Commensal anomiid bivalves on Late Cretaceous heteromorph ammonites from south‐west Japan

The heteromorph ammonite Pravitoceras sigmoidale from the Upper Cretaceous Seidan Formation (Izumi Group) in south‐west Japan is frequently encrusted by sessile anomiid bivalves. Fossils of P. sigmoidale with anomiids are often concentrated at the top of or just above turbidite sandstones. Projecting retroversal hooks and apertures of P. sigmoidale are usually intact, and some individuals are associated with jaw apparatuses near apertures. Anomiids are found on both sides and ventral peripheries of P. sigmoidale conchs, attached predominantly to body chambers. These modes of occurrence suggest that the encrustation by anomiids occurred not on post‐mortem floating or sunken carcasses but on live conchs and that these organisms were rapidly buried by turbidity current deposits shortly after death. Attachment to both flanks and ventral peripheries of the retroversal hooks may indicate that at least adult individuals of P. sigmoidale did not lie on the sea floor and did not drag their body chambers. It is suggested that fully mature individuals of this ammonite species lived for a long period of time after having formed the retroversal hook because a few generations of anomiids have colonized a single body chamber. Such colonization by anomiids is also observed on Didymoceras awajiense, which is considered to be the closely related ancestral species of P. sigmoidale. This anomiid–heteromorph ammonite commensal relationship might continue to persist in descendants during the course of evolution of these heteromorph ammonites.     

A description of a new species Gyrodactylus moldovicus sp. n. (Monogenea: Gyrodactylidae) from the European mudminnow Umbra krameri Walbaum, 1792 from the lower Dnester basin

Gyrodactylus moldovicus sp. n. found on gills, body and in nasal cavities of the European mudminnow (Umbra krameri) differs from G. slovacicus Ergens, 1963 also living on the this host by bigger size of the body, anchors and marginal hooks; from G. cylindriformes Mueller et Van Cleave, 1932 living on the American mudminnow Umbra limi--by bigger size of the body; from G. limi Wood et Mizelle, 1957 also from U. limi--by the form of ventral and dorsal bars and form of marginal hooks. It differs from other freshwater gyrodactylids by special type of marginal hooks which have a hook-like end of the blade. Gyrodactylus moldovicus, G. slovacicus and G. limi have marginal hooks of quite different morphological types. By the morphology of anchors, ventral and sometimes dorsal bars and also morphology of cirrus, G. moldovicus is most related to three species from Cyprininae: G. stankovichi Ergens, 1970, G. longoacuminatus Zitnan, 1964 f. typica and G. shulmani Ling, 1962.     

First direct evidence of ammonoid ovoviviparity

Ammonoids had high evolutionary rates and diversity throughout their entire history and played an important role in the high‐resolution sub‐division of the Mesozoic, but much of their palaeobiology remains unclear, including the brooding habitat. We present our study of the first recorded ammonite embryonic shell clusters preserved with calcified embryonic aptychi in situ within the body chambers of mature macroconch shells of the Early Aptian (Early Cretaceous) ammonite Sinzovia sazonovae. The following support the idea that the clusters are egg masses, which developed inside ammonite body chambers: the absence of post‐embryonic shells and any other fossils in these clusters, the presence of the aptychi in all embryonic shell apertures and peculiarities of adult shells preservation. These facts confirm earlier speculations that at least some ammonoids could have been ovoviviparous and that, like many modern cephalopods, they could have reproduced in mass spawning events. The aptychi of ammonite embryonic shells are observed here for the first time, indicating that they were already formed and calcified before hatching. Our results are fully congruent with the peculiar modes of ammonoid evolution: quick recovery after extinctions, distinct evolutionary rates, pronounced sexual dimorphism and the nearly constant size of embryonic shells through ammonoid history. We assume that adaptation to ovoviviparity may be the reason for the presence of these features in all post‐Middle Devonian ammonoids.     

