The history of Devonian-Carboniferous reef communities: Extinctions,effects, recovery

Summary Analysis of the taxonomic composition, diversity and guild structure of five “typical” reef and mud mound communities ranging in age from Late Devonian-Early Carboniferous indicates that each of these aspects of community organization changed dramatically in relation to three extinction events. These events include a major or mass extinction at the end of the Frasnian; reef communities were also effected by less drastic end-Givetian and mid-late Famennian extinctions of reef-building higher taxa. Peak Paleozoic generic diversities for reef-building stromatoporoids and rugose corals occurred in the Eifelian-Givetian; reef-building calcareous algal taxa were longranging with peak diversity in the Devonian. These three higher taxa dominated all reef-building guilds (Constructor, Binder, Baffler) in the Frasnian and formed fossil reef communities with balanced guild structures. The extinction of nearly all reef-building stromatoporoids and rugose corals at the end of the Frasnian and the survival of nearly all calcareous algac produced mid-late Famennian reef communities dominated by the Binder Guild. Despite the survival of most calcareous algae and tabulate corals, the mid-late Famennian extinction of all remaining Paleozoic stromatoporoids and nearly all shelf-dwelling Rugosa brought the already diminished Devonian reef-building to a halt. These Devonian extinctions differ from mass extinctions by the absence of a statistically significant drop in taxonomic diversity and by their successional and cumulative effects on reef communities. Tournaisian mud mounds contain communities markedly different from the frame-building communities in Late Devonian and Visean reefs. Mound-building biotas consist of an unusual association dominated by erect, weakly skeletonized members of the Baffler Guild (chiefly fenestrate Bryozoa; Pelmatozoa) and laterally expanded, mud-binding algae/stromatolites and reptant Bryozoa. The initial recovery to reefs with skeletal frameworks in the Visean was largely due to the re-appearance of new species of abundant colonial rugose corals (Constructor Guild) and fenestrate Bryozoa. This Frasnian-Visean evolution in the taxonomic composition and structure of the reef-building guilds is also expressed by abrupt changes in biofacies and petrology of the reef limestones they produced. Thus, “typical” Frasnian reef limestones with balanced guild structures are framestones-boundstones-bafflestones, Famennian reefs are predominantly boundstones, Tournaisian mud mounds are bafflestones and Visean reefs are bafflestones-framestones.     

Middle paleozoic waulsortian-type mud mounds in Southern Fergana (Southern Tien-Shan,commonwealth of independent states): The shallow-water atoll model

Summary Several Waulsortian-type mud mounds nearly 500 m thick and about 5 km long occur in the Middle Paleozoic carbonate section of the Aktur nappe in the mountains on the right bank of Isfara river. These buildups form a well developed barrier system that stretches along the South Ferganian carbonate platform margin and divides the carbonate complex into a fore-reef and a back-reef part. The time of the mounds' most active growth was from the Late Silurian (Ludlow) to the Middle Devonian (Eifel). Three main facies types can be recognized in the mud mounds: 1. micritic core facies, 2. sparitic flank facies and 3. loferitic capping facies. The central massive or crudely bedded part of the mounds consists of white or light grey clotted micrite. Macrofossils are rare. The sparitic flank facies in contrast consists of coarse and densely packed crinoidal wackestone-floatstones with some brachiopod shell debris. Solitary rugose corals, tabulate corals, stromato-poroids and fragments of mollusks are also abundant. The tops of the mounds are usually covered with loferitic pelmicrites or oolitic grainstone caps. Stromatactis-like structures are very rare and poorly developed in the South Ferganian mud mounds. However, almostin all such mounds horizons of calcitic breccias can be found. In order to explain all the features found in the Fergana mounds an ‘atoll-like’ model has been proposed which starts the evolution of the mud mounds with a small nucleus bioherm. The main stage of the evolution corresponds to an atoll-like structure developing on the surface of shallow water platforms. White clotted micrite of the mound core facies is interpreted as a accumulation of fine-grained sediment in an inner lagoon flanked by crinoidal bar deposits. The mound flank facies represents the atoll rim deposits from where the carbonate mud is derived. The capping loferitic facies is considered as tidal flat deposit that developed on top of the buildups during the last stage of its evolution. The knoll shape of the mounds is explained by the retreat of the atoll flanking crinoidal bars back into the inner lagoon during the rise in sea level. Stromatactis-like structures of small cavities filled with sparry calcite owe their existence to burrowing organisms. Calcitic breccias are interpreted as paleokarst collapse breccias. They indicate that the tops of the mud mound became subaerially exposed. Other evidence for a subaerial exposure can be seen in the occurrence of Variscian ‘black and white’ limestone gravel on the tops of some mud mounds. According toWard et al. (1970) these sediments were produced above the sea level at the edge of hypersaline lakes situated on islands.     

