Structure, ontogeny, and moulting of the olenid trilobite Ctenopyge (Eoctenopyge) angusta Westergård, 1922 from the upper Cambrian of Västergötland, Sweden

The genus Ctenopyge is known mainly from disarticulated sclerites and from rare complete specimens flattened in shales. Hitherto, very few specimens have been found preserved intact and in three dimensions. In a recently discovered fauna, however, in the Peltura minor Subzone in Västergötland, central Sweden, there occur several species of Ctenopyge , of which many are complete and superbly preserved; moreover they occur at all stages of growth. Of these the abundant Ctenopyge ( Eoctenopyge ) angusta Westergård, 1922 is described and reconstructed here as an adult, and the entire ontogeny is documented for all post–protaspid growth stages. Many characters typical of the adult, such as the long genal spines and the caudal spine, develop very early in ontogeny, and the relative dimensions of the cranidium do not greatly change during growth. Macropleural spines, however, develop later. The transitory pygidium, relatively large and shield–shaped in the early meraspid, later becomes very small as the ten thoracic segments are liberated; a median spine develops on the last thoracic segment only at the holaspid stage. Instar groupings can be clearly distinguished for the early stages. Recurrent associations of sclerites are interpreted as moulting configurations. As reconstructed, the genal spines are horizontal and parallel with the extended thorax; an adaptation which presumably allowed the trilobite to rest on the sea floor.     

Trilobite size-frequency distributions, recognition of instars, and phyletic size changes

Size-frequency analysis of over 5,000 Ordovician trilobites from the Teretiusculus Shales of the Builth inlier, central Wales, has revealed size distributions with counter intuitive shapes. Not only do most species show normal or slightly skewed distributions, despite the preponderance of moults, but there is no evidence of instar peaks. Such features can, however, be explained by reference to steady-state population structures of Recent marine arthropods, in which small individuals often form only a minor proportion of the post-larval population structure. Trilobite steady-state population structures would have differed in detail from species to species, but certain distribution shapes may have been characteristic of particular environments. These findings necessitate a reappraisal of previous work on trilobite size-frequency distributions, survivorship and recognition of instars. The Builth data also show the first clear evidence of phyletic size increase and parallel size changes in trilobites. ▭ Trilobites, size-frequency distributions, steady-state populations, instars, phyletic size changes.     

Ontogeny of the Upper Cambrian (Furongian) Olenid trilobite Protopeltura aciculate (Angelin, 1854) from Skåne and Västergötland,Sweden

Abstract: The ontogeny of the trilobite Protopeltura aciculata (Angelin, 1854) is described on the basis of material from the upper Cambrian (Furongian) of Andrarum (Skåne) and Hjelmsäter (Västergötland), 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.     

Rusophycus carleyi (James, 1885), Trace Fossils from the Lower Ordovician of Southern Morocco,and the Trilobites that Made Them

Thin-bedded, pyrite-rich, fine sandstones and mudstones of the Floian-Dapingian Upper Fezouata Formation contain abundant trace fossils Rusophycus carleyi in close association with a species of the asaphid trilobite Asaphellus. The sizes and shapes of this trilobite and the traces match closely. Five specimens have even been found where an articulated specimen of Asaphellus appears to be directly located over a specimen of Rusophycus carleyi within a thin bed of sandstone, suggesting that the trilobite animal may have been trapped on top of a trace that it had just made. Such intimate associations between a putative tracemaker and a trace are rare in the fossil record and particularly rare for Trilobita. The number of coxal impressions that form part of R. carleyi, eleven, matches the number expected for an asaphid trilobite (one for each of eight thoracic segments and one for each of three post-oral cephalic appendages). Impressions of the hypostome, thoracic tip impressions, cephalic margin, and pygidial margin in a few of the traces also match those of this asaphid trilobite. R. carleyi has been found in Ordovician strata of other parts of the world in association with asaphid trilobites.     

