Three nephromixial patterns in polychaete species currently assigned to the genus Pista (Annelida,Terebellidae)

A comparative morphological study of nephromixial systems in three Californian terebellid polchaetes currently assigned to the genus Pista shows that P. fimbriata has all attributes of the generic type, but that P. pacifica and P. elongata differ markedly. The features of typical Pista include (among others): two pairs of usually unequal branchiae, long-handled anterior uncini (hooks) of crested avicular (bird-head-like) from, muddy, unornamented tubes, one pair of anterior excretory nephromixia (ENMX), and two pairs of separate thoracic reproductive nephromixia (RNMX) with genital papillae on segments VI and VII. A review indicates that P. fimbriata shares these typical features with practically all adequately described Pista species. However, P. pacifica and P. elongata possess three pairs of branchiae, long-handled uncini of distinctive crochet-like form, and membranous tubes with apertural hoods. Both have two pairs of ENMX, the first supplied by one pair of ciliated renal funnels, the second by two pairs of such funnels. But they differ in their RNMX: P. pacifica has three pairs of complex RNMX, those on each side united by a common duct. P. elongata has 11–13 pairs of simple RNMX united by common ducts. Although these species do not fit into Pista, no genus has been found to accommodate them. Generic placement is complicated by the fact that no instances of intra-generic nephromixial variation have been reported in the Terebellidae, although inter-generic variation is well known. If they are congeneric, this would be the first example of intrageneric RNMX variation in Terebellidae. But if assigned to separate genera or subgenera on the basis of their RNMX, their similarity of anterior uncini might be attributable to parallel or convergent adaptation to life in comparable tubes. More evidence, including molecular analysis, is needed for phylogenetic studies of Terebellidae. © 1992 Wiley-Liss, Inc.     

Structure and formation of the uncini in Pectinaria koreni,Pectinaria auricoma (Terebellida) and Spirorbis spirorbis (Sabellida): inplications for annelid phylogeny and the position of the Pogonophora

Setation is an important taxonomic character of the Annelida. Within this taxon, Terebellida and Sabellida both have transverse rows of short, apically toothed setae which are situated inside the neuropodial rim. The apical spines are curved and their tips point anteriorly. These setae are termed uncini. In the terebellidans Pectinaria koreni, Pectinaria auricoma and in the sabellidan Spirorbis spirorbis, these setae arise from a follicle which consists of a chaetoblast and two follicle cells. The special structure of the uncini is a result of temporal modifications of the actin-filament system of the chaetoblast and changing spatial interactions between the chaetoblast and the follicle cells during the formation of these setae. Once the uncinus is formed, the microvilli are withdrawn and electron-dense material is deposited in the remaining canals. The microvilli are replaced by short processes of the chaetoblast, and the actin-filament system is replaced by a system of intermediate filaments which help to mechanically attach the uncinus to the follicle. Such uncini are also described in both pogonophoran groups, the Perviata and the Obturata (Vestimentifera). In several structural details they correspond to those of the species investigated in this paper, so that the hypothesis of a homology of the uncini seems to be justified. This hypothesis leads to the conclusion that uncini evolved in the common stem lineage of Pogonophora, Terebellida and Sabellida, implying a monophyletic origin of these three taxa. The uncini are compared to the hooked setae of the Arenicolida, Maldanida and Psammodrilida, which are also aligned in transverse rows inside the neurophodial rim. Hooked setae and uncini are hypothesized to be homologous. It, therefore, can be concluded that Arenicolida, Maldanida and Psammodrilida are closely related to the monophylum consisting of Terebellida, Sabellida and Pogonophora, and that these six taxa share a common ancestor, which evolved transverse rows of setae with apically curved spines and a formative site lateral to the edge of the neuropodial rim. According to the phylogenetic relationships proposed here, the Pogonophora are a subordinate taxon within the Annelida.     

