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A large number of complete specimens together with numerous disarticulated sclerites of the eodiscinid trilo- bite Sinodiscus changyangensis Zhang in Zhou et al., 1977 have been collected from the lower Cambrian Shuijingtuo Formation in Changyang, Hubei Province, South China. An ontogenetic series is established based on the immature and mature exoskeletons including the previously unknown pro- taspides and meraspides, in particular. No further substages can be differentiated in the protaspid specimens herein. Changes that took place during the meraspid period include the addition of postcephalic segments and prominent pygidial larval notches in early meraspid development which became progressively less distinct and disappeared in degree 2. Two holaspid stages are recognized based on the addition of a new pygidial segment, indicating that the start of the holaspid phase preceded the onset of the epimorphic phase and accordingly, its developmental mode is attributed to the protarthrous pattern. The trunk segmentation schedule of S. changyangensis is discussed, which is similar to other prim- itive eodiscoid trilobites, that is, as the boundary between the thorax and pygidium migrated posteriorly, there is no change in the number of the trunk segments. The processes of libera- tion of the thoracic segment and segment insertion into the pygidium are separated from one another, and the two differ- ent mechanisms, somitogenesis and tagmosis, progress inde- pendently during the ontogenetic development of the postcephalic region of these primitive eodiscinids.

Several specimens of polychaetes were collected on a sandy beach at depths of 0–1 m in the northeastern Venezuelan using a PVC corer (0.018 m2), taking sediment cores of 20 cm depth. Samples were sieved using a 0.5-mm open mesh. Specimens were fixed in 10 % formaldehyde and preserved and stored in 70 % ethanol. They were then examined using compound microscopes, and structures with taxonomic value were dissected. The characteristics of the specimens examined allowed to identify them as belonging to the genus Malacoceros. A new species of spionid polychaete is described from the Caribbean Sea, Malacoceros longiseta n. sp. This species clearly differs from all other species of Malacoceros Quatrafages (Mag Zool Paris 2:1–16, 1843), by the presence of lateral subterminal prostomial horns, extremely long notopodial capillary chaetae on the three first chaetigers (similar to the ones observed for Dispio uncinata Hartman), the length of the branchiae in relation to the notopodial lamellae, and the morphology of the pygidium.

Previous studies on preferences of phoretic deutonymphs of Uropodina for attachment sites have shown that they frequently select smooth and hydrophobic surfaces. The aim of our study was to provide the detailed morphological and topographical characteristics of beetle body surfaces to which deutonymphs frequently attach and to verify how the presence of setae and surface sculpture affects deutonymph attachment. The study was conducted on Uropoda orbicularis (Müller, 1776) and its common beetle carriers: Aphodius prodromus (Brahm, 1790), Aphodius fimetarius (Linnaeus, 1758), Onthophagus nuchicornis (Linnaeus, 1758) and Margarinotus carbonarius (Hoffmann, 1803). Morphology and topography of elytra, femora, propygidia and pygidia of beetles were analysed mainly using SEM methods supported with CLSM and AFM techniques. The hypothesis that deutonymphs may attach to surfaces covered with setae, if seta density is low enough not to disturb mite movement, was tested. The study revealed that deutonymphs attach to surfaces of various types as follows: (i) smooth, (ii) hairy, i.e., covered with setae, (iii) flat and (iv) sculptured. Smooth body parts and body parts covered with setae of low density were most frequently and intensively occupied with deutonymphs. Surfaces of high seta density were avoided by mites. Within elytra of Aphodius beetles, deutonymphs definitely preferred flat surfaces of elytral intervals. On the contrary, densely punctuated propygidium and pygidium in M. carbonarius were heavily infested with deutonymphs. We conclude that carrier surface morphology and topography are important for Uropodina deutonymph attachment, but these two factors cannot fully explain the observed relation.

The ability to enrol effectively evolved several times among trilobites. Here, we show that, unlike most redlichiid trilobites that could not enrol, both morphotypes of Eccaparadoxides pradoanus from the middle Cambrian of Spain enrolled so as to enclose most of the ventral surface beneath the exoskeleton and possessed specialized articulat- ing devices that facilitated this behaviour. The holaspid tho- rax of all E. pradoanus was divided into two principal regions. The boundary between these marked a transition from anterior segments with short pleural spines, fulcra and ridge-and-groove inner pleural regions to posterior seg- ments with longer, acuminate pleural spines that lack fulcra and inner pleural regions. Devices that aid articulation, such as fulcra with short articulating pleural surfaces, the petaloid articulating facet and long articulating half rings, are concentrated in the anterior region. These features, and the large number of specimens preserved in various degrees of enrolment, suggest an enrolment procedure in which the rear part of the trunk, containing both the posterior thorax and the pygidium, rotated as a single unit without signifi- cant internal flexure. As these posterior trunk articulations were apparently not required to permit enrolment, concen- trating flexure in the anterior may have presaged the cau- dalized condition seen in many derived trilobite groups that encapsulated, in which a larger proportion of the trunk segments were allocated to the mature pygidium, and there- fore unable to articulate.

Namalycastis jaya sp. n. (Polychaeta: Nereididae: Namanereidinae) is described from the southern coast of Kerala in southwest India. One important characteristic feature of the species is the lack of notochaetae in all parapodia, a characteristic that it shares with at least two other species, Namalycastis elobeyensis Glasby, 1999 and Namalycastis hawaiiensis Johnson, 1903. It differs from N. elobeyensis by virtue of its smaller antennae, unequal eye size, bilobed acicular neuropodial ligule and multi-incised pygidium rim. Moreover, it differs from N. hawaiiensis byfewer teeth on the serrated blades of the sub-neuroacicular falciger in chaetiger 10, and by possessing finely serrated falcigers in posterior segments. Beyond morphological analyses, molecular phylogenetics was used for the first time for Namalycastis to support population monophyly and recognition of the new species. The analysis, using both mitochondrial and nuclear data, corroborated the morphological analysis in suggesting that our specimens represent an as yet undescribed species, Namalycastis jaya sp. n., which forms a monophyletic group among the sampled nereidid taxa. Finally, a taxonomic key for Namalycastis species recorded from the Indian region is provided.

Orphnine scarab beetles (Orphninae) are widely distributed in the tropical and subtropical regions of the southern continents except for Australia. The catalogue of nominal taxa of orphnines includes 2 tribes, 15 genera, and 195 species. Diagnosis of the group, based on adult morphological characters, is as follows: antennae 10-segmented with 3-segmented club; mandibles with 2–4 scissorial teeth and well developed mola; labrum and mandibles protruding past clypeus and visible from above; scutellum well developed in winged species, reduced but distinct in wingless species; wings with distinct anal area; apices of anterior tibia in males without spur but normally with a few robust setae; anterior coxa with longitudinal hollow on anterior surface; tarsi with 2 similar claws; middle and hind tibiae with 2 apical spurs; abdominal sternite 2 with sub-triangular to rounded plectrum; dorsal surface of hind coxae with oval flat stridulatory file; pygidium partly hidden under elytra; parameres symmetrical; bursa copulatrix sacciform, membranous; spermatheca C-shaped, not sclerotized; accessory vaginal glands developed; abdomen with 2 sclerotized tergites (VII–VIII) and 6 visible sternites (III–VIII). Preliminary phylogenetic analysis based on 47 characters of adult morphology shows that the tribe Aegidiini Paulian is a natural, monophyletic group. The genus Stenosternus Karsch described from a single specimen from Sao Tome Island (Gulf of Guinea), is morphologically more similar to the New World taxa than to the Old World ones and is provisionally placed in Aegidiini. The tribe Orphnini Erichson seems non-monophyletic and has no synapomorphies. The genus Orphnus is apparently a polyphyletic group and it needs taxonomic revision. The hypothesis on sister-group relationship of Orphninae and Allidiostomatinae, based on molecular data, is not supported by the morphological characters. The stridulatory organs (the putative synapomorphy of Orphninae + Allidiostomatinae) are not identical in these groups; the mouthparts and female genitalia are essentially different. Orphninae have chewing mouthparts with large scissorial teeth and well developed mola, which is characteristic of generalist saprophagous species. Allidiostomatinae have mandibles with scissorial teeth and mola reduced; they also have sclerotized bursa copulatrix and sclerotized mandibular duct which opens on the dorsal side near condyle. Considering the present day development of alpha-taxonomy of most orphnine taxa, especially the speciose genus Orphnus, it seems premature to propose changes in higher classification of the subfamily. To clarify the phylogenetic position of the Orphninae among scarab beetles it is essential to include representative members of all taxa of orphnine lineage (sensu Browne, Scholtz, 1998) into the analysis.