Evaluating paedomorphic heterochrony in trilobites: the case of the diminutive trilobite Flexicalymene retrorsa minuens from the Cincinnatian Series (Upper Ordovician), Cincinnati region

Flexicalymene retrorsa minuens from the uppermost 3 m of the Waynesville Formation of the Cincinnatian Series (Upper Ordovician) of North America lived approximately 445 Ma and exhibited marked reduction in maximum size relative to its stratigraphically subjacent sister subspecies, Flexicalymene retrorsa retrorsa. Phylogenetic analysis is consistent with the notion that F. retrorsa retrorsa was the ancestor of F. retrorsa minuens. F. retrorsa minuens has been claimed to differ from F. retrorsa retrorsa"in size alone," and thus presents a plausible example of global paedomorphic evolution in trilobites. Despite strong similarity in the overall form of the two subspecies, F. retrorsa minuens is neither a dwarf nor a simple progenetic descendant of F. retrorsa retrorsa. More complex patterns of global heterochronic paedomorphosis, such as a neotonic decrease in the rate of progress along a common ontogenetic trajectory with respect to size, coupled with growth cessation at a small size, "sequential" progenesis, or non-uniform changes in the rate of progress along a shared ontogenetic trajectory with respect to size, can also be rejected. Rather, differences between these subspecies are more consistent with localized changes in rates of character development than with a global heterochronic modification of the ancestral ontogeny. The evolution of F. retrorsa minuens from F. retrorsa retrorsa was largely dominated by modifications of the development of characters already evident in the ancestral ontogeny, not by the origin of novel structures. Factors promoting size reduction in F. retrorsa minuens appear to have been specific to this subspecies, because other co-occurring taxa, including other trilobite species, do not show marked differences in mean size.     

Trilobite taphonomy and ecology in Upper Ordovician carbonate buildups in Dalarna, Sweden

A major ecological grouping of the trilobites in the Upper Ordovician Boda Limestone buildups in Dalarna is outlined, based primarily on the modes of occurrences in the Jutjärn quarry. The so-called pockets are analysed with regard to their trilobite contents and distribution within a carbonate buildup. The pockets are of two general types. One consists of narrow opened fissures along bedding planes. The matrix of these may contain either of the peculiar blind Isocolus sjoegreni or Ityophorus undulatus , both of which appear to have been cavernous species, possibly belonging to the Bathyuracea (or Leiostegiacea). In addition, small proetids used the fissures for moulting. The other kind of pocket is a cavity largely filled with sediment from the surface of the buildup. The size is considerable, in rare cases up to many tens of cubic metres. Although the buildups housed about 90 trilobite species, each of these pockets is dominated by one or two out of seven large species. Of these, six are illaenids, the seventh the illaenid-like cheirurid Holotrachelus punctillosus. Usually only pygidia and cranidia are found. These species may have lived gregariously around the rims of the cavities. The remaining species, about 85% of the trilobite fauna, appear to have lived on the surface of the buildups. The largest group is the proetids with some 20 species (in addition to those known from pockets). Cheirurinids and lichids contribute with some 15 species each, and there are six odontopleurids. The distinctness of the 'community' is indicated by the circumstance that only three species are shared with contemporaneous level-bottom faunas in Sweden.     

Reconstruction of the shape and optics of the lenses in the abathochroal‐eyed trilobite Neocobboldia chinlinica

Until now, the structure and optics of the calcite lenses in abathochroal trilobite eyes have not been investigated. So, the relationship of the abathochroal eye to other types of trilobite eyes has remained unclear. We have reconstructed the exact shape and optics of the lenses in the eodiscid trilobite Neocobboldia chinlinica to determine the mechanism of its abathochroal eye. The distal lens surface has a convex profile, while on the proximal lens surface there is a small central bulge, resulting in an undulating profile. Due to this bulge, the curvature and refractive power of the central region of the lens are greater than those of the peripheral zone. Consequently, the lens is bifocal. However, Neocobboldia could not take advantage of this bifocal property of its tiny lenses because of the diffraction of light and the infinite depth of field in object space. For the same reason, it is also sure that the undulating lower surface of the abathochroal lens did not evolve as a Huygensian profile, correcting for spherical aberration, as suggested earlier. This undulation is a result of the presence of the central bulge, the evolutionary significance of which remains enigmatic. On the basis of our results, we have outlined an evolutionary scenario for development of the optics of the lenses in trilobite eyes.     