Can the dietary conservatism of predators compensate for positive frequency dependent selection against rare,conspicuous prey?

The evolution of conspicuous colouration in prey is puzzling because such coloration attracts the attention of predators. Anti-apostatic selection, in which rare prey forms are predated disproportionately often, is a second potential obstacle to the evolution of conspicuous colouration in prey, as bright novel prey forms are likely to be very rare when they first appear in populations. It has recently been postulated that dietary conservatism in predators, an extended feeding avoidance of novel prey, would allow novel conspicuous prey to survive and multiply despite anti-apostatic and conspicuousness effects. We tested this hypothesis for a novel prey type arising in an otherwise cryptic population, providing a direct test of whether anti-apostatic selection or the predators’ wariness to attack the novel prey type is the more important force acting on the novel conspicuous prey. We conducted our experiment in the “Novel World”; an experimental system designed to test predators’ foraging decisions in a large landscape. We found that the conspicuous, novel prey suffered high initial costs of conspicuousness compared with cryptic prey, since most of these prey were attacked during the first “generation”, with no opportunity to “reproduce”. However, a subset of the 17 birds (24%) were following a dietary conservative foraging strategy and they were reluctant to eat the novel prey. Interestingly these birds were not more neophobic or less explorative. Our data demonstrate how difficult it is for the novel conspicuous prey to survive in cryptic populations, but they also highlight the importance of the predator’s foraging strategies in helping to promote the evolution and maintenance of aposematism.     

Typostrophism in Palaeozoic Ammonoids?

The anti-Darwinian “Typostrophe Theory” of O.H.Schindewolf can be put to the test by revisiting the ammonoid examples on which this macroevolutionary model was founded. It is shown that none of the three theoretical elements saltationism, internalism, and cyclism can be supported by empirical data obtained from ammonoid research. Putative saltations (“Typogenesis”) were feigned because of the lack of knowledge of intermediate forms. Internalistic and orthogenetic development (“Typostasis”) can only be favoured by neglecting possible functions of morphological characters. Preprogrammed extinction of “degenerated” clades (“Typolysis”) is unlikely when ruling out anthropocentric views regarding ammonoid morphology. In terms of evolution of Palaeozoic ammonoids, there is no basis for the preference of the “Typostrophe Theory” or some of its composing elements, including the “Type Concept” and “Proterogenesis”, over the Darwinian evolutionary model and the Modern Synthesis.      

Anthracoporella mounds in the Late Carboniferous Auernig Group,Carnic Alps (Austria)

Summary A heretofore undocumented example of skeletal mounds formed by the dasycladacean algaAnthracoporella spectabilis is described from mixed carbonate-clastic cycles (Auernig cyclothems) of the Late Carboniferous (Gzhelian) Auernig Group of the central Carnic Alps in southern Austria. The massive mound facies forms biostromal reef mounds that are up to several m thick and extend laterally over more than 100 m. The mound facies is developed in the middle of bedded limestones, which are up to 16 m thick. These limestones formed during relative sea-level highstands when clastic influx was near zero. The mound facies is characterized by well developed baffler and binder guilds and does not show any horizontal or vertical zonation. Within the massive mound faciesAnthracoporella is frequently found in growth position forming bafflestones and wackestones composed of abundantAnthracoporella skeletons which toppled in situ or drifted slightly.Anthracoporella grew in such profusion that it dominated the available sea bottom living space, forming ‘algal meadows’ which acted as efficient sediment producers and bafflers. BecauseAnthracoporella could not provide a substantial reef framework, and could not withstand high water turbulence, the biostromal skeletal mounds accumulated in shallow, quiet water below the active wave base in water depths less than 30 m. The massive mound facies is under- and overlain by, and laterally grades into bedded, fossiliferous limestones of the intermound facies, composed mainly of different types of wackestones and packstones. Individual beds containAnthracoporella andArchaeolithophyllum missouriense in growth position, forming “micromounds’. Two stages of mound formation are recognized: (1) the stabilization stage when bioclastic wackestones accumulated, and (2) the skeletal mound stage when the sea-bottom was colonized byAnthracoporella and other members of the baffler and binder guilds, formingAnthracoporella bafflestones and wackestones of the mound facies. A slight drop in sea-level led to the termination of the mound growth and accumulation of organic debris, particularly calcareous algae, fusulinids, crinoids and bryozoans, forming well bedded limestones, which overlie the mound facies     