The morphology and anatomy of some important structures of S. oryzae have been described. All major differences between the two sexes, in respect of habitus, rostrum, olfactory sensilla of antenna, eyes, pronotum, mesonotum, metanotm, metendosternite, abdomen, propygidium, pygidium, proventriculus, male and female genitalia have been fully discussed and figured. The main differences between male and female of S. oryzae are, in a side view, the ventral surface of the female is in a straight line, but in the male it is curved posteriorly; the rostrum in the male is smaller and thicker and less curved than in the female and is dorsally, dull and closely punctured, but in the female, shining and sparsely punctured and The number of ommatidia in male (~ about 167) is more than female (~ about 154). Other differences in the rest of structures are shown below.

The ontogeny of the trilobite Protopeltura aciculata (Angelin, 1854) is described on the basis of material from the upper Cambrian (Furongian) of Andrarum (Skane) and Hjelmsater (Vastergotland), Sweden. P. aciculata is present in the Parabolina brevispina and Parabolina spinulosa zones. Protopeltura aciculata is represented by all stages of growth, from early protaspides to holaspides, although most of the specimens are disarticulated and precise degrees are unknown. The cranidia have therefore been allocated to five morphological groups. Cuticular sculpture of the cranidia changes throughout ontogeny. Large tubercles are present in earlier stages, disappear gradually in middle meraspid stages and are replaced with a very faint granulation. The transitory pygidium, relatively large and shield-shaped with upwardly and backwardly directed marginal spines in early meraspides, later becomes very small, triangular-shaped and lacking spines as a late meraspid and holaspid. The development of hypostomes and librigenae is also described. Protopeltura aciculata shows major intraspecific variations throughout development, especially regarding the pygidium where variation is much less constrained than in many other olenids. This high developmental plasticity may be a survival strategy for a trilobite living in a stressed environment. Protopeltura inhabited a dysoxic environment, possibly unusually prone to localised spreading of anoxic or toxic water. Some morphs may have been less vulnerable than others to such stresses, surviving by chance and thus enabling the species to continue.

GraduateUniversityofChineseAcademyofSciences,Beijing100049, ChinaSummaryParasite-host associations among insects and mammals orbirds are well attended by neontological studies [1]. AnEocene bird louse compression fossil [2, 3] and severalfleaspecimensfromEoceneandOligoceneambers[4–8],re-ported to date, are exceptionally similar to living louse andflea taxa. But the origin, morphology, and early evolutionof parasites and their associations with hosts are poorlyknown[9,10]duetosparserecordsofputativeectoparasiteswith uncertain classification in the Mesozoic, most lackingmouthpart information and other critical details of the headmorphology [11–15]. Here we present two primitive flea-likespecies assigned to the Pseudopulicidae Gao, Shih et Renfamilia nova (fam. nov.), Pseudopulex jurassicus Gao, Shihet Ren genus novum et species nova (gen. et sp. nov) fromthe Middle Jurassic [16] and P. magnus Gao, Shih et Rensp.nov.fromtheEarlyCretaceousinChina[17].Theyexhibitmany features of ectoparasitic insects. Large body size andlong serrated stylets for piercing tough and thick skin orhides of hosts suggest that these primitive ectoparasitesmight have lived on and sucked the blood of relatively largehosts, such as contemporaneous feathered dinosaurs and/or pterosaurs or medium-sized mammals (found in the EarlyCretaceous, but not the Middle Jurassic).ResultsHerein, we report from the mid-Mesozoic of China two well-preserved fossil ectoparasitic insects, exhibiting manyfeatures of ectoparasitic insects, e.g., wingless body coveredwith stiff, posteriorly directed setae and bristles, short andbeaded antennae, reduced eyes, absent ocelli, robust mouth-parts with serrated stylets (laciniae), scythe-like pretarsalclaws, and ctenidia on the tibiae. These specimens sharewith extant fleas, especially with the most basal flea lineageTungidae [18], some striking similarities, but given a lack ofsome diagnostic features of Siphonaptera, their systematicpositions are deemed as Order incertae sedis.TaxonomyOrder Incertae sedis.Family Pseudopulicidae Gao, Shih et Ren fam. nov.Diagnosis. Robust and elongate piercing-sucking mouth-parts with serrated stylets; antennae with more than15 segments, flagellomeres compact; thorax relatively small,apterous; legs elongate with numerous stout setae, distinctctenidia on the tibiae, and pretarsus with strong scythe-shaped claws; body covered with stiff, posteriorly directedsetae and bristles.Remarks. Different from extant fleas in the absence of pro-notal and genal ctenidia on body (except for Tungidae), lackof the uniquely modified jumping hind legs, distinct ctenidiaon the tibiae, more developed eyes, detailed structure ofmouthparts with serrated stylets, antennae with more than15 segments, not laterally compressed abdomen, and largerbody size. Although Tarwinia [11] possesses many similardiagnostic characters, it is tentatively excluded from Pseudo-pulicidae fam. nov. due to lacking mouthpart information.Pseudopulex Gao, Shih et Ren, gen. nov.Type species. Pseudopulex jurassicus Gao, Shih et Ren sp.nov.Etymology. The generic name is a combination of prefixpseudo-, indicating ‘‘with visual similarity to,’’ and Latin wordof Pulex, meaning ‘‘flea.’’ Gender masculine.Species included. The type species and Pseudopulexmagnus Gao, Shih et Ren sp. nov.Diagnosis. Same as familial diagnosis.Pseudopulex jurassicus Gao, Shih et Ren, sp. nov., seeFigures 1A–1J and Figures S1A–S1C available onlineHolotype. CNU-NN2010001, a well-preserved almost com-plete body in ventral view (Figures 1A and 1B; Figures S1Aand S1B). Deposited in the Key Lab of Insect Evolution andEnvironmental Changes, Capital Normal University, Beijing,China.Locality and Age. Jiulongshan Formation, the Late MiddleJurassic (Bathonian-Callovian boundary, 165 million yearsago [Mya]) [16]; Daohugou Village, Ningcheng County, InnerMongolia, China.Etymology. The specific epithet jurassicus derives fromLatin ‘‘jurassicus,’’ referring to the age of this fossil.Diagnosis. Different from P. magnus by the number ofantennal segments, presence of tibiae ctenidia, and maxillarylaciniae. Serrated teeth of P. jurassicus cover the entiresurface of stylet, whereas those of P. magnus cover only theedges.Description. Body length 17.00 mm excluding antennae.Head small and trapezoid shape. Eyes reduced and ocelliabsent. Antennal flagellum 16 segments visible, short andstout, slightly compressed, connected to both sides of headin clear depressions (Figure 1G). Mouthparts specialized,long and strong (at least 3.44 mm), much longer than twicethe length of head, labrum unclear, maxillary palps threesegments visible as preserved, labial palps four segmentspreserved, but the exact number unknown, maxillary laciniaeforming some deeply notched serrations, extending frombase to tip (Figures 1A–1F). Thorax small, about one-fifth ofthe abdomen in length. Abdomen not laterally compressed,with eight segments based on eight spiracles (segmentationis not clear), covered with dense and long bristles directingposteriorly, possessing a pygidium with sensilia on the ninthtergite (Figure S1C). Legs unusually long, coxae slightly