Calcareous Concretions and Non-Calcified Hooks in the Body Wall of Nemertean Worms

Two types of refractile inclusions in the integument of nemertean worms have been studied by X-ray microanalysis and electron microscopy. The body wall of Tetrastemma bilineatum contains ovoid concretions that are 10–20 μm long. Qualitative X-ray microanalyses indicate that the concretions are composed of calcium, phosphorus, barium and perhaps magnesium and sodium. In Zygonemertes sp. and Z. virescens minute hook-like structures occur throughout the epidermis. The hooks develop by one week post-fertilization and appear to be non-calcified in both larvae and adults. Each hook is formed within a large vacuole and consists of several fused subunits. The subunits lack well defined cortical and medullary regions and seem to be produced by the rough endoplasmic reticulum of the epidermal secretory cell. The possible relationships between the calcareous concretions and proboscidial stylets of nemerteans are discussed and the structure of the integumentary hooks is compared to that of rhabdite-like bodies secreted by other invertebrates.     

Development of secondary sexual characters and their relationship to ontogeny and seasonal reproductive period in Hyphessobrycon igneus (Ostariophysi: Characiformes)

Sexual dimorphism in size, anal‐fin shape and coloration of Hyphessobrycon igneus, Characidae, were examined. Males were more frequent at larger body sizes, confirming body size as a sexually dimorphic trait. Anal‐fin shape and the colour of all fins were the same for females and juveniles, differing only in adult males. Likewise, only adult males had bony hooks on fin rays; larger and more sexually mature males had the most numerous and developed hooks and hooks were most developed in degree and number during peak reproductive periods. Fin hooks regressed in number and developmental degree after the reproductive period, but restarted development with the beginning of the new reproductive period without completely disappearing. Results show that bony hooks have a development and regression cycle related to reproductive seasonality.     

A morphological gradient revisited: cave mollies vary not only in eye size

In the Mexican Cueva del Azufre, cave-dwelling Atlantic mollies (Poecilia mexicana, Poeciliidae) have been reported to exhibit a pronounced gradient in eye size from front cave chambers towards rear cave chambers. Here, we re-examined this morphological gradient using a geometric morphometric analysis of body shape to test for variation along the gradient in traits other than eye size. Our results indicate that fish within the cave also vary in head size. Whereas differences in eye size appear to be related to differences in the amount of light received among cave chambers, variance in head size coincides with differences in hydrogen sulfide concentrations. Consequently, the morphocline within the Cueva del Azufre is more complex than previously assumed and raises interesting new questions on the mechanisms maintaining these morphological variations on such a small spatial scale.     

Morphological description of three marine ciliates (Ciliophora, Scuticociliatia), with establishment of a new genus and two new species

Three marine scuticociliates, Falcicyclidium fangi nov. gen., nov. spec., Falcicyclidium atractodes nov. spec., and Cristigera media Kahl, 1928 were investigated using live observation and silver impregnation methods. The genus Falcicyclidium is distinguished by the combination of: (i) dorsoventrally flattened body, (ii) hook-like (falciform) paroral membrane, (iii) anterior end of paroral membrane posterior to anterior end of membranelle 1, and (iv) multiple caudal cilia. Falcicyclidium fangi nov. spec., the type of the new genus, can be recognized by the combination of its large size, extremely dorsoventrally flattened (3:1) body, consistently 10 somatic kineties, and the broad, elongate buccal area occupying 60% of the body length. Falcicyclidium atractodes nov. spec. is mainly characterized by a unique spine projecting from both the anterior and posterior end. The uncommon form, Cristigera media is redescribed based on the population from Qingdao, the statistic data and additional features, especially the morphology of the living cells, are documented.     