Trilobite ‘pelotons’: possible hydrodynamic drag effects between leading and following trilobites in trilobite queues

Energy saving mechanisms in nature allow following organisms to expend less energy than leaders. Queues, or ordered rows of individuals, may form when organisms exploit the available energy saving mechanism while travelling at near‐maximal sustainable metabolic capacities; compact clusters form when group members travel well below maximal sustainable metabolic capacities. The group size range, given here as the ratio of the difference between the size of the largest and smallest group members, and the size of the largest member (as a percentage), has been hypothesized to correspond proportionately to the energy saving quantity because weaker, smaller, individuals sustain the speeds of stronger, larger, individuals by exploiting the energy saving mechanism (as a percentage). During migration, small individuals outside this range may perish, or form sub‐groups, or simply not participate in migratory behaviour. We approximate drag forces for leading and following individuals in queues of the late Devonian (c. 370 Ma) trilobite Trimerocephalus chopini. Applying data from literature on Rectisura herculea, a living crustacean, we approximate the hypothetical walking speed and maximal sustainable speeds for T. chopini. Our findings reasonably support the hypothesis that among the population of fossilized queues of T. chopini reported in the literature, trilobite size range was 75%, while the size range within queues was 63%; this corresponds reasonably with drag reductions in following positions that permit c. 61.5% energy saving for trilobites following others in optimal low‐drag positions. We model collective trilobite behaviour associated with hydrodynamic drafting.     

Positional specification in the segmental growth pattern of an early arthropod

In many arthropods, there is a change in relative segment size during post-embryonic development, but how segment differential growth is produced is little known. A new dataset of the highest quality specimens of the 429 Myr old trilobite Aulacopleura koninckii provides an unparalleled opportunity to investigate segment growth dynamics and its control in an early arthropod. Morphometric analysis across nine post-embryonic stages revealed a growth gradient in the trunk of A. koninckii. We contrastively tested different growth models referable to two distinct hypotheses of growth control for the developing trunk: (i) a segment-specific control, with individual segments having differential autonomous growth progression, and (ii) a regional control, with segment growth depending on their relative position along the main axis. We show that the trunk growth pattern of A. koninckii was consistent with a regional growth control producing a continuous growth gradient that was stable across all developmental stages investigated. The specific posterior-to-anterior decaying shape of the growth gradient suggests it deriving from the linear transduction of a graded signal, similar to those commonly provided by morphogens. A growth control depending on a form of positional specification, possibly realized through the linear interpretation of a graded signal, may represent the primitive condition for arthropod differential growth along the main body axis, from which the diverse and generally more complex forms of growth control in subsequent arthropods have evolved.     

Body size and cell size in Drosophila: the developmental response to temperature

In Drosophila, like most ectotherms, development at low temperature reduces growth rate but increases final adult size. Cultures were shifted from 25 degrees C to low (16.5 degrees C) or to high (29 degrees C) temperature at regular intervals through larval and pupal stages, and the flies of both sexes showed an increase or decrease, respectively, in the size of thorax, wing and abdominal tergite. Size changes in the wing blade resulted from changes in the size of the epidermal cells (with only a small increase in cell number in males reared at low temperature). The temperature-shifts became less effective as they were made at successively later developmental stages, demonstrating a cumulative effect of temperature on adult size. The thorax and wing develop from the same imaginal disc, with most cell division occurring in larval stages, but they differ in timing of temperature sensitivity, which extends only to pupariation or into the late pupal stage, respectively. Growth of the adult abdomen occurs largely after pupariation but its size is temperature-sensitive through both larval and pupal stages. We discuss growth control in Drosophila and the likely effects of temperature on food assimilation, growth efficiency and allocation of nutrients to the production of different tissues.     

THE EXTRAORDINARY TRILOBITE FENESTRASPIS (DALMANITIDAE, SYNPHORIINAE) FROM THE LOWER DEVONIAN OF BOLIVIA

Abstract:  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 .     

Early larval ontogeny of the Permo-Carboniferous temnospondyl Sclerocephalus

The early larval development of the temnospondyl Sclerocephalus sp. is analyzed, based on 38 specimens from the Lower Rotliegend (Permo-Carboniferous boundary) of the Saar-Nahe Basin (south-west Germany). The study focuses on the smallest larval specimens, which exemplify changes in both proportions and ossification patterns. In comparison with dissorophoid larvae, the skull ossifies more fully and at a much faster rate; the smallest specimens already have completely formed circumorbital bones that are sutured throughout. Sculpturing undergoes two marked changes, first from uniformly pitted to pits of variable size and regional differentiation, and finally to the origin of ridges. The palate of small larvae differs from that of larger specimens in patterns of dentition, having more teeth including a denticle field on the cultriform process. The mandible of small larvae is described for the first time, being narrower than in adults and having three dentigerous coronoid elements. The smallest specimens have poorly ossified neural arches, lack vertebral centra, and have faintly ossified humeri, femora, and very poorly developed distal elements. The posterior ribs, metapodia, and phalanges appeared after the dermal elements of the pectoral girdle, whereas the scapulocoracoid and ischium are absent throughout the larval period. Early growth and differentiation of the limbs and the ilium illustrates the developmental patterning of the appendages, which proceeded from proximal to distal. Dermal squamation is uniform in small stages, consisting of round or oval osteoderms with pronounced growth rings; in large larvae, they start to differentiate in certain body regions.     