Phylogenetic analysis of the Amphitritinae (Polycnaeta: Terebellidae)

The phylogenetic relationships of the Amphitritinae (Polychaeta: Terebellidae) were studied using parsimony analysis of 22 external morphological characters. To choose outgroups to polarize the characters, I carried out a preliminary analysis of the relationships of the four terebellid subfamilies and the Trichobranchidae. The single most parsimonious tree from the analysis supports monophyly of the Terebellidae by the presence of ventral glandular shields. However, this character is homoplasious within the Terebellomorpha, and further evaluation of the Terebellidae is recommended. Artacama and Thelepus were chosen as outgroups for the analysis of amphitritine genera. The generic level analysis yielded seven equally parsimonious trees, which are consistent in their topologies except for the relationships among seven genera in one large clade. In all trees, Artacama is the sister taxon to a large clade within the Amphitritinae; the Artacaminae is therefore synonymized with the Amphitritinae, which is diagnosed by the presence of double rows of uncini. Within the Amphitritinae, the status of several monotypic genera is questioned; plesiomorphic character states indicated by the analysis are discussed. The results presented are offered as working hypotheses of the relationships among amphitritine genera. The large number of homoplasies indicated by the analysis emphasizes the need to further evaluate these hypotheses using additional characters. With a robust phylogenetic hypothesis of amphitritine relationships, a re-classification of the group based on apomorphic character states can be undertaken, and questions regarding the evolution of morphological characters, reproductive modes, or biogeographical patterns can be properly addressed.     

Early scale development in Heterodontus (Heterodontiformes; Chondrichthyes): a novel chondrichthyan scale pattern

In two species of Heterodontus, H. portusjacksoni and H. galeatus, the first scales to develop form two opposing rows along the caudal fin axis on both the left and right sides of the fin. The opposing rows originate from an initial scale located on either side of the posterior tip of the caudal fin, with subsequent scales erupting in a posterior to anterior direction along the tail axis. These scale rows may strengthen tail movements, providing aeration in the egg case, but are lost later in ontogeny. Development of subsequent body scales shows a more irregular origin and arrangement, from anterior to posterior, to cover the dorsal and ventral lobes of the caudal fin. Although the early developmental pattern of the scale associated with the Heterodontus caudal fin has not been previously described, several chondrichthyan taxa, including chimeroids, likewise possess ordered rows of flank scales early in ontogeny that are subsequently lost. These ordered scales contrast with previous suggestions that chondrichthyan scale development is entirely random. Instead, regulated and sequential development of scales may be a plesiomorphic character for both chondrichthyans and osteichthyans, with the less organized arrangement in later ontogenetic stages being a derived condition within Chondrichthyes.     

Colony Growth Pattern in Electra pilosa (Linnaeus) and Comparable Encrusting Cheilostome Bryozoans

Colony growth pattern is described in E. pilosa, an abundant cheilostome bryozoan commonly found as an epiphyte of Laminaria. Each zooid has 4 potential budding loci—one distal, two lateral and one proximal. The ancestrula buds daughter zooids from all of these loci; the two lateral buds appear first, followed by the distal bud and, after a long delay, the proximal bud. The laterally budded zooids curve inwards as they grow to form a triad with their distally budded sibling zooid. ‘Mature’ multiserial colonies growing on flat substrata consist of a series of radially diverging sectors. Each sector has an axis, generally of 3 parallel rows of zooids, flanked by wings consisting of rows of zooids originating as lateral buds from the section axis which infills the area between the axes. Occasional colonies occur with uniserial or semiuniserial growth patterns. These resemble colonies of the obligatory uniserial species Pyripora catenularia and poorly fed colonies of the related Conopeum tenuissimum, which is normally multiserial like E. pilosa. The ‘composite multiserial’ colonies of E. pilosa differ in several respects from ‘unitary multiserial’ colonies characteristic of most sheet-like cheilostomes, including the well-known Membranipora membranacea. Composite and unitary multiserial growth patterns may have evolved independently from uniserial ancestors.     

Innervation pattern and electric organ discharge waveform in Gymnotus carapo (Teleostei; Gymnotiformes)

The electrogenic organ (EO) of Gymnotus carapo has two main portions: a posterior region consisting of four bilaterally arranged electrocyte rows; and an anterior portion composed of only two. The lateral row (LR) of the anterior portion contains doubly innervated electrocytes with axon terminals from different nerves on their rostral and caudal faces. The LR is continuous with the most dorsal row of the caudal region. This row also contains doubly innervated electrocytes. The medial row (MR) electrocytes of the anterior region and ventral rows of the caudal region are exclusively caudally innervated. All caudal faces of the anterior or abdominal region are supplied by two nerves which originate from spinal roots VIII to XXI. Roots I to VII give origin to pure rostral nerves whose electromotor axons terminate on the rostral surfaces of the first seven LR electrocytes. A given doubly innervated electrocyte is supplied on its caudal face by a nerve originating several segments (usually seven) posterior to the spinal root supplying its rostral face. Transections of the spinal cord at the level of root VIII isolate the activity of the rostral surfaces of the first electrocytes. The EO discharge (EOD) then appears as a head negative deflection which arises from abdominally located electrocytes. Its monophasic character reveals that the activity remains restricted to the rostral electrocyte surfaces. Damage of the abdominal portion of the EO abolishes the first negative deflection of the normal pulse. Transections of the spinal cord at the level of root XXI isolate the activity of the whole abdominal portion of the EO. Since both doubly and singly innervated electrocytes remain active, the EOD appears biphasic. Comparative studies have shown that the EOD of Hypopomus sp. lacks any early negative wave and correspondingly all its electrocytes are exclusively caudally innervated.     