Erratum

Feet of two-toed sloths (Choloepus) are long, narrow, hook-like appendages with only three functional digits, numbers II, III, and IV; Rays I and V are represented by metatarsals. Proximal phalanges of complete digits are little more than proximal and distal articulating surfaces. All interphalangeal joints are restricted, by interlocking surfaces, to flexion and extension. Ankle and transverse tarsal joints, however, allow extreme flexion and inversion of foot. Powerful digital flexion is augmented by several muscles from extensor compartment of leg. Intrinsic foot musculature is reduced to flexors and extensors but these, with the exception of lumbricals, are large and well developed. Choloepus uses its feet much like hooks with distal phalanges and covering claws forming the “hook” element. These hook-like appendages are seemingly best suited for supports less than 50 mm in diameter suggesting that two-toed sloths may prefer supports of this size in their natural habitat.     

Investing in attachment: evolution of anchoring structures in acanthocephalan parasites

Secure attachment to host tissues is essential for survival and reproduction in parasitic organisms. The production of elaborate attachment structures must be costly, however, and investments in attachment should be approximately proportional to the likelihood that a parasite will be dislodged. In the present study, relative investments in attachment as a function of body size and the type of host used were examined across 138 species of acanthocephalans. These worms live anchored to the intestinal wall of a vertebrate host by inserting their hooked proboscis into host tissues. Taking proboscis volume into account, there is a negative interspecific relationship between the number of hooks borne on the proboscis and their mean length, reflecting a trade‐off between hook number and hook length. This supports the assumption that hooks are costly to produce, because any given species cannot simultaneously maximize both the relative number and relative length of the hooks it produces. There is a positive relationship between total worm size and total hook length, but it is weak, with a slope indicating that, as total body volume increases, total hook length also increases but at a slower rate. Indeed, relative investments in attachment, measured as hook length per unit body volume, decrease as worm size increases. Independently of total body size, investments in hook production are higher in species exploiting endothermic hosts, especially birds, than in those living in ectothermic hosts. Given the greater amounts of food passing through the gut of endotherms, and the richer and denser communities of intestinal parasites that they harbour, they are likely to select for greater investments in attachment. These results support the prediction that investments in attachment are influenced by the probability of being dislodged, and allow comparisons with other groups of intestinal parasites such as cestodes or trematodes. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 637–645.     

Seasonal variations of opisthaptoral hard parts of Gyrodactylus salaris Malmberg, 1957 (Monogenea: Gyrodactylidae) on parr of Atlantic salmon Salmo salar L. in the River Batnfjordselva,Norway

Seasonal variations in size and shape of the marginal hooks, the anchors and the ventral bar of the opisthaptor of Gyrodactylus salaris Malmberg, 1957 were studied. The G. salaris specimens were collected from parr of Atlantic salmon Salmo salar L. in the River Batnfjordselva, north-western Norway. Samples were taken at roughly monthly intervals during a two-year period. The marginal hooks, the anchors and the ventral bar showed considerable seasonal variation in size, but varied only slightly in shape. The variation in 12 of the 14 characters measured showed a significant regression to the variation in the water temperature. The total length of the marginal hooks of G. salaris can be considerably longer than the previously reported maximum of 40 m for the species.     

Acanthocephaloides cyrusi n. sp. (Acanthocephala: Arhythmacanthidae) from southeast African teleost fishes

A new species,Acanthocephaloides cyrusi, is described from the fishesSolea bleekeri andPomadasys commersoni from Lake St. Lucia, Natal, South Africa. It is distinguished from the other species in the genus by the more marked sexual dimorphism in length, the arrangement of hooks, the proboscis with the longest hooks at the anterior-most extremity and the greater size of the proboscis hooks and body spines. An acanthella, which may represent this species, was found in the tanaidApseudes digitalis.     