Sedimentology and stromatoporoid palaeoecology of Frasnian (Upper Devonian) carbonate mounds in southern Belgium

Da Silva, A.‐C., Kershaw, S. & Boulvain, F. 2011: Sedimentology and stromatoporoid palaeoecology of Frasnian (Upper Devonian) carbonate mounds in southern Belgium. Lethaia, Vol. 44, pp. 255–274. Stromatoporoids are the most abundant large skeletal organisms in middle Frasnian carbonate mound environments of southern Belgium. They occur in environments ranging from flank and off‐mound, mound core, shallow mound and restricted mound. A detailed log and comprehensive sampling of stromatoporoids in a single section cutting through all middle Frasnian mound levels in La Boverie–Rochefort Quarry, near Rochefort and Dinant reveals a stromatoporoid assemblage comprising 10 genera; 472 samples, containing an overall total of 3079 stromatoporoids (including complete and fragmented specimens) have been studied. The following list gives abundance using numbers of specimens and areas of total stromatoporoid area on outcrop surfaces (% number; % area in cm²): Actinostroma (0.4; 9.2), Amphipora (15.5; 1.7), Atelodictyon (0.2; 4.4), Clathrocoilona (0.3; 0.5), Euryamphipora (13.7; 0.7), Idiostroma (2; 1.9), Salairella (1.2; 9.6), branching Stachyodes (43.2; 59.1), laminar Stachyodes australe (1.9; 1.3), Stictostroma (4.8; 13.1) and Trupetostroma (0.2; 0.8), showing that Stachyodes is approximately half of the total assemblage. Deeper environments contain more abundant low profile forms, shallow water facies contain more domical and bulbous forms; branching forms are ubiquitous. Low profile stromatoporoids are likely to have been important sediment stabilizers that may have led to expansion of the carbonate factory, and they may have therefore contributed to the structural building of the mounds. Stromatoporoid‐coral intergrowths are observed in only Stictostroma suggesting that there is a close biological relationship between them; however, stromatoporoid skeletons in almost all cases appear to be unaffected by the presence of intergrown corals, suggesting they were commensals. □Frasnian, Late Devonian, mounds, palaeoecology, stromatoporoid.     

Subtropical coral-reef associated sedimentary facies characterized by molluscs (Northern Bay of Safaga, Red Sea, Egypt)