The genera <Emphasis Type="Italic">Bispira</Emphasis> Krøyer, 1856 and <Emphasis Type="Italic">Stylomma</Emphasis> Knight-Jones, 1997 (Polychaeta,Sabellidae): systematic revision,relationships with close related taxa and new species from Australia

A cladistic analysis undertaken to test monophyly of Bispira and Stylomma (including new species of both genera) and to ascertain relationships with related taxa reveals that Bispira is paraphyletic without the inclusion of Pseudobranchiomma and Branchiomma, and that Stylomma is monophyletic due to the presence of a structure on the dorsal basal flanges which has never been described before. New species of Bispira and Stylomma from Australia are described. Bispira serrata n. sp., differs from the other species of the genus in having serrated radiolar flanges whereas all other Bispira species have smooth flanges or lack them, and also by the shape of the thoracic and abdominal uncini, which are short-handled and have 2–3 rows of big teeth above the main fang, instead of having medium-sized handles and small, numerous rows of teeth over the main fang. Stylomma juani n. sp., is characterized by the presence of serrated radiolar flanges and unpaired compound eyes along the branchial radioles, whereas other species in the genus have smooth radiolar flanges and subterminal compound radiolar eyes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: K. Martens     

ON THE ANATOMY AND MORPHOLOGY OF LATERAL BRANCH SYSTEMS OF ARCHAEOPTERIS

A large collection of specimens, consisting primarily of Archaeopteris macilenta, forms the basis for an anatomical and morphological study of the lateral branch systems. Emphasis is placed on the main axes of the branch systems, which are found to be characterized by a variable number of orthostichies of lateral appendages, ranging from 7 to probably 11. The number of orthostichies seems to correspond to the number of ribs of the stele. There may be an equal number of rows of leaves on either side of the plane in which the two rows of lateral branches lie, or one side may have one or two more leaf orthostichies than the other. In all specimens for which both part and counterpart were available, however, there is a greater density of leaves on one side (hypothesized to be the adaxial) than on the other, caused primarily by an apparent abortion of leaf primordia early in development on the side of least density. Leaves and branches occur in the same ontogenetic spiral and in one specimen comprise a ‘phyllotaxis’ of 2/9 and contact parastichies of 5 + 9. On both the main and lateral axes, leaves are characterized by long, decurren bases which essentially ensheath the axrs on which thay are borne. Phyllotaxis and the correlated stelar form apparently vary considerably within Archaeopteris. It is suggested that Archaeopteris was more closely related to the coniferophytes than to any other gymnosperms.     

Chaetal arrangement provides no support for a close relationship of Sabellidae and Sabellariidae (Annelida)