Chamber growth in ammonites inferred from colour markings and naturally etched surfaces of Cretaceous vascoceratids from Nigeria

Cretaceous Vascoceras and Jurassic Lytoceras show colour markings and etched surfaces representing original organic membranes between the septa. The main difference between the formation of ammonite and Nautilus chambers involved the continuous secretion of a gelatinous cameral liquid to support the ammonite mantle when it moved forward. The gel containing cyclically secreted membranes. here named pseudosepta, resembled the intra-cameral structures of the cuttlebone in Sepia. Pseudoscpta are attached to the shell wall in pseudosutures (Pseudoloben) which are particularly visible in the saddles of the septal suture and tend to mimic them. Their shape suggests reconstruction of posterior mantle shape during translocation. Drag-bands (Schleppstreifen) are spiral markings formed by the overlapping pseudosepta along the axial traces of the foliole folds. The chamber of ammonites was formed by a locally muscular mantle in a tripartite cycle: (1) the mantle initially remained attached to the saddles of the completed septal suture while muscular tissue within the umbilical lobes was contracted and rapidly reattached to the side of the lateral saddles; (2) the whole mantle subsequently crept forward by secreting a gelatinous matrix which contained telescoped membranes, with an adhesive function on pseudolobc flanks; (3) the mantle almost ceased to move within the sites of future lobules, but expanded and crept on before forming the mural and 'gutter' ridges of the septum. □ Ammonites, chamber growth, vascoceratids, LYTOCERAS, Nigeria.     

Peltidiphonte gen. n., a New Taxon of Laophontidae (Copepoda: Harpacticoida) from Coral Substrates of the Indo-West Pacific Ocean

A new genus of the harpacticoid family Laophontidae is described and named Peltidiphonte gen. n. Eight new species are assigned to this genus; they were collected from different locations in the Indo-West Pacific Ocean, including the Comoros, the Kenyan coast, the Red Sea, the Andaman Islands, the northern coast of Papua New Guinea, the Solomon Islands and the northeastern coast of Australia. Most of the specimens were collected from dead coral substrates, suggesting a close affinity between the members of the new genus and this substrate. Peltidiphonte gen. n. can easily be discriminated from other genera of the family by the extremely depressed body and by the shape of the antennule, bearing two (or three) processes on the first segment and a hook-like process along the outer margin of the second segment. An identification key for the new genus is provided.     

Taphonomy and ichnology of cephalopod shells in a Maastrichtian chalk from Western Australia

The post-mortem history of a prolific Maastrichtian ammonite and nautiloid fauna preserved as phosphatic steinkerns in chalk of the upper Miria Formation of Western Australia is described. Sediment infilling of phragmocones, required for their fossilisation, was accomplished by means of perforations in the shell wall induced by the activity of abundant endoliths. These include borings ascribed to clionid sponges (Entobia), thallophytes, polychaete worms (including Caulostrepis and probable Maeandropolydora), phoronids (Talpina and Gnathichnus) and others of conjectural origin. Sediment infilling by this mechanism is considered to be more applicable to the taphonomy of phragmocones in general than sediment entry through the siphuncle and to be indicative of low sedimentation rates for the hosting strata. Nonheteromorph ammonites, and the nautiloid Cimomia, are preserved almost exclusively as phragmocones in the upper Miria Formation, and are numerically subordinate to the heteromorph Eubaculites which, together with Glyptoxoceras, is preserved predominantly as body chambers. This phragmocone/body chamber preservational contrast is attributed to the influence of shell shape on pre-burial mechanical abrasion. The apparent dominance of Eubaculites is considered to be largely a preservational artifact and ascribed to the ease with which body chambers were infilled, and thereby favoured for steinkern formation, relative to phragmocones. Phragmocones, or parts thereof. not filled with sediment were eliminated from the fossil record by diagenetic aragonite dissolution. Many ammonites with open umbilici have the early whorls missing. This we attribute to the trapping of sediment in the umbilicus prior to burial, preventing endolith attack whereby the early whorls avoided a sediment infilling. However, the protected inner whorls of involute ammonites and Cimomia, which have closed umbilici, and the inner whorls of evolute ammonites where a cemented umbilical plug supported the mouldic cavity left by shell dissolution, were preserved and were commonly infilled with calcite spar later in diagenesis. The dearth of cephalopod fossils in chalk underlying the upper Miria Formation is ascribed to diagenesis in which aragonite dissolution was not preceded by cementation.     