Summary The shallow marine subtropical Northern Bay of Safaga is composed of a complex pattern of sedimentary facies that are generally rich in molluscs. Thirteen divertaken bulk-samples from various sites (reef slopes, sand between coral patches, muddy sand, mud, sandy seagrass, muddy seagrass, mangrove channel) at water depths ranging from shallow subtidal to 40m were investigated with regard to their mollusc fauna >1mm, which was separated into fragments and whole individuals. Fragments make up more than 88% of the total mollusc remains of the samples, and their proportions correspond to characteristics of the sedimentary facies. The whole individuals were differentiated into 622 taxa. The most common taxon,Rissoina cerithiiformis, represented more than 5% of the total mollusc content in the samples. The main part of the fauna consists of micromolluscs, including both small adults and juveniles. Based on the results of cluster-, correspondence-, and factor analyses the fauna was grouped into several associations, each characterizing a sedimentary facies: (1) “Rhinoclavis sordidula—Corbula erythraeensis-Pseudominolia nedyma association” characterizes mud. (2) “Microcirce sp.—Leptomyaria sp. association” characterizes muddy sand. (3)”Smaragdia spp.-Perrinia stellata—Anachis exilis—assemblage” characterizes sandy seagrass. (4) “Crenella striatissima—Rastafaria calypso—Cardiates-assemblage” characterizes muddy seagrass. (5) “Glycymeris spp.-Parvicardium sueziensis-Diala spp.-assemblage” characterizes sand between coral patches. (6) “Rissoina spp.-Triphoridae —Ostreoidea-assemblage” characterizes reef slopes. (7) “Potamides conicus—Siphonaria sp. 2—assemblage” characterizes the mangrove. The seagrass fauna is related to those of sand between coral patches and reef slopes with respect to gastropod assemblages, numbers of taxa and diversity indices, and to the muddy sand fauna on the basis of bivalve assemblages and feeding strategies of bivalves. The mangrove assemblage is related to those of sand between coral patches and the reef slope with respect to taxonomic composition and feeding strategies of bivalves, but has a strong relationship to those of the fine-grained sediments when considering diversity indices. Reef slope assemblages are closely related to that of sand between coral patches in all respects, except life habits of bivalves, which distincly separates the reef slope facies from all others.     

Reconstructing atoll-like mounds from the Frasnian of Belgium

A succession of Frasnian mounds on the southern border of the Dinant Synclinorium (Belgium) was investigated for their facies architecture, sedimentary dynamics and palaeogeographic evolution. Seven mound facies were defined from the Arche (A) and Lion (L) members, each characterized by a specific range of textures and association of organisms (A2/L2: red or pink limestone with stromatactis, corals and crinoids; A3/L3: grey, pink or green limestone with stromatactis, corals and stromatoporoids; A4/L4: grey limestone with corals, peloids and dasycladaceens; A5/L5: grey microbial limestone; A6/L6: grey limestone with dendroid stromatoporoids; A7/L7: grey laminated limestone with fenestrae; and A8/L8: grey bioturbated limestone). Laterally equivalent sediments include substantial reworked material from the buildups and background sedimentation. Textures and fossils suggest that A2/L2 and A3/L3 facies developed close to storm wave base, in a subphotic environment. Facies A4/L4, occurring near fair weather wave base in the euphotic zone, includes lenses of A5/L5 with stromatolitic coatings and thrombolithes. A6/L6 corresponds to a slightly restricted environment and shows a progressive transition to fenestral limestone of A7/L7. This facies was deposited in a moderately restricted intertidal area. A8/L8 developed in a quiet lagoonal subtidal environment. The mounds started with A2/L2 or A3/L3 in which microbial lenses and algal facies A4/L4 became progressively more abundant upwards. Following 20 m of laterally undifferentiated facies, more restricted facies occur in the central part of the buildups. This geometry suggests the initiation of restricted sedimentation, sheltered by bindstone or floatstone facies. The facies interpretation shows that after construction of the lower part of the mounds during a transgression and a sea-level highstand, a lowstand forced reef growth to the margin of the buildups, initiating the development of atoll-like crowns during the subsequent transgressive stage. The persistence of restricted facies results from the balance between sea-level rise and reef growth.     

Depth-related ecological zonation of a carboniferous carbonate ramp: Upper Viséan of Béchar Basin,Western Algeria