Male and female reproductive systems of Stolas conspersa (Germar) (Coleoptera, Chrysomelidae, Cassidinae). The male and female reproductive systems of Stolas conspersa (Germar, 1824) are described and illustrated for the first time. The male reproductive system shows no difference from the subfamily pattern, which is a tubular well-developed median lobe; "Y", "V" or T-shaped tegmen; reduced pygidium; internal sac membranous and tubular; flagellum generally well developed needle-like structured and gastral spiculum absent. However, the female differs from the pattern proposed for Stolas in two aspects: ovary with 28 ovarioles and a reduced ampulla with indistinct velum.Sistema reprodutivo masculino e feminino de Stolas conspersa (Germar) (Coleoptera, Chrysomelidae, Cassidinae). O sistema reprodutivo do macho e fêmea de Stolas conspersa (Germar, 1824) são descritos pela primeira vez. Neste estudo, foi observado que o macho não diverge do padrão proposto para a subfamília, que é edeago tubular e bem desenvolvido; tégmen em forma de "Y", "V" ou "T"; pigídio reduzido; saco interno membranoso e tubular; flagellum geralmente bem desenvolvido, em forma de agulha e espículo gastral ausente. Por outro lado, o sistema reprodutor feminino diverge do padrão usual proposto para o gênero Stolas em dois aspectos: ovário com 28 ovaríolos e ampola reduzida com velum não distinto.

The Emu Bay Shale Konservat-Lagerstätte (Cambrian Series 2, Stage 4) on Kangaroo Island, South Australia, is the source of two new non-biomineralized artiopodan arthropods. Squamacula buckorum n. sp. is the first record outside of China of a genus otherwise known only from its type species, S. clypeata, from the Chengjiang biota. The Australian species displays the long cephalic doublure and spiniform exopod setae that are apomorphic for this genus, provides new information on the alimentary tract and midgut glands (the latter preserved as three-dimensional, permineralized structures), and indicates interspecific variability in trunk segment numbers. The distribution of Squamacula strengthens the biogeographic connections between early Cambrian “Burgess Shale-type” biotas of Australia and South China. Australimicola spriggi n. gen. n. sp. represents a monotypic genus resolved in a cladistic analysis of Cambro-Ordovician artiopodans as most closely related to or within Conciliterga (a clade containing Helmetia, Kuamaia, Kwanyinaspis, Rhombicalvaria, Saperion, Skioldia, and Tegopelte). Compared with other members of this clade from Chengjiang and the Burgess Shale, the new genus is diagnosed by an elongate trunk with 23 thoracic tergites having spatulate pleural tips and a small pygidium possessing a single, elongate pair of pleural spines, with specimens also showing a hypostome attached to an anterior (or prehypostomal) sclerite, antennae, short endopods, an annulated alimentary tract, and a series of three-dimensional, permineralized midgut glands. An alternative relationship between Australimicola and the Early Ordovician–Early Devonian Cheloniellida explains the shared anterior flexure of trunk pleurae but forces dubious homologies in other characters, such as dorsally-articulated furcae versus spines.

Phylogenetic relationships among species of the family Olenidae (Trilobita, Ptychopariida) were traditionally established by the direct comparison of morphological features and the evaluation of stratigraphic placement. In this study, a cladistic analysis of Parabolinella Brogger is conducted in order to test the supposed monophyly of the genus and to determine the relationships of its species. Twenty-one taxa (sixteen of this genus) from the late Cambrian and Tremadocian of Baltica, Avalonia, Laurentia, South China and the western margin of Gondwana were revised. Forty non-ordered characters were considered in the analysis, 34 belonging to the cephalon, four to the pygidium, and two to the thorax. Some of these characters were specially defined for this study. The phylogenetic analysis was performed under unweighted parsimony. The obtained tree is partially consistent with the stratigraphic record of the studied species and shows correspondence with global late Cambrian and early Ordovician palaeogeography. Parabolinella may have originated in deep water sites of the Oaxaca region, Mexico and then migrated to other regions. The genus diversified in outer shelf facies of the Baltica Province during the late Cambrian and Tremadocian, whereas it would have dispersed to the South China Province in early Tremadocian times. Copyright © 2012 John Wiley & Sons, Ltd.

Introduction Polychaetes are segmented marine worms with body segments separated by a complete or incomplete septum. In most polychaetes the whole body cavity is filled with gametes during the breeding season. Platynereis dumerilii (Pl. dumerilii), which has an incomplete septum was shown to develop a single gonadal structure for gamete production located in the neck region. However, in Perinereis nuntia (Pe. nuntia), which has a complete septum separating each segment, the developmental feature of gametes remains unknown. To clarify this, the marker gene vasa was used to trace the development of germ cells throughout the life stages of Pe. nuntia. Results In three-segmented juveniles, Pn-vasa was expressed in the parapodia and in the two cells localized in the pygidium. During the addition of a new segment, Pn-vasa positive cells in the pygidium increased from two to four and two new Pn-vasa positive cells were found in the newly-generated segment. In adults, Pn-vasa was expressed in a large cell cluster at the distal end of the parapodia, in smaller cell clusters (which had an elongated form in the trunk area of the parapodia), and in oocytes in the coelomic cavity. This may suggest that germ cells settle in the parapodia and later translocate into the coelomic cavity to develop into oocytes. Conclusion Our observations will help in understanding the mechanism of germ cell development in all body segments of Pe. nuntia. We hypothesize that primordial germ cells are supplied from the pygidium to every newly-generating segment which later settle in the parapodium. This will explain how polychaetes can generate gametes in each body segment, even those that are independently separated with a complete septum. Electronic supplementary material The online version of this article (doi:10.1186/s40851-015-0035-y) contains supplementary material, which is available to authorized users.

A restudy of the Burgess Shale arthropod Emeraldella brocki suggests novel interpretations of its morphology. We show that the morphology is more plesiomorphic than previously assumed, particularly regarding tagmosis. The cephalon probably only incorporates three limb-bearing postantennular segments. The trunk is not differentiated and consists of 12 tergite-bearing segments and a styliform telson. Limb structure is generally similar to that of other artiopods except for a tripartite exopod and a high degree of differentiation of podomere proportions along the body. A phylogenetic analysis of 20 fossil arthropod taxa based on 36 characters renders E. brocki as a basal taxon within a monophyletic group that comprises all artiopods included. Autapomorphies of this taxon are a filiform antennula and a bilobate exopod that carries lamellae proximally. Trilobites are nested within a group of artiopods sharing a pygidium. Agnostus pisiformis is retrieved as the sister taxon to the stem-lineage crustacean Oeland...