Description and morphological variability of <Emphasis Type="Italic">Neoechinorhynchus beringianus</Emphasis> n. sp. (Acanthocephala: Neoechinorhynchidae) from north-eastern Asia

Neoechinorhynchus beringianus sp. n. is described from Pungitius pungitius L. in north-eastern Russia. Since 1986, when it was first found, it was reported as 'N. pungitius Dechtiar, 1971'. However, this new species differs from the latter in having an egg shell without a prolongation of the fertilisation membrane, a larger proboscis and proboscis hooks, a subterminal position of the female genital pore and a more slender trunk, and it occurs in a different site in the intestine. N. beringianus has a small, stout body with an asymmetrical position of the proboscis, which is located ventrally to and at an angle with the longitudinal axis of the body. The proboscis is wider than long, the hooks are of equal size in each circle but diminish in size posteriorly, whereas the lemnisci are subequal in length. It differs from those species of Neoechinorhynchus Stiles & Hassall, 1905 with somewhat similar characteristics in body length, proboscis size and proportions, proboscis hook lengths, egg size, size-ratio of the cement gland and testes, and the number of giant nuclei in the tegument and lemnisci. In different geographical populations of the new species, the sizes of both the proboscis and proboscis hooks exhibit some variation.     

The climbing habit in palms: Biomechanics of the cirrus and flagellum

Climbing palms in the Arecoideae (Desmoncus) and Calamoideae (rattan palms) both evolved cirrate leaves armed with hooks and grapnels for climbing. Some species of Calamoideae develop a different climbing organ known as the flagellum, which also bears hooks. The present study indicates that geometry and mechanical properties of the cirrus vary between species. Cirrate leaves are constructed to optimize bending and torsion in relation to the deployment of recurved hooks. Hook development, size, and strength vary along cirri and flagella and are consistent with observations of these attachment organs functioning as a ratchet mechanism: hooks increase in strength toward the base of attachment organs and always fail before the axis in strength tests. Hook size and strength differ between species and are related to body size and ecological preference. Larger species produce larger hooks, but smaller climbing palms of the understory deploy fine sharp hooks that are effective on small diameter supports as well as large branches and trunks. The ephemeral nature of climbing organs in palms provides a challenge to their life-history development, particularly in terms of mechanical constraints and remaining attached to the host vegetation; these differ significantly from many vines and lianas having more perennial modes of attachment.     

Acanthocephalus?amini n. sp. (Acanthocephala: Echinorhynchidae) from the freshwater fish Cichlasoma?urophthalmus (Günther) (Cichlidae) in Mexico

Acanthocephalus amini n. sp. (Palaeacanthocephala: Echinorhynchidae) is described from the intestine of Cichlasoma urophthalmus (Günther) (Pisces: Cichlidae) collected in the Río Champotón, a river in Campeche State, Mexico. It is the fourth species of Acanthocephalus Koelreuther, 1771 described from North American freshwater fishes, although two other species are known from South America. The new species is distinguished from other members of Acanthocephalus by features of its trunk, which is small, clavate, slightly expanded medially and bluntly pointed posteriorly. It is further distinguished by having a cylindrical proboscis armed with 13–14 longitudinal rows of 11–12 stout hooks; the apical and medial proboscis hooks are almost uniform in size and shape, decreasing in size towards the base; the posteriormost hooks are smaller, straighter and more slender than the anterior and middle hooks; and the lateral rows of hooks are more widely spaced, forming a conspicuous longitudinal area devoid of hooks. Furthermore, the lemnisci are saccate and shorter than the proboscis receptacle; and the neck is very short with a thick collar of trunk tegument, which encircles the base of proboscis. In males, the testes are in the middle third of trunk, diagonal, spherical and small relative to the body size, and there are six clavate cement glands. In females, the uterus forms a conspicuous, elongate, cylindrical egg reservoir. The new species is most similar to A. alabamensis Amin & Williams, 1983, but can be distinguished by its swollen, clavate trunk, the largest proboscis hooks being present apically and medially, smaller testes, a shorter male reproductive system relative to body size and females with a prominent uterus. They have different hosts and geographical distribution. The new species can be differentiated from Brasacanthus sphoeroides Thatcher, 2001, a similar species in a monotypic echinorhynchid genus, because the latter is larger, has smaller proboscis hooks and its lemnisci are longer than the receptacle.     