Summary Following the demise of the stromatoporoid-coral reef community in Late Frasnian time, Lower Carboniferous carbonate shelf profiles possessed a ramp geometry, with major organic buildups represented by mud-rich mounds. Microfacies petrography of the exceptionally well-preserved Upper Viséan (Lower Carboniferous) carbonate ramp of the Béchar Basin, Algerian Sahara, may well contribute significantly to our understanding of the paleoecological zonation of Carboniferous non-rimmed platforms, and of the still enigmatic mounds commonly referred to as Waulsortian banks or mounds. Facies are grouped into two broad groups: (a) a mound facies group which comprises sponge wackestone-bafflestone, sponge-fenestellid bafflestone-wackestone, crinoid wackestone-packstone, and bedded flanks of intraclastic wackestone-packstone, all four facies composing the actual mud-rich mounds, and (b) a supramound facies group composed of coral-microbial framestone, crinoid packstone-grainstone, algal-foraminiferal grainstone and oolite grainstone. Calcareous algae are important bathymetric indicators and are used to delineate three bathymetric zones based on light penetration: the aphotic zone, which contains no calcareous algae; the dysphotic zone, where there is little ambient light, and which is characterized by the presence of red algae (Fasciella, Ungdarella, Stacheia, Epistacheoides) and absence of green algae; and the euphotic zone, which receives the full spectrum of sunlight, and is characterized by the occurrence of both green algae (Koninckopora, Kamaenella, Kamaena, Palaeoberesella, Calcisphaera, Anthracoporellopsis, Issinella, Exvotarisella) and red algae. Integration of algal zonation, distribution of the other biota, and recurrence of distinct assemblages, enable recognition of seven depth-related benthic assemblages. Together with the physical properties of the facies, the benthic assemblages were used to define seven bathymetric zones, from upper to lower ramp: (1) algal assemblage (upper ramp); (2) crinoid-ramose bryozoan assemblage (mid-ramp); and (3) productid brachiopod assemblage, (4) colonial rugose coral-microbial encruster assemblage, (5) crinoid-fenestellid assemblage, (6) sponge-fenestellid, and (7) sponge assemblage (lower ramp). The vertical zonation of the mud-rich mounds and associated facies differ from that reported from the classical Upper Tournaisian-Lower Viséan Waulsortian mound-bearing successions.     

Voiceprint identification and its application to sociological studies of wild Japanese monkeys (Macaca fuscata yakui)

Japanese monkeys often exchange the particular vocal sound, “coo,” especially when they feed or move as a group. It was considered that the “coo” sound had no positive social meaning, perhaps because the “coo” sound network and its function were hidden behind other behavioral observations. For identification of the vocalizer only from hearing the “coo” sound, three phonetic values, i.e., the “fundamental,” “duration,” and “formants,” plus other characteristics were used as indices of voiceprints. The results indicated that these were effective for identifying the vocalizer in two-thirds of the adults in the study troop which was composed of 12 adults and 16 immature members. The “coo” sound exchange network among the troop members (adults) was drawn on the basis of the voiceprint identification. The network showed three characteristics as follows: (1) matriarchs of the kin-groups frequently exchanged “coo” sounds with each other; (2) the other females exchanged “coo” sounds mostly within their own kin-groups; and (3) males seldom participated in the “coo” sound exchange. This suggests that “coo” sound exchange plays a central role for the matriarch of kin-groups in binding each kin-group and, ultimately, in binding all members together into an organized troop.     

Compostional variations and patterns of condont reworking in Late Devonian and early carboniferous calciturbidites (Moravia, Czech Republic)

