-most end of the spider embryo. In embryos, in which Dll was silenced by RNAi, the distal part of the prosomal appendages was missing and the labrum was completely absent. Thus, Dll also plays a crucial role in labrum formation. However, the complete lack of labrum in RNAi embryos may point to a different nature of the labrum from the segmental appendages. Our data show that the expression of Dll in the appendages is conserved among arthropods, and furthermore that the role of Dll is evolutionarily conserved in the formation of segmental appendages in arthropods.     

Characterization of a Novel <Emphasis Type="Italic">Bacillus thuringiensis</Emphasis> Phenotype Possessing Multiple Appendages Attached to a Parasporal Body

Bacillus thuringiensis is a bacterium best known for its production of crystal-like bodies comprised of one or more Cry-proteins, which can be toxic to insects, nematodes or cancer cells. Although strains of B. thuringiensis have occasionally been observed with filamentous appendages attached to their spores, appendages in association with their parasporal bodies are extremely rare. Herein we report the characterization of Bt1-88, a bacterial strain isolated from the Caribbean that produces a spore–crystal complex containing six long appendages, each comprised of numerous thinner filaments approximately 10 nm in diameter and 2.5 μm in length. Each of the multi-filament appendages was attached to a single, small parasporal body located at one end of the bacterial spore. Biochemical tests, 16S rDNA gene sequencing, and the identification of two Cry proteins by partial protein sequencing (putatively Cry1A and Cry2A), unambiguously identified Bt1-88 as a strain of B. thuringiensis. Bt1-88 represents the second reported strain of B. thuringiensis possessing a parasporal body/appendage phenotype characterized by one or more long appendages, comprised of numerous filaments in association with a parasporal body. This finding suggests that Bt1-88 is a member of a new phenotypic class of B. thuringiensis, in which the parasporal body may perform a novel structural role through its association with multi-filament appendages.     

Inclusion appendages associated with the intraerythrocytic rickettsial parasiteAnaplasma marginale are composed of bundled actin filaments

Summary Anaplasma marginale, a tick-borne rickettsia that infects erythrocytes of cattle, occurs within a parasitophorous vacuole or inclusion body. A tail-like inclusion appendage, composed of multiple filaments, occurs in association with the inclusion body membrane. The composition and function of the inclusion appendage have not been determined. In this study, theA. marginale inclusion appendage in bovine erythrocytes was found to be composed of actin filaments as determined by labeling with rhodamine-conjugated phalloidin. Electron microscopy studies revealed that theA. marginale inclusion appendages differed from F-actin tails reported previously in association with other pathogens in eukaryotic cells because these highly ordered structures were organized into regularly occurring striations, and the appendages were adhered directly to the parasitophorous vacuole membrane. In addition, actin appendages have not been described previously in erythrocytes. The potential role of the inclusion appendage associated withA. marginale in bovine erythrocytes and recently fed ticks is discussed.     

Tube foot preservation in the Devonian crinoid Codiacrinus from the Lower Devonian Hunsrück Slate,Germany

Fossilized tube feet are described on Codiacrinus schultzei Follmann from the Lower Devonian Hunsrück Slate of Germany. This is the first definitive proof of tube feet on any fossil crinoid. Three lightly pyritized, flattened tube feet are preserved in a single interray of this cladid crinoid. The tube feet were at least 7 mm long. Their preservation is very similar to the tube feet reported previously from a Hunsrück ophiuroid, except that the Codiacrinus tube feet have small papillae, similar to living crinoids.