A new species of Acanthobothrium van Beneden, 1849 (Cestoda: Tetraphyllidea) from Rioraja castelnaui (Chondrichthyes: Rajoidei) in coastal waters of Argentina

Acanthobothrium marplatensis n. sp. is described from the spiral intestine of the skate Rioraja castelnaui taken off Mar del Plata, Argentina. The new species is compared with those species that it most closely resembles morphologically, to species reported from other skate hosts and to species described from the same geographical region. A. marplatensis is most similar to A. paulum, A. benedeni, A. costarricense, A. fogeli, A. himanturi, A. lintoni, A. monksi, A. olseni, A. puntarenasense and A. mathiasi. All of these species share a similar combination of characters (12–40 proglottids, 20–50 testes and total length of 2–10mm). A. marplatensis can be differentiated from these species and those described from skates by a combination of the following characters: 4.79–8.44 (6.18) mm long, with 18–30 (24) proglottids, spinose cephalic peduncle 160–338 (227) long, bothridial hooks with total length 93-134 (115), 24–39 (32) testes per proglottid, cirrus-sac curved anteriorly, ovarian lobes never reaching the level of the cirrus-sac and vaginal sphincter absent. Geographically, the new species can be distinguished from A. zapterycum and three different species designated as Acanthobothrium sp. reported from the coast of Uruguay and Argentina by the size of the worms, number of proglottids, position of the genital pores and size of the bothridial hooks. A. marplatensis showed the greatest preference for chambers 2 and 3 of the eight chambers of the spiral intestine. The mean intensity of infection in individual hosts increased with host size and was greater in spring and summer than in cold seasons. The intensity of infection was unrelated to host sex.     

Reappraisal of the latest Albian (Mortoniceras fallax Zone) cephalopod fauna from the classical Salazac locality (Gard,southeastern France)

The Salazac locality (Gard, southeastern France) is renowned for the richness of its cephalopod fauna (especially ammonites) from the Mortoniceras fallax Zone (uppermost Albian, Lower Cretaceous). However, most ammonite species have paradoxically been scarcely illustrated up to now. Furthermore, the rare assessments of ammonite taxonomic diversity are presumably inaccurate, as they either result from a drastic typological approach or could not benefit from the most recent advances in modern concepts of ammonoid intraspecific and ontogenetic variations. In this work, we document an original cephalopod assemblage from Salazac represented by abundant, well-preserved specimens (564 studied ammonites, including ca. 50% of heteromorphs) constituting a high diversity. The sample size and quality of this new material allowed some extensive morphological and biometric analyses whenever possible (i.e., for subdatasets in which a significant number of measurements is available) and to thoroughly assess intraspecific and ontogenetic variations for a few species. This led to determine the diagnostic value of various shell characters commonly used to characterize both well-documented and poorly-defined species. In particular, we show that a significant part of the variation in the number of ribs per whorl within the heteromorphic genera Mariella and Turrilitoides is directly related to shell size and that the strength of tubercles covaries negatively with the number of ribs in Mariella, therefore challenging the diagnostic value of these characters. We further suggest that rib density covaries with whorl shape in Mariella, following Buckman’s first rule of covariation. In total, one species of nautilid and 38 taxa (25 genera) of ammonites are described, among which five species are documented from Salazac for the first time. In turn, this study provides valuable data for future phylogenetic, biostratigraphical, biogeographical and broad-scale diversity studies.