A complete molted exoskeleton of the asteropygine phacopid trilobiteGreenops widderensisLieberman and Kloc, 1997 from the Middle Devonian (Givetian) Widder Formation in southwestern Ontario, Canada that has suffered predatory trauma provides insights into the sequence of regeneration of segments. The molt configuration is such that it is possible to interpret the molting technique used by the trilobite. Predatory trauma affected four areas of the exoskeleton. The pygidium shows loss of the spinose margin on one side and damage to a single spine on the other; one genal spine has been broken and partially regrown; and the posterior of the glabella has been removed. It is thought that the first three traumas occurred during life, as these areas affected show signs of exoskeletal regeneration. The fourth trauma probably occurred to the exuvium. Analysis of the degree of regeneration of the pygidial pleurae indicates that there was an anteroposterior polarity to the regeneration. Other examples in the literature suggest that this regeneration polarity pattern may have been widespread in trilobites. It is suggested that, as in modern arthropods and annelids, this sequential regeneration was under the control of segmentation polarity genes.

Three thousand seven hundred disarticulated remains together with several articulated specimens of the Cambrian Series 2 ptychopariid trilobite Strenuaeva inflata Ahlberg and Bergstrom, 1978 have been collected from the Tornetrask area, northern Sweden. The material provides significant new data on the morphology, ontogeny, moulting and enrolment of the species. Two distinct morphotypes, possibly an expression of sexual dimorphism, are recognized. The morph with a pair of bulbs in the frontal area, interpreted as brood pouches, is considered to represent females. Statistical treatment of the length/width ratio in cranidia reveals isometric growth during ontogeny for both morphotypes. The transition from the meraspid to holaspid ontogenetic period has been established through recognition of the successive development of the number of thoracic segments in articulated late meraspides. Throughout its life cycle, S. inflata went through 11 meraspid degrees and at least 17 holaspid growth stages. Inferred moult ensembles and exuviae reveal the successive opening of cephalic sutures and the function of the rostral plate during exuviation. As in other ellipsocephalid trilobites in which enrolment is known, the pygidium and two or three thoracic segments of S. inflata are concealed beneath the cephalon (spiral enrolment) during complete enrolment. (Less)

Five species of Pauropoda belonging to four genera have been identified from New Zealand. The genus Samarangopus is reported from there for the first time. Four new species are described by the first author: one in Pauropodidae, Decapauropus whanganuiensis n. sp., two in Eurypauropodidae, Samarangopus plicatus n. sp. and S. constellatus n. sp., and one in a new family, Eirmopauropodidae, Eirmopauropus distichos n. gen., n. sp. The new family Eirmopauropodidae is diagnosed as having head and pygidium free, not covered by 1st and 6th tergite respectively; head and tergites sclerotised with setae in regular transversal rows.

Esteve, J., Zamora, S., Gozalo, R. & Linan, E. 2010: Sphaeroidal enrolment in middle Cambrian solenopleuropsine trilobites. Lethaia, 10.1111/j.1502-3931.2009.00205.x Fifty specimens belonging to species of Solenopleuropsis and Pardailhania from Spain and France demonstrate sphaeroidal enrolment in Cambrian trilobites for the first time. These solenopleuropsines show novel coaptative structures in different regions of the exoskeleton: in the cephalon there are vincular furrows and notches; in the thorax an articulating facet is developed at the pleural margins, with a ball and socket connection on the adaxial most portion, and an articulating half-ring axially; the pygidium possesses an articulating facet. The interaction of these coaptative structures resulted in a sphaeroidal enrolment that was a progressive act from the first articulation between the occipital ring and the first segment to the pygidial articulating facet. A similar type of sphaeroidal enrolment is observed in the Devonian trilobite Phacops. Both Cambrian and Devonian trilobites developed a vincular furrow in the ventral surface of the cephalon to close their bodies tightly. In both cases, this is probably a convergent adaptation to protect against predators and obrution. Indeed, the enrolled trilobites are very common in obrution deposits restricted to shallow and soft muddy substrates. □Coaptative structures, convergence, Murero Formation, Pardailhania, Solenopleuropsinae, Solenopleuropsis.

Denierota juarezi, new species, is described from the Sierra Madre del Sur in Guerrero, Mexico. The only other congener, D. kraatzi (Haag-Rutenberg), is allopatric and ranges from Chiapas into Central America. The new species differs from D. kraatzi primarily in the structure of the male trochanters and pygidium.

The paper records the rediscovery of the rare Urodontidius enigmaticus Louw, 1993 in South Africa, based on specimens reared from galls in the succulent leaves of Ruschia versicolor. The original account of some of the morphological characters of the species is corrected, and its habitus, antennae, pygidium and genitalia are illustrated. Its life history and galling habit on its host plant are described and illustrated, and its larva is compared with those of the genera Urodontellus Louw and Urodontus Louw, which represent different larval types with different life histories. The silk-spinning habits of the Urodontellus larva are briefly described. A tribute to the late Schalk Louw is presented, together with a list of his publications on weevils.

A revision of paradoxidid trilobites reveals that previous identifications of specimens from Sardinia and Spain as the Nordic trilobite species Paradoxides brachyrhachis Linnarsson, 1883, are mistaken. The southern species, occurring also in France, is here referred to Eccaparadoxides mediterraneus (Pompeckj, 1901). Main differences are seen in the preocular field, pleural furrow and pygidium. The species P. brachyrhachis is referred with question to the genus Mawddachites Fletcher 2007.

Articulated meraspid and holaspid exoskeletons of Guangxiaspis guangxiensis from the Guole Township, Jingxi County, Guangxi Province, China, are preserved in mudstone deposited during an obrution event. The species has a short dorsal pre-cranidial median suture that splits ventrally into a pair of posteriorly divergent connective sutures. The rostral plate of G. guangxiensis is thus triangular in outline, as in the co-occurrent Shergoldia laevigata, which also bore a conterminant hypostome. These two taxa appear to be closely related. The cephalic venter of Shergoldia laevigata has recently been interpreted to suggest a diphyletic origin of the median suture within the order Asaphida, but Guangxiaspis guangxiensis, Shergoldia laevigata and other tsinaniid trilobites display several characters reminiscent of members of the non-asaphide suborder Leiostegiina. These include swellings adjacent to the margins of the L1 glabellar lobe, the shape and furrows of the glabella, a semi-circular pygidium with a long and thin axis, and macrospinous first opisthopleurae of the holaspid pygidium. Based on these characters and on other new information on the early ontogeny of other tsinaniids, all these taxa likely belong within Leiostegiina. This suggests that the median suture arose independently in corynexochide and asaphide trilobites. The degree of convergence between S. laevigata and members of the derived asaphide family Asaphidae was remarkable. Guangxiaspis guangxiensis shows marked morphological change during both meraspid and holaspid ontogeny and might include more than a single morphotype.

The first completely enrolled adult scutelluids assigned to the Paralejurus brongniarti group (Pragian) and to Paralejurus rehamnanus (Emsian) from southern Morocco are described. The Pragian taxon exhibits volvation in the sphaeroidal mode and a simple morphological configuration of the external thoracic pleurae with edge to edge fit between each other and against both the posterior edge of the genae and the anterior edge of the pygidium, respectively. Evolutionary trends towards perfect tightness of enrollment and the avoidance of lateral shearing are obvious in later Moroccan Paralejurus such as P. rehamnanus and allies from the Lower/Middle Devonian transition. In these forms, coaptative devices, such as submarginal grooves on the posterior edges of the genae and external thoracic pleurae, ventral fulcral processes and sockets, as well as bulging of the anterolateral pygidial doublures and corresponding depressions in genal doublures, were developed. The possible role of terrace ridges to assist in coaptation and/or to obtain hermetic closure of articulation gaps during volvation is discussed. Morphological and behavioural traits point to a semi-endobenthic lifestyle of Paralejurus with enrollment being an exceptional defence strategy against predatory attack. These traits that differentiate Paralejurus from all other scutelluid taxa are regarded as ecological rather than phylogenetical in origin. Consequently, the assignment of Paralejurus to the subfamily “Paralejurinae” within the Scutelluidae is favoured.