Sabellid and sabellariid polychaetes are regarded as sister groups in a number of recent phylogenetic analyses. This is based mainly on a shared specific arrangement of chaetae referred to as chaetal inversion. Remarkably, the uncini have a notopodial position in the abdomen, whereas capillary chaetae occur in the neuropodia in both taxa in contrast to the situation in putative relatives. However, in sabellids uncini and capillary chaetae change their position completely at the border between thorax and abdomen, whereas uncini are missing in the parathorax of Sabellariidae. Due to this difference the significance of the chaetal inversion for systematics has been subject to discussion for years. Serial semithin sections of parapodia of the Sabellidae Sabella pavonina, Branchiomma bombyx, Fabricia stellaris, and of the Sabellariidae Sabellaria alveolata were studied in order to obtain detailed information on their chaetal arrangement and sites of chaetal origin. SEM investigations and computer-aided 3D-reconstructions provide deep insight into the spatial organization of the rami. Though differing externally, the principal chaetal arrangement and the location of the formative sites turned out to be almost identical within the species of Sabellidae. Most chaetae are aligned in straight transverse rows with a dorsal site of origin within neuropodia and a ventral one in notopodia as is common in sedentary polychaetes. Semicircular and spiral arrangements are revealed to be modified transverse rows. Only in thoracic notopodia does an additional dorsocaudal formative site form distinct rows. The chaetal inversion in Sabellidae is additionally characterized by an abrupt change of capillary chaetae and uncini along with a sudden change of the parapodial morphology at the border between thorax and abdomen. All chaetae of S. alveolata are aligned in transverse rows with the same location of the formative sites as in sabellids and other sedentary polychaetes. However, in contrast to sabellids the chaetae are not inverted across a parathoracic abdominal border. Moreover, there is no inversion of the parapodial structure from parathorax to abdomen and the neuropodial chaetal composition changes gradually from parathorax to abdomen. The chaetal arrangement in Sabellariidae thus cannot be described as inverted along the body-axis as in Sabellidae. Evolutionary steps implied by the assumption of an inverted chaetal pattern in a supposed common ancestor are discussed. It is concluded that the specific chaetal arrangement of Sabellidae and Sabellariidae arose independently and therefore provides no support for a sistergroup relationship of sabellids and sabellariids.     

Ultrastructure of opisthosomal chaetae in Vestimentifera (Pogonophora, Obturata) and implications for phylogeny

Schulze, A. 2000. Ultrastructure of opisthosomal chaetae in Vestimentifera (Pogonophora, Obturata) and implications for phylogeny. — Acta Zoologica (Stockholm) 82 : 127–135
The posterior segmented body region of Vestimentifera bears rows of uncini that function to anchor the animal within its tube. SEM studies of five vestimentiferan species reveal intraspecific and interspecific variation in the number of chaetigerous segments and the arrangement of uncini within a given segment. The portion of an uncinus that extends beyond the epidermis comprises two opposing groups of teeth that probably correspond to the capitium and subrostral process of polychaete uncini, and a distinct protuberance between them, interpreted as a rostrum. In Ridgeia piscesae , the uncini are formed by chaetal follicles, consisting of a chaetoblast, a follicle cell and an epidermis cell. The chaetal shaft is elongate and composed of up to 40 hollow cylinders that are invaded at their base by microvilli from the apical part of the chaetoblast. Opisthosomal chaetae in perviate Pogonophora are usually restricted to four per segment and are of a rod-shaped type. It is hypothesized that the rod-shaped chaetae represent reduced hooked chaetae probably derived from a condition such as found in Monilifera. Uncini of Pogonophora, Sabellida, Terebellida and Oweniida are considered homologous but details of chaetal design may be due to functional adaptations and thus do not represent reliable characters for phylogenetic studies on higher taxonomic levels than genera or potentially families.     

Morphological and histological changes of dermal scales during the fish‐to‐tetrapod transition

Witzmann F. (2011). Morphological and histological changes of dermal scales during the fish‐to‐tetrapod transition. —Acta Zoologica (Stockholm) 92 : 281–302. The gastral scales of limbed tetrapodomorphs evolved from the ‘elpistostegid’‐type of scale by an enlargement and differentiation of the articulation facets and a shortening and broadening of the keel. These changes caused a tighter connection between gastral scales within a scale row and a greater overlap between the rows. Dorsal round scales of limbed tetrapodomorphs developed from a gastral scale‐type by an alteration of the ontogenetic pathway. The posterolateral direction of scale rows in ‘elpistostegids’ was retained in the gastral scalation of most limbed tetrapodomorphs, whereas the arrangement of round dorsal scales is modified to a transverse orientation. Both gastral and dorsal scales of limbed tetrapodomorphs consist solely of parallel‐fibred bone with circumferential growth marks. The proportionally larger overlap surfaces of gastral scales and their mode of articulation in the ventral midline indicate that the body of limbed tetrapodomorphs might have been more flexible than that of their finned relatives. The alteration of dermal scales was one of the most rapid morphological changes during the fish‐to‐tetrapod transition. Once established, gastral and dorsal scales were retained as a conservative character in different lineages of basal tetrapods, in both the amphibian and the amniote lineages.     