Summary Compositional variations and grain-size properties of both carbonate constituents and conodonts as an alternative component group were used for interpreting the processes governing the deposition of upper Famennian and middle Tournaisian calciturbidites in Moravia, Czech Republic. Both the composition and grain-size properties of conodont element associations showed to be markedly dependant on facies type of their host sediment. Upper Devonian calciturbidite successions deposited on flanks of wide, Moravian-Silesian carbonate platform are composed mainly of echinoderm-and peloid-rich wacke/packstones and intraclastic float/rudstones (fine-grained calciturbidites, “normal” calciturbidites with Tab Bouma sequences, debris-flow breccias) with abundance of shelf-and shelf margin conodont taxa and epipelagic and “mesopelagic” conodonts. Upper Devonian calciturbidites deposited on slopes of volcanic sea-mounts are composed of echinoderm-and peloid-rich wacke/packstones and float/rudstones with increased proportion of intraclasts and volcanigenic lithoclasts (fine-grained calciturbidites, normal calciturbidites), yeilding abundant conodont associations with higher proportion of “mesopelagic” taxa compared to the platform-flank examples. Middle Tournaisian calciturbidite succession composed of crinoid-, peloid-, intraclast-and lithoclast-rich lime mudstones, wacke/packstones and float/rudstones (normal calciturbidites and debris-flow breccias) yielded conodont element associations rich in shelt-and shelf-margin taxa, “mesopelagic” conodonts and reworked Middle-and Upper Devonian conodonts. In general, the ratio of shelf-and shelf margin conodont taxa to “mesopelagic” taxa is distinctly lower in finegrained calciturbidites than it is in normal calciturbidites and debris-flow breccias. Grain-size properties (mean grain size and sorting) and percentage of fragmented conodont elements, too, are markedly dependant on the facies type: in fine-grained calciturbidites the values of mean grain-size and fragmentation are low and the sorting is good to very good whereas in normal calciturbidites and debris-flow breccias the values of mean grain-size and fragmentation are distinctly higher and the sorting is poorer. The interdependence of facies type and composition and grain-size properties of conodont element associations in gravity-flow deposits is explained as resultant from hydrodynamic sorting during turbidity current flow and final deposition of the bed. Compositional variations observed in our sections may thus be attributed to facies variability (coarsening-and thickening-upward trends) rather than to sea-level fluctuations (highstand shedding of carbonate platforms). On the other hand, significant enrichment in reworked conodont taxa in middle Tournaisian normal calciturbidites compared to scarcity and/or absence of such conodonts in essentially identical facies of upper Famennian age indicate sea-level to be the major control governing such compositional variations, with low relative sea-level stand in middle Tournaisian and high relative sea-level stand in upper Famennian. Thorough analysis of conodont evolution, palaeoecology and taphonomy, with emphasis on understanding the processes of deposition of their host rock, are recommended for any biostratigraphic and biofacies study to be done in carbonate sediments deposited under strong hydrodynamic regimes, such as calciturbidites, temperstites, debris-flow deposits, shelf-edge oolitic sands, tidal-channel facies etc.     

The Jackprot Simulation Couples Mutation Rate with Natural Selection to Illustrate How Protein Evolution Is Not Random

Protein evolution is not a random process. Views which attribute randomness to molecular change, deleterious nature to single-gene mutations, insufficient geological time, or population size for molecular improvements to occur, or invoke “design creationism” to account for complexity in molecular structures and biological processes, are unfounded. Scientific evidence suggests that natural selection tinkers with molecular improvements by retaining adaptive peptide sequence. We used slot-machine probabilities and ion channels to show biological directionality on molecular change. Because ion channels reside in the lipid bilayer of cell membranes, their residue location must be in balance with the membrane’s hydrophobic/philic nature; a selective “pore” for ion passage is located within the hydrophobic region. We contrasted the random generation of DNA sequence for KcsA, a bacterial two-transmembrane-domain (2TM) potassium channel, from Streptomyces lividans, with an under-selection scenario, the “jackprot,” which predicted much faster evolution than by chance. We wrote a computer program in JAVA APPLET version 1.0 and designed an online interface, The Jackprot Simulation , to model a numerical interaction between mutation rate and natural selection during a scenario of polypeptide evolution. Winning the “jackprot,” or highest-fitness complete-peptide sequence, required cumulative smaller “wins” (rewarded by selection) at the first, second, and third positions in each of the 161 KcsA codons (“jackdons” that led to “jackacids” that led to the “jackprot”). The “jackprot” is a didactic tool to demonstrate how mutation rate coupled with natural selection suffices to explain the evolution of specialized proteins, such as the complex six-transmembrane (6TM) domain potassium, sodium, or calcium channels. Ancestral DNA sequences coding for 2TM-like proteins underwent nucleotide “edition” and gene duplications to generate the 6TMs. Ion channels are essential to the physiology of neurons, ganglia, and brains, and were crucial to the evolutionary advent of consciousness. The Jackprot Simulation illustrates in a computer model that evolution is not and cannot be a random process as conceived by design creationists.     