Sphaerodorids belonging to the genus Ephesiopsis were collected on the outer continental shelf and slope off the coast of São Paulo State, during the REVIZEE/Score Sul/Benthos Program. Ephesiopsis guayanae Hartman and Fauchald 1971 from off Guyana is the only species previously known in this genus. Ephesiopsis shivae sp. nov., differs from E. guayanae in having a longer body, macrotubercles with smaller terminal papillae, and simple chaetae not as pointed distally; eyes are absent in the new species. Four macrotubercles with terminal papillae were observed on the pygidium of E. shivae sp. nov. Such macrotubercles were not reported in E. guayanae, although the specimens are complete; the presence of these structures in the new species necessitating emendation of the genus diagnosis.

We used morphological data for a phylogenetic analysis of the genus Petaloptoctus (Maldanidae: Polychaeta). We found three most parsimonious phylogenetic trees with length = 37, CI = 0.89 and RI = 0.92. The genus Petaloproctus is monophyletic, and is supported by a strongly arched prostomium, pinnate capillary setae, and an anal cup with a reduced dorsal border. Our proposal for the systematisation of the subgroups of Petaloproctus is (( P. tenuis ( P. terriculus + P. neoborealis )) ( P. ornatus ((( P. cirratus + P. dentatus ) ( P. macrodentatus ( P. borealis ( P. vallejoi n. sp. + P. socialis )))))). We also regard Nicomachinae as a monophyletic group with the following synapomorphies: prostomium short and arched; nuchal grooves short and curved; cephalic plate lost; anal pore opening on margin of anal plate. Monophyly of Micromaldane and Nicomache was also supported in the analysis. Petaloproctus vallejoi n. sp. has 19 setigerous and one asetigerous pre-anal segment. The prostomium is rounded, forming a keel with curved laterals. Nuchal grooves are short and deep, strongly curved outwards. Each neuropodium has one acicular spine on setigers 1-3. The pygidium has a large ventral border, and a reduced dorsal border. The anus is terminal, and close to the margin of the anal plate, surrounded by divergent folds.

The hitherto poorly known, monotypic trilobite genus Fenestraspis from the Lower Devonian of Bolivia is revised and its original assignment to the Synphoriinae supported. The thoracic morphology of the genus remains very poorly known. Fenestraspis is morphologically unusual because of the development of extensive fenestrae in the pleural region of the pygidium and apparently of the thorax; the presence of upwardly directed spines on the cephalon, thorax and pygidium; and the exceptionally large and highly elevated eyes with the palpebral rim projecting outwards above the visual surface. The function of the fenestrae remains uncertain. If they formed openings in the body of the trilobite in life they may have allowed circulation of oxygenated water to the limb exites so that respiration could have been maintained while the trilobite was enrolled. If they were covered with a flexible membrane, they may have been secondary respiratory structures or had a sensory function. The Synphoriinae is regarded as a subfamily of the Dalmanitidae rather than as an independent family of the Dalmanitoidea as proposed by some authors. The type species of the poorly known monotypic genus Dalmanitoides from the Lower Devonian of Argentina is illustrated photographically for the first time and compared with Fenestraspis.

Several thousand disarticulated remains together with a few complete enrolled specimens of the lower Cambrian eodiscoid trilobite Calodiscus lobatus (Hall, 1847) have been collected at two outcrop areas in Sweden. The material reveals new details of morphology and morphogenesis during ontogenetic development. Size-frequency analyses show that the material from the Fanan rivulet in Jamtland, central Sweden, represents a natural population dominated by juveniles, whereas the material from Gislovshammar in Scania, southern Sweden, has been sorted during postmortem transport. Three stages of protaspid development can be traced and defined as well as all subsequent ontogenetic stages for the cephalon, hypostome and pygidium. The early meraspid pygidium has a pronounced larval notch, which persists, though becoming progressively less distinct in later meraspides. The number of axial rings in the transitory pygidium increases throughout meraspid development until a third and final thoracic segment is liberated. During ontogeny the articulating half-rings are strongly developed, and both meraspides and holaspides were capable of full sphaeroidal enrollment and outstretched postures. The hypostome undergoes some dramatic modifications; in M0 the anterior margin is axe-shaped, by M1 the area of attachment greatly decreases and the hypostome becomes more elongated and pear-shaped, before attaining its adult form, which has an overall resemblance to that of polymerid trilobites. During ontogeny, the hypostome changes from a conterminant attachment to a natant condition, thereby mirroring hypostomal evolution within trilobites generally. The morphology, ontogeny, enrollment, hypostomal development and the presence of calcified protaspides suggest polymerid rather than agnostoid affinities of the eodiscoids.

nae and a conspicuous pygidium with two short, well-developed digitiform anal cirri. These two new species constitute the first report of polychaetes living in symbiosis with chemosymbiotic bivalves in the south-eastern Pacific.

The genus Ovatoryctocara Tchernysheva, 1962, and its key species Ovatoryctocara granulata Tchernysheva, 1962, are revised. Ovatoryctocara granulata occurs near the base of the Ovatoryctocara Zone and ranges up into the lower portion of the Kounamkites Zone in the Siberian Platform. O. granulata also appears in southeastern Guizhou, South China, but O. granulata in northern Greenland may represent an indefinite species. Specimens of Ovatoryctocara from Newfoundland cannot be identified to species level. Specimens including two cranidia and three pygidia from the lower part of the Aoxi Formation at Yaxi Village, Shizhu Town, eastern Tongren, northeastern Guizhou, were previously assigned to O. granulata, which is now reassigned as a new species O. yaxiensis sp. nov. It bears the following main features: glabella club-shaped, slightly expanded medially, with four pairs of lateral furrows, of which S1–S3 are triangular pits, S4 is shallow, connecting with axial furrow; shorter palpebral lobe situated a little anterior to the midway of facial suture across the fixigenae, longer posterolateral area (exsag.); semielliptical pygidium consisting of seven axial rings with a terminal piece and with eight pairs of marginal tips giving a sawtooth-like shape of the lateral margins in dorsal view. Although O. granulata is a widely distributed species, the FAD of O. granulata, for a global stage boundary, still has some disadvantages. First, its distribution is not as wide as that of Oryctocephalus indicus (Reed, 1910). Second, specimens of O. granulata are common only in Siberia. Third, the stratigraphic range of the species has not been studied in detail in all continents. Fourth, O. granulata is relatively small and easily affected by post-burial distortion. Last but not least, there exists obviously a facies change between the Ovatoryctocara Zone (lower Amgan Stage; deeper water facies) and the underlying Anabaraspis splendens Zone (Toyonian Stage; shallow water facies) in Siberian Platform. Nevertheless, the stratigraphic correlation utility of O. granulata in South China and Siberia is discussed.