Phylogeny of western Mediterranean Leptodirini, with an emphasis on genital characters (Coleoptera: Leiodidae: Cholevinae)

Abstract The tribe Leptodirini (Leiodidae: Cholevinae) is one of the largest radiations of Coleoptera in the subterranean environment. Although subjected to systematic and evolutionary studies, the phylogeny remains poorly understood. We assessed the phylogeny of the western Mediterranean lineages (Iberian Peninsula, Pyrenees and Sardinia) based on a cladistic analysis of fourteen characters of external morphology and twenty characters of the male and female genitalia, studied in 182 species belonging to thirty‐nine genera. We tested the monophyly of the traditional two main divisions of the group (infraflagellates and supraflagellates), as well as that of some ‘phyletic series’. The final matrix contained fifty‐eight terminal taxa, twenty‐four of which had different character state combinations. The strict consensus of the sixty most parsimonious trees recovered a monophyletic Leptodirini, but not their separation into infraflagellates and supraflagellates. The supraflagellates formed a paraphyletic group with respect to the infraflagellates (corresponding to our sampled ‘Speonomus’ series), with Notidocharis sister to all other included Leptodirini, and Speonomidius sister to Leptodirini excluding Notidocharis. The series ‘Spelaeochlamys’, including the Sardinian genera but excluding Pseudochlamys, was recovered as monophyletic with weak support. The ‘Quaestus’ series formed a polytomy with Pseudochlamys plus the ‘Speonomus’ series (including Bathysciola), which was recovered as monophyletic with strong support. Speonomus, Bathysciola, Quaestus and Troglophyes were para‐ or polyphyletic. Our results suggested the respective monophyletic origin of the Leptodirini from the Pyrenees (Pseudochlamys plus the ‘Speonomus’ series) and the Mediterranean coast plus Sardinia (series ‘Spelaeochlamys’). On the contrary, the Leptodirini of the Atlantic north coast of the Iberian Peninsula (series ‘Quaestus’ and ‘Speonomidius’) were not monophyletic.     

Pollen morphology of Zehneria s.l. (Cucurbitaceae)

A growing body of experimental data obtained from sporoderm ontogenetic studies led to the appearance of the ‘micellar’ hypothesis. The hypothesis is that the sequence of sporoderm developmental events represents the sequence of self-assembling micellar mesophases, initiated by genomically given physico-chemical parameters, which are then picked up by physico-chemical self-assembly. However, besides morphological evidence, the best proof of this hypothesis would be an experimental modelling of sporoderm-like patterns. The main idea of this study is to remove the influence of the genome, selecting substances and their concentrations for simulations to replace it, and then to trace what ‘pure’ self-assembly is capable of constructing. Our aim in this study was to simulate mainly young structures in sporoderm development, i.e. the glycocalyx and the primexine. Several polysaccharide gels (as a callose substitute) and surfactants (as glycocalyx and sporopollenin monomer substitutes) were mixed at different concentrations and combinations, thermally set and left to condense. A number of patterns were obtained in colloidal solutions in the course of condensation, simulating structures at different stages of exine development. Their structures were observed and analysed with transmission electron microscopy (TEM). Our first experiments on the modelling of biological patterns in vitro have shown encouraging results.     

Ontogeny of the jaws of monogonont rotifers: the malleate trophi of Rhinoglena and Proalides (Ploima,Epiphanidae)

Information on the embryonic development of the malleate trophi in Epiphanidae (Rotifera, Monogononta, Ploima) is presented, based on scanning electron microscopy observations in Rhinoglena fertoeensis, R. frontalis, R. kutikovae, R. tokioensis, and Proalides tentaculatus, to contribute to the understanding of this structure of high evolutionary and functional relevance in Rotifera. The first observable and distinctly sclerotized structures were a double row of median transversal sclerites along the longitudinal axis, wherein the future unci, rostellar scleropili, cristae rami, and basal apophyses became recognizable. Fulcrum and manubria arose subsequently; the fulcrum sclerites were longitudinally ordered in a double layer. The rami chambers developed last as lamellar structures. Unci appeared as separate thin, elongate elements, the primary uncini, developing to uncus plates by transversal growth and apposition of sclerite material on the shafts of the uncini. The heads of the uncini showed their greatest development after fusion of their shafts into uncus plates. The interjacent spaces between the heads functioned as a mold, organizing bundles of sclerites which developed into the uniseriate, zigzag‐shaped cristae rami. The fulcrum attained its definite shape by elongation of the double layer of rod‐shaped sclerites into appressed sclerofibrillae. Manubria became visible as a proximal ridge of sclerites, whereupon a triangular lamella composed of crisscross‐oriented sclerites developed distally, growing out to the manubrial chambers. Ramus chambers originated from two longitudinal amorphous lamellae incorporating the median rami sclerites and closing from distal to proximal; subbasal chambers were formed before the basal chambers.     