Paleoecology of Pennsylvanian phylloid algal buildups in south Guizhou,China

Pennsylvanian phylloid algal reefs are widespread and well exposed in south Guizhou, China. Here we report on reefs ranging from 2 to 8 m thickness and 30–50 m lateral extension. Algae, the main components, display a wide spectrum of growth forms, but are commonly cyathiform (cup-shaped) and leaf-like (undulate plates). The algal reef facies is dominated by boundstone. Algal thalli form a dense carpet whose framework pores are filled with marine cement and peloidal micrite. The peloidal matrix is dense, partly laminated or clotted with irregular surfaces and often gravity defying. Algal reefs in Guizhou differ from examples reported to date by the high biodiversity of organisms other than phylloids: e.g., the intergrowth of algae with corals (some of which are twice the size of algal thalli) and numerous large brachiopods. This contrasts to previous views that phylloid algal “meadows” dominated the actual seafloor, excluding other biota. Also, the pervasive marine cements (up to 50%) including botryoidal cement are noteworthy. Algal reefs developed at platform margins, a depositional environment similar to that of modern Halimeda mounds in Java, Australia and off Bahamas, and to that of time-equivalent examples reported from the Canadian Arctic Archipelago. Whereas nutrients appear decisive in the growth of Halimeda reefs, algal reefs reported herein seemingly grew under conditions of low nutrient levels. Overall, algal reefs in Guizhou challenge previous views on growth forms, diversity patterns, and depositional environments and add to the spectrum of these partly puzzling biogenic structures.     

Seasonal structural and functional changes in the photosynthetic apparatus of evergreen conifers

We conducted a detail study of the photosynthetic apparatus in assimilating organs of three introduced evergreen conifer species: Taxus cuspidate S. et Z. ex E. (Far-Eastern yew), Thuja occidentalis L. (arbovitae “green”), and Th. occidentalis f. “Reingold” (arbovitae “yellow”) at various times in their life cycle. We studied the potential photosynthesis rate; composition and ratios of pigments, including primary carotenoids; the violaxanthin cycle (VC) activity, the synthesis of a secondary carotenoid, rhodoxanthin; and chloroplast ultrastructure. In winter and spring, β-carotene and lutein (primary carotenoids) contents were relatively constant in yew and arbovitae “yellow”. In December, the VC in yew was balanced and in arbovitae “yellow” unbalanced. In arbovitae “yellow”, the zeaxanthin pool was heterogeneous, and only part of it took part in the VC. It can be assumed that the other part of the pool can be oxidized to form a secondary carotenoid, rhodoxanthin. This secondary carotenoid was also accumulated in arbovitae “green”; its synthesis took place during the season, when the photosynthesis rate of plants was the lowest, and a significant chloroplast reorganization occurred (the number of thylakoids in grana decreased and plastoglobules appeared). We suppose that rhodoxanthin forms a filter for the light under the conditions of high insolation in winter. Thus, the evergreen conifer plants studied, which are adapted to growing at high latitudes where temperature is low and insolation is high in winter and spring, have a system for protecting the photosynthetic apparatus against photodestruction. In the basis of this system, the primary and secondary carotenoids lie, whose content changes during the year.     

Models for spatial polymerization dynamics of rod-like polymers

 We investigate the polymerization kinetics of rod-like polymer filaments interacting with a distribution of monomer in one spatial dimension (e.g. along a narrow tube). We consider a variety of possible cases, including competition by the filament tips for the available monomer, and behaviour analogous to “treadmilling” in which the polymer adds subunits to one end and loses them at the other end so as to maintain a constant length. Applications to biological polymers such as actin filaments and microtubules are discussed. Received: 16 March 1999     

Transforming Our Thinking about Transitional Forms

A common misconception of evolutionary biology is that it involves a search for “missing links” in the history of life. Relying on this misconception, antievolutionists present the supposed absence of transitional forms from the fossil record as evidence against evolution. Students of biology need to understand that evolution is a branching process, paleontologists do not expect to find “missing links,” and evolutionary research uses independent lines of evidence to test hypotheses and make conclusions about the history of life. Teachers can facilitate such learning by incorporating cladistics and tree-thinking into the curriculum and using evograms to focus on important evolutionary transitions.     

An initial attempt to map carbonate buildups using Ground-Penetrating Radar: An example from the Upper Jurassic of SW-Germany

Summary Although carbonate buildups host important hydrocarbon reservoirs and industrial minerals, relatively little is known about their three-dimensional architecture. Ground-penetrating radar (GPR, “Georadar”) studies provide the opportunity to reconstruct both their internal anatomy and external geometry. This short paper reports an initial attempt to map some Upper Jurassic sponge/algal buildups in the Upper Jurassic of SW-Germany using georadar. Several distinct radar facies types can be distinguished and calibrated to lithofacies in adjacent quarry outcrops. Mapping of the radar facies types reveals the potential to reconstruct the distribution of “buildup” versus other facies. Moreover, it highlights the need of obtaining more closely spaced radar lines in order to arrive at a high-resolution image.     