The Scarabaeinae of the Indonesian island of Sulawesi have always been marginally treated, the only two directly relevant synopses being Boucomont (1914) and Balthasar (1963a, b), the latter author leaning heavily on his French predecessor as far as Sulawesi taxa are concerned. This applies also to the major ingredient of the subfamily, the subcosmopolitan genus Onthophagus Latreille, 1802. Splitting up this megagenus (ca 2000 named species, depending on one’s opinion on subgenera, see below) into smaller, regionally limited operational units – only few wellfounded subgeneric names being available – has been common practice to make its complex taxonomic diversity more manageable. Recently we published the first of a series of reviews on Sulawesi Onthophagus groups (Krikken & Huijbregts 2008). In the present review we treat the Sulawesi species having – in both sexes – a pair of unconnected, upright paraocular protrusions, and unite them in what is here called the Onthophagus lindu group. In this group ten species are recognized, all new, which are keyed, described, and illustrated below. Seven species with paraocular protrusions from Sulawesi are not included in the O. lindu group since one or more of the following attributes, absent from the O. lindu group, justify a position in other known groups: (1) paraocular protrusions connected by a distinct ridge or lamina; (2) ridge delimiting pygidium from propygidium absent; (3) base of the first elytral interstria with swelling; (4) eye foramina (in full-face view) very broad, Sulawesi Onthophagus with paraocular protrusions: ten new species, with a key (Coleoptera: Scarabaeidae: Scarabaeinae)

The early Cambrian Chengjiang biota in Yunnan, southern China has yielded many articulated exoskeletons of the spiny redlichiid trilobite Eoredlichia intermedia, of which some have their appendages exceptionally well preserved. Both of the paired uniramous antennae of a medium-sized holaspis consist of 46–50 short segments (articles), each of which bears a fine spine near its inner edge. Behind the antennae there are twenty-one pairs of biramous limbs: three pairs are situated underneath the cephalon, one pair underneath each of the fifteen thoracic segments, and probably three pairs underneath the small pygidium. The endopod consists of a broad basis and seven podomeres, of which the last is divided into three terminal spines. The exopod is blade-like, and according to one interpretation, is dorsally hinged to the basis of the endopod; an alternative suggestion being that both the endopod and exopod are split from the basis, the latter being independent and not forming part of the endopod. The exopod has a prominent anterior rim, and possesses about forty long filaments along the posterior margin, and short setae along the rounded distal lobe. The basic appendage features of the redlichiid trilobites, and likewise the gut, are comparable to those of other known Cambrian polymerid trilobites that belong to more distantly related clades.

A new spionid polychaete, Malacoceros jennicus, is described from the crater of a hydrothermally active submarine volcano near Grenada in the Caribbean, Lesser Antilles Arc region. The new species is characterized by having prominent lateral frontal horns on the prostomium, paired eye spots, neurosetae that include simple capillaries, inferior sabre setae, numerous bidentate hooded hooks, and a pygidium bearing four digitiform anal cirri surrounding the anus. Worms were collected in regions of diffuse venting with a suction sampler deployed on an ROV. Specimens representing a range of sizes including sexually mature and reproductive individuals containing eggs and sperm were found. Stable isotope analysis of 13C and 15N indicate mixed feeding on photosynthetic and hydrothermal vent source material.

Adults ofPseudopolydora rosebelaesp. nov. inhabit silty tubes on muddy bottoms in shallow water in southern Brazil, states of São Paulo and Rio de Janeiro. They are rare and extremely delicate, attaining 20 mm long for 55 chaetigers. The worms are distinctive by their colourful yellow and black pigmentation on the anterior part of body and palps, prominent transverse hood on the dorsal anterior edge of chaetiger 3, and lack of coloured respiratory pigment in blood. Of 12 examined individuals, all were females. Oogenesis is intraovarian; oocytes develop from chaetigers 14–15 to chaetigers 24–36. Recently laid oocytes were about 150 μm in diameter, with embryos and developing larvae found in capsules inside female tubes in March–June. Broods comprised up to 23 capsules with 400 propagules. Capsules were joined to each other in a string and each attached by a single thin stalk to the inner wall of the tube. Larvae hatched at the 4-chaetiger stage and fed on plankton. Pelagic larvae are unique amongPseudopolydorain having large ramified mid-dorsal melanophores from chaetiger 3 onwards. Competent larvae are able to settle and metamorphose at the 15-chaetiger stage, but can remain planktonic up to 18 chaetigers. They have one pair of unpigmented ocelli and three pairs of black eyes in the prostomium, unpaired ramified mid-dorsal melanophores on chaetiger 1 and on the pygidium, ramified lateral melanophores on chaetigers 5–10, prominent yellow chromatophores in the prostomium, peristomium, on dorsal and ventral sides of chaetigers and in the pygidium. Branchiae are present on chaetigers 7–10, and gastrotrochs are arranged on chaetigers 3, 5, 7 and 12. Provisional serrated bristles are present in all notopodia, and hooks are present in neuropodia from chaetiger 8 onwards. Two pairs of provisional protonephridia are present in chaetigers 1 and 2, and adult metanephridia are present from chaetiger 4.

All recent work has assigned the genera Cheilocephalus Berkey, 1898 and Oligometopus Resser, 1936 to the family Cheilocephalidae Shaw, 1956, a small group of largely Laurentian trilobites from the Late Cambrian Steptoean Stage. New data from Nevada and Newfoundland indicate that the Cheilocephalidae is polyphyletic and that Cheilocephalus and Oligometopus are allied with different superfamilies. The Cheilocephalidae, which includes Cheilocephalus and Pseudokingstonia Palmer, 1965, is assigned to the Dameselloidea Kobayashi, 1935, and a pygidium from the Cow Head Group of Newfoundland indicates that Cheilocephalus first appears in the late Marjuman. The Dameselloidea disappeared on other continents at an extinction interval at the end of the Marjuman, but Cheilocephalus survived and underwent modest diversification in Laurentia in the succeeding Steptoean Stage. Oligometopus is transferred to the leiostegioidean family Leiostegiidae Bradley, 1925. The genus shows closest affinities with such Gondwanan genera as Idamea Whitehouse, 1939, and its appearance in mid-Steptoean strata records immigration of the Leiostegoidea into shelf-margin environments of Laurentia.

In this paper one new genus and three new species Bellimordella capitulifera gen. et sp. nov., B. longispina gen. et sp. nov., B. robusta gen. et sp. nov., all placed within the subfamily Praemordellinae of the family Mordellidae, are described and illustrated. The specimens were recovered from the Yixian Formation of Beipiao City, Liaoning Province, NE China. The key to the species of this new genus is given. Furthermore, we discuss evolution of the pygidium and possible food for those ancient mordellids.

This study deals with the Dasyvalgus species known from South India, Sri Lanka and Taiwan. A new species, Dasyvalgus schulzei sp.n. from the Anamalai Hills in Southern India, is here described and illustrated. Its relationship to the other Dasyvalgus species is given. Xenoreoderus occidentalis Arrow, 1910, is transferred to Dasyvalgus. A key to the identification of the male Dasyvalgus of South India and Sri Lanka is presented. Another new species, Dasyvalgus sauteri sp.n. from Taiwan, is described and a key to the species distributed on the island is given. A lectotype of D. formosanus Moser is designated. The parameres of Dasyvalgus kanarensis Arrow, Dasyvalgus taiwanus Miyake and Dasyvalgus formosanus Moser as well as the pygidium of the latter species are figured for the first time.

The study of previously unidentified material of Spionidae from the Grand Caribbean region resulted in the discovery of five new species in four genera: Boccardia salazari sp. nov., Dipolydora anatentaculata sp. nov., D. contoyensis sp. nov., Polydora quintanarooensis sp. nov. and Pseudopolydora floridensis sp. nov. Boccardia salazari is characterized by two types of major spine in chaetiger 5, one with expanded end bearing cusps without bristles, the other, simple, falcate; D. anatentaculata is characterized by an occipital antenna, and major spines simple, falcate, without subterminal protuberance; D. contoyensis is characterized by six pairs of branchiae on chaetigers 11–16, major spines with two large accessory teeth on concave and convex sides in chaetiger 5, and pygidium small, glandular, cuff‐shaped with dorsal notch; Polydora quintanarooensis is characterized by two eyes, modified spines of chaetiger 5 falcate, with a lateral tooth, and gizzard‐like structure present at segment 16, and Pseudopolyd...