Chaetal arrangement in Orbiniidae (Annelida,Polychaeta) and its significance for systematics

The systematic position of Orbiniidae within Polychaeta is still uncertain. In order to provide additional comparative data, we investigated the chaetal arrangement in this family, which is considered valuable for polychaete systematics. Specimens of Scoloplos armiger, Orbinia latreillii, and Pettibonella multiuncinata were examined by SEM and serial sections analysed by computer aided 3D-reconstructions. The obtained data suggest that the chaetal arrangement of Orbiniidae resembles that of other sedentary polychaetes in only a few respects. Transverse rows are only present in the main, anterior part of the chaetal patches of thoracic neuropods. The position of the formative site indicates homology with the transverse rows of several sedentary polychaete taxa. The chaetal patches thus differ significantly from those known in Apistobranchidae. Independent rows with an own caudal formative site, which run along the caudoventral edge of the chaetal patches, resemble the neuropodial ventral longitudinal rows known in Spionidae and related taxa. The abdominal neuropodia of S. armiger and O. latreillii bear longitudinal rows of chaetae. These are reorientated during ontogenetic chaetiger transformation and become the transverse rows of the thoracic chaetal patches. 3D reconstruction of S. armiger revealed that the notopodial chaetal bundles are organized in rows as well. Notopodia and abdominal neuropodia bear deep reaching supportive chaetae. They are the first chaetae formed during neuropodial development and reside dorsally to the longitudinal row of capillary chaetae. Neither position nor structure indicates homology with the supportive chaetae of other sedentary polychaetes. Spionidae and related taxa are thus the only sedentary polychaetes, which specifically resemble Orbiniidae in certain aspects of their chaetal arrangement. Dedicated to Prof. Dr. Wilfried Westheide on the occasion of his 70th birthday.     

Occurrence of early post-hatching Jurassic Nautilida in Normandy, France: palaeobiologic, palaeoecologic and palaeobiogeographic implications

Nautilida are present and sometimes abundant in various sedimentary environments of the Jurassic series (Sinemurian to Kimmeridgian) in Normandy, but in the majority of cases the individuals are mature or at a late stage of ontogenetic development. Comparative analysis of the palaeobiogeographic distribution of Nautilida according to their ontogenetic stages shows that early post-hatching specimens are present only in particular palaeoenvironments which are rarely visible and laterally limited in outcrop. The sedimentologic and palaeoecologic originality of the Carixian, Aalenian and Bajocian deposits bearing these peculiar specimens is related to their palaeogeographic position along the western border of the Anglo-Paris Basin, on the eastern margin of the Armorican Massif, where Jurassic deposits overlie Palaeozoic rocky reefs or shoals. The egg-laying places of the Jurassic Nautilida show palaeoenvironmental features similar to those of the recent Nautilus, having been controlled by the water temperature and the nature and morphology of the sea bottoms. A hard, anfractuous and stable substratum, low sedimentation rates and warm and well-oxygenated marine waters appear to have been required for egg-laying and for the embryonic development of these cephalopods. The discovery of early post-hatching fossil Nautilida is fundamental to a better understanding of their palaeobiogeographic distribution at local to global scales. The stenotopic nature of their egg-laying places explains the segregation between newly hatched individuals and the rest of the population and leads also to a distinction between the notions of ‘dispersal potential’ and ‘settlement potential’ for populations of Nautilida. Three palaeobiogeographic maps show that Nautilida were absent in the Jurassic Boreal realm. This exclusion could be closely dependent on the thermal requirements that govern their reproduction. By comparison, the contemporary occurrence of ammonoids in this realm appears to reflect differences in reproductive strategies.     