Macroevolution via secondary endosymbiosis: a Neo-Goldschmidtian view of unicellular hopeful monsters and Darwin’s primordial intermediate form

Seventy-five years ago, the geneticist Richard Goldschmidt hypothesized that single mutations affecting development could result in major phenotypic changes in a single generation to produce unique organisms within animal populations that he called “hopeful monsters”. Three decades ago, Sarah P. Gibbs proposed that photosynthetic unicellular micro-organisms like euglenoids and dinoflagellates are the products of a process now called “secondary endosymbiosis” (i.e., the evolution of a chloroplast surrounded by three or four membranes resulting from the incorporation of a eukaryotic alga by a eukaryotic heterotrophic host cell). In this article, we explore the evidence for Goldschmidt’s “hopeful monster” concept and expand the scope of this theory to include the macroevolutionary emergence of organisms like Euglena and Chlorarachnion from secondary endosymbiotic events. We argue that a Neo-Goldschmidtian perspective leads to the conclusion that cell chimeras such as euglenids and dinoflagellates, which are important groups of phytoplankton in freshwater and marine ecosystems, should be interpreted as “successful monsters”. In addition, we argue that Charles Darwin had euglenoids (infusoria) in mind when he speculated on the “primordial intermediate form”, although his Proto-Euglena-hypothesis for the origin of the last common ancestor of all forms of life is no longer acceptable.     

Proposal for an integrated approach for the assessment of cross-media aspects relevant for the determination of “best available techniques” bat in the european union

The EC Directive concerning integrated pollution prevention and control (IPPC Directive 96/61/EC) obliges all Member States to make an integrated assessment of the impacts on the environment “as a whole”, as regards granting permission for and the operation of environmentally relevant industrial installations. The determination of “Best Available Techniques” BAT plays an essential role in the material transformation of the IPPC-Directive. An integrated approach for the assessment of cross-media aspects of techniques for the determination of BAT is outlined in this paper, which is grounded on the basic concept of Life Cycle Assessment (LCA), emphasising the need for decision support. The proposed assessment approach is applied to a case study sinter production in an integrated iron and steel works, which forms the base for several recommendations concerning further research.     

Linking limitation to species composition: importance of inter- and intra-specific variation in grazing resistance

Short-term responses of producers highlight that key nutrients (e.g., N, P)—or combinations of these nutrients—limit primary production in aquatic and terrestrial ecosystems. These discoveries continue to provide highly valuable insights, but it remains important to ask whether nutrients always predominantly limit producers despite wide variation in nutrient supply and herbivory among systems. After all, predictions from simple food chain models (derived here) readily predict that limitation by grazers can exceed that by nutrients, given sufficient enrichment. However, shifts in composition of producers and/or increasing dominance of invulnerable stages of a producer can, in theory, reduce grazer limitation and retain primacy of nutrient limitation along nutrient supply gradients. We observed both mechanisms (inter- and intra-species variation in vulnerability to herbivory) working in a two-part mesocosm experiment. We incubated diverse benthic algal assemblages for several months either in the presence or absence of benthic macro-grazers in mesocosms that spread a broad range of nutrient supply. We then conducted short-term assays of nutrient and grazer limitation on these communities. In the “historically grazed” assemblages, we found shifts from more edible, better competitors to more resistant producers over enrichment gradients (as anticipated by the food web model built with a tradeoff in resistance vs. competitive abilities). However, contrary to our expectations, “historically ungrazed” assemblages became dominated by producers with vulnerable juvenile forms but inedible adult forms (long filaments). Consequently, we observed higher resource limitation rather than grazer limitation over this nutrient supply gradient in both “historically grazed” (expected) and “historically ungrazed” (not initially expected). Thus, via multiple, general mechanisms involving resistance to grazing (changes in species composition or variation in stage-structured vulnerability), producer assemblages should remain more strongly or as strongly limited by nutrients than grazers, even over large enrichment gradients. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.