The Chrysobothris femorata (Olivier, 1790) species group is currently comprised of five species from North American localities north of Mexico. One previously described and synonymized species, C. quadriimpressa Gory & Laporte, 1837 is resurrected. In addition, neotypes are designated for C. femorata , C. rugosiceps Melsheimer, 1845 and C. viridiceps Melsheimer, 1845 as the type specimen for each of these species is lost. A key is included to facilitate identification of the 12 species, 11 of which have deciduous trees as hosts and one species from woody goldenrod, Chrysoma pauciflosculosa (Michx.). Six new species of Chrysobothris are described herein: C. mescalero, C. seminole, C. wintu, C. comanche, C. caddo, and C. shawnee . The new species are each fully described and the elytra, male genitalia, and female pygidium (8 th abdominal tergite) of all species are illustrated.

(Grube, 1870), a sabellid polychaete, were submitted to different anaesthetization and fixation procedures: a control group with living animals, immediate fixation with formaldehyde 4% (FO), ethanol 100% (AE) or ethanol 70% (ET), and anaesthetization with magnesium chloride (MC), refrigeration (RE), menthol crystals (ME) or freshwater (FW). Twenty-three morphometric variables of the body were measured and analysed with size-free multivariate statistics. The first three canonical variables explained 80% of the variation, being significantly correlated with 19 measured variables, mainly those related to the shape of the body setigers. Besides morphometric variables, some morphological characters commonly used in polychaete taxonomy also presented variation among fixation procedures, such as the release of parapodial elements and the integument. Among the procedures tested, AE, ET and FW were the best in approximating the shape of living specimens. Specimens submitted to FO and RE had shorter, wider, and thicker bodies, whereas those submitted to MC and ME were more elongated, narrower, flatter, and had wider pygidia than living specimens. Lengths of branchial crown, setiger 1 and pygidium seem to be the most informative morphometric characters for taxonomic purposes since they did not present deformation due to fixation procedures.INTRODUCTIONSystematists are often interested in quantifying morpho-logical differences among species, conspecific populations, or ontogenetic stages (Strauss & Bookstein, 1982). Body shape represents an important set of animal characters that are used for several purposes, such as studies in taxonomy, ecology, evolution, growth and morphological abnormali-ties (Rohlf & Marcus, 1993; Lestrel, 2000). Therefore, mor-phometrics became a fundamental tool to assess shape and shape variation in biology (Rohlf, 1990).Numerous taxonomic studies have used multivariate morphometrics successfully to analyse differences among species and even among populations (e.g. Debuse et al., 2001; Jordaens et al., 2002; O’Reilly & Horn, 2004). However, there are few morphometric studies on polychaete worms, such as Mackie (1984), Fauchald (1991), Sigvaldadottir & Mackie (1993) and Martin et al. (2003). For soft-bodied animals or structures, body shape depends on the degree of relaxation (Gustus, 1972). Hence, the measurements of some soft-bodied animals or structures after fixation may not reflect their real shape, due to deformation. Consequently, morphometric differences observed among groups of animals submitted to different methods of fixation and/or anaesthetization could be merely methodological artefacts (Howe, 2002).The effects of fixatives on body shape are difficult to predict due to variability related to the type and concentration of fixative, life stage, life habit and other factors (Sagnes, 1997). Methodological artefacts owing to anaesthetisation and fixation procedures are noticeable when comparing material from different origins, such as museum and fresh material. Even for museum specimens, the lack of information concerning anaesthetization and prior fixation methods could restrict comparisons among specimens when shape information is necessary.Experimental studies on potential morphological modifications related to methods of fixation are still scarce for most animal groups (Fowler & Smith, 1983; Kruse & Dalley, 1990; Quinonez-Velazquez & Chaumillon, 1996; Sagnes, 1997; Jordaens et al., 2002) and absent in polychaetes. Mostly, studies regarding anaesthetization methods are more engaged in the maintenance of living animals for transportation rather than on shape recovery of dead or fixed animals (Heasman et al., 1995; White et al., 1996; Bower et al., 1999).In the present study, the effects of different methods of anaesthetization and fixation on morphometric variables were assessed in the polychaete marine worm

Despite being reported widely from basal Sunwaptan strata, Dellea? punctata Palmer, 1965, is known from only four figured cranidia. New material from Nevada, Utah and Oklahoma, including librigenae and pygidia, indicate that specimens assigned previously to D.? punctata are best interpreted as a complex of related species that are assigned to a new genus, Bartonaspis. This genus is diagnosed by punctate sculpture, a pair of fossulae at the anterior corners of the glabella, and by a subelliptical pygidium with a short convex axis composed of three segments. Three new species, B. fredericksoni, B. palmeri, and B. wilsoni, are described and B. punctata (Palmer) is restricted to its holotype. Bartonaspis is known only from the Irvingella major Zone, and its first appearance defines the base of the Sunwaptan Stage. Dellea Wilson, 1949, is related to Bartonaspis and is differentiated from it primarily by pygidial morphology. In contrast to Bartonaspis, the pygidium of Dellea has an axis composed of four, rather than three, segments and a narrow concave border.

Seven species of the marine enchytraeid genus Grania Southern, 1913 are described from sediments sampled during the 2003 International Workshop on the Marine Flora and Fauna of Esperance Bay and the Recherche Archipelago, on the southern coast of Western Australia. Two species are new to science, the euryhaline Tasmanian G. dolichura Rota and Erseus, 2000 represents a new record for the state, and the remaining four species were known from other parts of Western Australia. Grania quaerens sp. n. is recognized by having a high chaetal index (= 5 short chaetal foot), small coelomocytes, penial apparati with long whip-like terminal stylets, conspicuous spermathecae with ectally bulbous ducts, and ectally granulated ampullae housing sperm rings in their ental region. Grania sperantia sp. n. is readily distinguishable by the complete lack of lateral chaetae, a multiple-banded pattern of the clitellum, extremely long sperm funnels, and the intrasegmental location of the spermathecal pores. The latter new species and four others in the collection (G. bykane Coates, 1990, G. crassiducta Coates, 1990, G. dolichura, and G. ersei Coates, 1990) are remarkable in possessing the head organ, a sensory structure unique to Grania that was not noted previously in Western Australian species. When considering the whole genus, the geographic pattern of the head organ appears southern-centred: of the 17 species of Grania reported to possess it, as many as 13 inhabit the southern latitudes. The seventh species of the Esperance collection, G. vacivasa Coates and Stacey, 1993, is notable for the kind of items found in its gut and the unusual appearance of its pygidium.

Two species of Tachygonidius are described from the Atlantic coast of Brazil: T. fluminensis sp. nov. (type locality: Restinga da Marambaia, Rio de Janeiro) and T. paulensis sp. nov. (type locality: Boa Esperança do Sul, São Paulo). The former species develops as a leaf miner in a species of Ochnaceae. Sexual dimorphism related to the form of the declivous portion of pygidium, deeper in male and less deep in female, is reported for the first time in T. fluminensis. A key for the four known species of Tachygonidius is provided.