Setal Structure, Functions and Interrelationships in Spirorbidae (Polychaeta, Sedentaria)

Typical thoracic setae are composed of longitudinally arranged cylinders ending as teeth at the oblique distal surface. Most thoracic 'blades' are rounded in cross section and show bilateral asymmetry. Some have teeth in oblique rows ("cross striations") on the side adjoining the tube wall. Sickle setae found posteriorly on the thorax have distal portions like abdominal setae. These are flat, each composed mainly of a palisade of spatulate units. On one side cylindrical elements may also be present, in patterns which are characteristic of some taxa. Uncini are made of rows (sometimes single) of shorter cylindrical elements with hooked ends pointing anteriorly. Anterior thoracic hooks are usually blunt, but pointed in the genus Janua. The collar setae assist withdrawal into the tube, the abdominal setae help with emergence, whilst other thoracic setae act as distance pieces, pushing the body to anchor the thoracic uncini against the opposite wall. The other uncini are brought into use by the prehensile abdomen. The flaccid ends of the sickle setae are readily lost in some taxa and their development or loss may reflect unimportant variations in strength of the adjacent abdominal field. Fin and blade setae are not very different from simple setae and the natural grouping of the genera into subfamilies shows that changes from one form of collar seta to another have occurred independently several times.     

Ontogeny of the redlichiid trilobite Eoredlichia intermediata from the Chengjiang Lagerstätte,lower Cambrian,southwest China

A detailed investigation of the morphology and ontogeny of the redlichioid trilobite Eoredlichia intermediata (Bulletin of the Geological Society of China, 3–4, 1940, 333) from the lower Cambrian Yu'anshan Member of the Heilinpu Formation, in Kunming, Yunnan Province, southwest China, is presented. The new material comprises a relatively complete ontogenetic series ranging from the early meraspid to the holaspid period, which reveals more details on morphological variation such as the appearance of bacculae in some holaspid specimens, contraction and disappearance of fixigenal spines, and macropleural spines of the first and second thoracic segments, which are all documented for the first time and can also be used as developmental markers defining ontogenetic phases. Two distinct morphotypes, possibly an expression of intraspecific variation or sexual dimorphism, are distinguished by the morphology of pleural spines of the second thoracic segment in meraspid degree 14 and holaspides. The trunk segmentation schedule of E. intermediata is also discussed and conforms to the protarthrous developmental mode. The distinction of the thoracic region into two portions can be observed during late meraspid and early holaspid periods, which might be considered as a reference for a better understanding on the relationship of tagmosis and growth segmentation among the Cambrian redlichiid trilobites.     

Ultrastructural Observations on the Spermatozoa of Two Temnocephalids (Platyhelminthes)

The spermatozoa of two Temnocephalidae collected in Uruguay, Temnocephala iheringi Haswell, 1893 (Host: Pomacea canaliculata) and Temnocephala axenos Monticelli, 1899 (Host: Parastacus varicosus), were studied with a transmission electron microscope. In both species the spermatozoon is made up of a long sperm body which bears at one extremity two free flagella of the 9+‘1’ flatworm pattern. The sperm body contains the nucleus, mitochondria, dense bodies and parallel, cortical, longitudinal singlet microtubules. Along a part of the sperm body the palissade of the microtubules displays a spiral pattern in transverse sections. A part of the perimeter of the cell is thus lined by two overlapping rows of microtubules. This spiral pattern of the singlets is considered as a synapomorphy of the family Temnocephalidae. The singlet microtubules are interconnected by two kinds of links: tangential links between neighbouring singlets in the same row and radial links between singlets belonging to two rows. The presence of these links suggests that this structure could be a motile system of singlets.     

Minireview: On the homology of the protocoel in Cephalochordata and 'lower' Deuterostomia

Hypotheses regarding the homology of the protocoel in planktonic deuterostome larvae and mesodermal structures in ontogenetic stages of cephalochordates are evaluated. The prevalent ‘classical’ hypothesis describes the protocoel as being homologous with the diverticula of Hatschek, which, on the left side, develop into the preoral pit, subsequently into Hatschek’s pit and groove (in part). This hypothesis is based mainly on the position of Hatschek’s diverticula anterior to the rest of the mesoderm during their enterocoelic origin. It is shown here that during development the mesodermal segment that develops into Hatschek’s nephridium is the most anterior one prior to formation of Hatschek’s diverticula, and this segment assumes an anteriormost position after differentiation of Hatschek’s diverticula. Additional similarities between this segment and protocoels are: (i) presence of endomesodermal cells with podocytic extensions, (ii) excretory function, (iii) relatively early ontogenetic origin, (iv) probable lack of association with nervous structures, (v) probable ectodermal origin of a portion of the canal, and (vi) position relative to the mouth opening. Therefore, homology between the protocoel and the segment that becomes Hatschek’s nephridium is proposed. It is concluded that a glandular structure homologous to the diverticula of Hatschek and anterior to the protocoel/Hatschek’s nephridium is a synapomorphy of notochordates or chordates.