Ontogenetic Evidence for Dental Homologies and Premolar Replacement in Fossil and Extant Caenolestids (Marsupialia)

It is generally accepted that the South American marsupial family Caenolestidae is characterized in part by the absence or noneruption of the third deciduous premolar (dP3) in both jaws, although juvenile stages have rarely been identified in extant or fossil representatives of the family. Published illustrations of the dentary of the Miocene caenolestid Stilotherium suggested to us, however, that P3 erupted relatively late during ontogeny, after the eruption of M4. In extant marsupials, this eruption sequence appears to represent the plesiomorphic state and this pattern is generally associated with the eruption of dP3 earlier in ontogeny, and its subsequent replacement by the erupting P3. Therefore, we suspected that a dP3 erupted in earlier ontogenetic stages of caenolestids; to test this hypothesis we searched the mammalogy collections of three museums for evidence of dP3 in juvenile specimens of caenolestids. Examination of more than 180 specimens of the three extant genera of caenolestid marsupials resulted in the identification of only nine juvenile or subadult stages of dental eruption. Seven specimens of Caenolestes and Rhyncholestes corroborated our hypotheses of late eruption of P3 in Caenolestidae. In addition, the two youngest specimens of Caenolestes possessed a tiny, rudimentary dP3, measuring about 0.4 to 0.5 mm in greatest length, associated with a mesiolingual eruption pit containing the apex of the larger P3 in very early phases of eruption above the alveolar margins. The tiny dP3 is clearly nonfunctional in occlusion, and it is questionable whether it erupted above the gun margins in life. Comparison of the dentaries of subadult caenolestids with four dentaries of the Miocene genus Stilotherium corroborated our initial impression that the fossil genus exhibited evidence of a late-erupting P3, comparable to the condition in extant caenolestids. We suggest that examination of other specimens of juvenile dentitions, skulls, and skeletons in museum collections can provide additional insight into the developmental and evolutionary biology of mammals.     

NOTE ON A NEW PROETID FROM UPPER DEVONIAN OF KWANGSI

In 1950,some interesting specimens of small trilobites have been forwardedthe writer for examination by Prof.C.C.Y(?) which form the subject of the presentcommunication.The specimens were collected by Prof.Y(?) from the PisoliticLimestone Series near Fonglingshan,Lingchuan-Hsien,Kwangsi province.The trilobite fossils were found in association with some ostracods and smallChonetes.The ostracods have been recently identified by Y.T.Hou as Richterinasubstriatula Hou,R.subhemisphaerica Hou and Healdia lingchuanensis Houassigning to the Upper Devonian age.It is interesting that the evidence of thetrilobites points also to the Upper Devonian age.     

Presumed postlarval pentacrinoids from the Lower Devonian Hunsrück Slate,Germany

Several tiny crinoids with crowns as small as 1 mm, or less, in width are newly recognized from the Hunsrück Slate of southwestern Germany. The presence of erect arms above an amorphous calyx in some specimens can be inferred. Based on comparison with the size and gross morphology of developmental stages in living crinoids, these tiny Hunsrück crinoids are judged to be at an early postlarval stage that is analogous to the pentacrinoid stage just after development from the stalked, but armless, smaller cystidean larval stage found in both living comatulids and isocrinids. Some of these tiny crinoids have a stalk up to 4 mm long attached to a now pyritized former substrate. Their clustered occurrence suggests gregarious settlement of larvae. Taxonomic identification of these presumed pentacrinoids is not possible, even to the sub‐class level, although they are preserved with larger juveniles of the cladids Propoteriocrinus and Lasiocrinus. These larger juveniles exhibit 3‐D pyritized calcite plates, whereas the probable pentacrinoids appear to be preserved as flattened, micro‐crystalline pyritized dermal tissues that enclosed lightly calcified, porous ossicles. The pentacrinoids were likely buried within weeks or months of hatching, based on developmental stages in similar‐sized living crinoids. These tiny crinoids, presumably pentacrinoids, are a further example of the extraordinarily detailed preservation of delicate tissues in pyrite from the Hunsrück Slate. They are most likely the pentacrinoid stage from one or more of the crinoid taxa (30 genera) present in the Hunsrück Slate. Assuming these are not microcrinoids, they are the first report of pentacrinoids from the fossil record and document that a Palaeozoic sister group to modern crinoids had similar developmental stages.     