We herein report a fossilized polychaete annelid, Guanshanchaeta felicia gen. et sp. nov., from the Lower Cambrian Guanshan Biota (Cambrian Series 2, stage 4). The new taxon has a generalized polychaete morphology, with biramous parapodia (most of which preserve the evidence of chaetae), an inferred prostomium bearing a pair of appendages, and a bifid pygidium. G. felicia is the first unequivocal annelid reported from the Lower Cambrian of China. It represents one of the oldest annelids among those from other early Paleozoic Lagerstätten including Sirius Passet from Greenland (Vinther et al., Nature 451: 185–188, 2008) and Emu Bay from Kangaroo island (Parry et al., Palaeontology 57: 1091–1103, 2014), and adds to our increasing roll of present-day animal phyla recognized in the early Cambrian Guanshan Biota. This finding expands the panorama of the Cambrian ‘explosion’ exemplified by the Guanshan Biota, suggesting the presence of many more fossil annelids in the Chengjiang Lagerstätte and the Kaili Biota. In addition, this new taxon increases our knowledge of early polychaete morphology, which suggests that polychaete annelids considerably diversified in the Cambrian. Electronic supplementary material The online version of this article (doi:10.1007/s00114-015-1285-4) contains supplementary material, which is available to authorized users.

Well preserved parts of Telephina bicuspis described from dark limestone concretions of Llanvirn age, in the Oslo Region, Norway, exhibit many features associated with a pelagic life style for this trilobite. The free cheek has a huge holochroal eye with perfectly square lenses and a long genal spine which is unique in being directed vertically upwards rather than ventrally. Additional spines include an occipital spine, the macro spine on the rachis of the 6th thoracic segment and paired spines from each of the two rachial rings of the pygidium. These spines may have had a hydrodynamic and stabilizing function. The genal spine could have had a sensory function or been a dimorphic feature. The hypostoma is figured for the first time and a reconstruction is given for the complete exoskeleton.

A new species of Polygordiidae, Polygordius jouinae sp. nov., along with its distribution, habitat, and reproduction is described. Polygordius jouinae sp. nov., the first North American Polygordius, is a dominant member of macrofaunal communities on the inner continental shelf, and in bays and harbours from Massachusetts to southern New Jersey. It is distinguished from most other Polygordius species by its non-inflated, heavily ciliated pygidium, absence of pygidial glands, and a conical (rather than rounded) prostomium. The 18S SSU rDNA from P. jouinae sp. nov. was sequenced and represents the first named Polygordius species with a DNA reference in GenBank. Spearman rank correlation of sediment grain size with density of P. jouinae sp. nov. at a New Jersey site showed that density was significantly (PP. jouinae sp. nov. is an important macrofaunal species given its widespread distribution and its fidelity for coarse sand habitats. Thus, its relative abundance may be useful as an indicator of changing sedimentary conditions.

Evidence about ontogeny of the miomerid trilobite groups Eodiscina and Agnostina is summarizied and unified in terminology. There appear to be strong similarities in the juvenile development of both suborders. They both have prolonged meraspid degree 0 and 1 stages, including two to six molts. The degree 0 stage tends to shorten from six to four molts in Eodiscina, and probably from three to two in Agnostina, while degree 1 tends to lengthen whithin Eodiscina. During degree 0 and 1 moltings, none of the segments are liberated into the thorax, and one segment is added into the transitory pygidium. The liberation of thoracic segments marks the beginning of the meraspid degrees and the holaspid period. This developmental scenario contrasts with that of the classical polymerid—in the latter case, the liberation of thoracic segments is accompanied by insertion of new segments in the transitory pygidium. In the Eodiscina, during meraspid degree 2, the polymerid scheme is present.

Paradoxides jemtlandicus Wiman, 1903, from strata near the boundary between the oelandicus Stage and the paradoxissimus Stage of the middle Cambrian in central Sweden, is redescribed using more complete specimens than were known hitherto. It is one of the earliest species of the genus Paradoxides sensu stricto, and though the form of the dorsal exoskeleton and the transverse shape of the hypostome are normal for the genus, compared with typical Paradoxides such as P. paradoxissimus it shows features that can be regarded as primitive. These features include the shape of the glabella, the lack of strong fusion of the rostrum with the hypostome, the acute inner spine angle of the librigena and the short pygidium; they are reminiscent of some species of Hydrocephalus.

A paper by renowned eighteenth century Swedish naturalist, Carl Linnaeus, published in 1759, contained the first fossil reconstruction of what was later called a trilobite and was an early milestone in invertebrate palaeontology. The earliest work devoted solely to Palaeozoic trilobites (Arthropoda), Linnaeus' “Petrificatet Entomolithus paradoxus …” delved, in great detail, into the classification of this genus intermedium between known marine crustaceans. Linnaeus depicted on two folding, copper-engraved plates the fossil animal (comprising the cephalon, thorax and pygidium with pleural spines and furrows) from the best-preserved material in limestone thus far recovered, including an erroneous interpretation of antennae in relation to this Swedish material – however, confirmed in the latter part of the nineteenth century with well-preserved Silurian trilobites.

Two sympatric species earlier differentiated by means of starch gel electrophoresis within Polydora cf. ciliata from the Sea of Japan are here identified and their adult and gamete morphology are described and illustrated. One species bores into various shells and coralline algae while the other bores only into gastropod shells occupied by hermit crabs. Both species have the prostomium anteriorly incised to rounded, 0–4 eyes, the caruncle extending to the end of chaetiger 2, heavy falcate spines of chaetiger 5 with a lateral tooth, branchiae beginning on chaetiger 7, and the pygidium disc-like to cup-shaped with a dorsal gap to only an incision. Males of the two species differ in aggregations of spermatids. The more opportunistic borer, which occurs in a greater number of hosts, always has 8-cell-aggregations of spermatids. These worms are here referred to as P. calcarea. The other species, in which spermatids are always interconnected in tetrads, is here described as a new one, P. manchenkoi sp. nov. The two species also differ in that prominent brownish-yellow pigment appears on the posterior segments after fixation in formaldehyde in P. manchenkoi sp. nov. but not in P. calcarea. It is suggested that spermatid aggregates and spermatozoan morphology be included in future polychaete descriptions when possible.

Prionospio patagonica inhabits temporary silty tubes intertidally and shallow subtidally in brackish water estuarine environments in southern Chile. The species is gonochoristic with the female:male ratio being close to 2:1. Females and males release gametes into water. Pelagic planktotrophic larvae were caught in the plankton in the River Valdivia estuary in October–November and in March. Development of the adult morphology is described and illustrated beginning from the 2-chaetiger larval stage. One pair of lateral eyes first appears in the early larva, and shortly after that the right median eye develops. The left median eye appears after settlement, in juveniles with 10–11 segments. Developed 6-segment larvae have three dark red eyes, short palps of equal length, no nototrochs, one pair of small cells with grasping cilia on the pygidium, a small ciliated pit, gastrotrochs on segments 2–6, long serrated bristles in notopodia, adult capillaries in noto- and neuropodia on segments 2–5, single hooks in both rami on segment 6, and one pair of provisional papillae on the pygidium. A ventral buccal bulb is present below the short oesophagus and two pairs of provisional protonephridia are present in segments 1 and 2. The wall of the anterior part of the midgut has a characteristic brown pigment. The 6-segment larvae, about 400 μm long, settle and undergo gradual metamorphosis. In adults, hooks are gradually lost from noto- and neuropodia but sabre chaetae appear from segment 7 and retain their anterior position as growth proceeds. Up to 14 pairs of cirriform branchiae develop from segment 2, and three adult cirri appear on the pygidium. The afferent and efferent arms of each branchial blood loop are interconnected by capillary loops. A greenish heart body is present in the main dorsal blood vessel in anterior segments. Up to 21 pairs of metanephridia develop in anterior sterile segments, beginning with segment 4. Transparent gonoducts are present in fertile segments.