A dual upstream open reading frame-based autoregulatory circuit controlling polyamine-responsive translation

A novel form of translational regulation is described for the key polyamine biosynthetic enzyme S-adenosylmethionine decarboxylase (AdoMetDC). Plant AdoMetDC mRNA 5' leaders contain two highly conserved overlapping upstream open reading frames (uORFs): the 5' tiny and 3' small uORFs. We demonstrate that the small uORF-encoded peptide is responsible for constitutively repressing downstream translation of the AdoMetDC proenzyme ORF in the absence of increased polyamine levels. This first example of a sequence-dependent uORF to be described in plants is also functional in Saccharomyces cerevisiae. The tiny uORF is required for normal polyamine-responsive AdoMetDC mRNA translation, and we propose that this is achieved by control of ribosomal recognition of the occluded small uORF, either by ribosomal leaky scanning or by programmed -1 frameshifting. In vitro expression demonstrated that both the tiny and the small uORFs are translated. This tiny/small uORF configuration is highly conserved from moss to Arabidopsis thaliana, and a more diverged tiny/small uORF arrangement is found in the AdoMetDC mRNA 5' leader of the single-celled green alga Chlamydomonas reinhardtii, indicating an ancient origin for the uORFs.     

Die paradoxe Fruchtbarkeit der Strahlenflosser

The paradox fecundity of Actinopterygii More than fifty percent of about 70,000 vertebrate species are bony fish or Actinopterygii. These are characterized by two prominent and unique features: very tiny early life stages, independent of species or size, and a seemingly paradox fecundity, depending on size and age of the spawning fish. Usually even small fish produce quantities of eggs outnumbering those of other vertebrates and many invertebrates by two to more than five orders of magnitude. This suggests an unusual reproduction strategy of bony fish compared to other animal species. Due to their abundance, growth rate and consumption habit, little fish convert large amounts of short‐lived plankton biomass to long‐lived fish biomass. This generates a huge depot of food which is exploited by all carnivores living in or at aquatic environments. Not least or even first of all bony fishes themselves exist on this biomass store when grown to a larger size and prefer larger food. This behaviour likely implies an energy gain in a habitat where most biomass is produced by tiny and short lived plankton organisms and stored as fish biomass.     

A laboratory model of abiotic peloid formation

Peloids are rounded grains of micritic calcite whose origin has been attributed to various biological and abiotic mechanisms. To constrain abiotic parameters that favour the formation of peloids, we precipitated calcite crusts in the absence of microorganisms. Clotted opaque fabrics that formed during the initial stages of the experiment consisted of ～10 µm peloids, while compact clear sparitic crusts precipitated in subsequent stages. The increasing supersaturation of the solution in time is responsible for this morphological succession. Initially, peloids form by the radial growth of spar crystals around a small number of nuclei. As the supersaturation increases, more spar crystals nucleate and aggregate nonradially into compact crusts. Rounded clotted precipitates are a consequence of the growth in suspension and geopetal settling, and isopachous crusts grow in the absence of these processes. Although peloids are commonly assumed to have a microbial origin, our results show that very similar morphologies can be created by purely abiotic mechanisms. Thus, the biological origin of rounded micritic calcite grains in the rock record must be verified against the abiotic null‐hypothesis in each specific case.     

Developmental traits and life strategy of redlichiid trilobites

The developmental mode of four redlichiid trilobites is summarized, based upon exceptionally well-preserved, articulated specimens from Cambrian Series 2 (stages 3 and 4) strata in southwestern China and South Australia. These relatively complete developmental sequences indicate a balanced rate in segment increase and addition to the thorax during the meraspid phase, which might explain why most redlichiids possess micropygous body patterning during ontogeny. In addition, an analysis of the size distribution, developmental strategy, and distribution of specimen numbers at different growth stages reveals a distinct developmental strategy during the redlichiid life cycle. A relatively short pre-holaspid and a prolonged holaspid phase in these redlichiid taxa offers insight into the developmental control and life strategy in these primitive arthropods.