Early Emsian ammonoids from the eastern Anti-Atlas (Morocco) and their succession

From 20 sections in the Tafilalt (eastern Anti-Atlas, Morocco), 13 early Emsian ammonoid species of the families Mimosphinctidae, Mimoceratidae, Teicherticeratidae, Mimagoniatitidae, and Auguritidae have been discovered. The following taxa are newly described:Chebbites n. gen.,Chebbites reisdorfi n. sp.,Chebbites undosus n. sp.,Gracilités maghribensis n. sp., Palaeogoniatitinae n. subfam.,Irdanites n. gen.,Irdanites kaufmanni n. sp.,Irdanites korni n. sp.,Lenzites gesinae n. sp.,Weyeroceras n. gen.,Weyeroceras angustus n. sp. In addition, specimens ofErbenoceras solitarium, Metabactrites formosus, andRuanites obliquecostatus are described and figured from Morocco for the first time. The ammonoid generaConvoluticeras, Erbenoceras, Mimosphinctes, Palaeogoniatites, Talenticeras, andTeicherticeras and the families Mimosphinctidae, Mimoceratidae, Teicherticeratidae, and Mimagoniatitidae are reviewed.Taskanites andLaganites are regarded as subjective junior synonyms ofConvoluticeras. In Morocco, the early Emsian ammonoid succession differs from what can be expected from a phylogenetic point of view. Several species belonging to different families of various phylogenetic stages appear within one stratigraphical unit. In contrast to previous opinions however,Gyroceratites laevis appears stratigraphically earlier thanMimagoniatites fecundus, which agrees in this case with the phylogenetic succession of the taxa. Stratigraphical data from early Emsian ammonoids are still insufficient as basis for a more detailed stratigraphical subdivision of the early Emsian.     

华北高产农田生态系统中蚯蚓种群次生演替规律

通过对山东省桓台县不同土壤肥力的农田生态系统中蚯蚓种群的调查发现 ,该地区农田生态系统中有 7种蚯蚓 ,高肥力土壤中蚯蚓相对较丰富 ,种群密度可达 83.83条 / m2 ,梯形流蚓为优势种 ;低肥力土壤中蚯蚓种群密度 4 0 .18条 / m2 ,优势种是天锡杜拉蚓。梯形流蚓和湖北远盲蚓在高肥力土壤中的种群密度要明显高于低肥力土壤 ;而其他几种蚯蚓种群数量变化不大 ,随着土壤肥力的演变 ,低肥力土壤中天锡杜拉蚓的优势地位随着土壤肥力的提高逐渐被梯形流蚓所代替。合理的投入特别是农田有机物投入可以加速农田生态系统中生态演替 ;试验中不同处理间蚯蚓的种群生物量有以下趋势 :化肥投入 <化肥与麦秸还田 <化肥与玉米麦秸全还田 <化肥与玉米麦秸全还田以及有机肥的施用。无论在高肥力还是低肥力的土壤上都表现出相同的规律 ,但蚯蚓种群组成并没有明显差异     

The mode of life in ammonoids

The following structural features clearly indicate that ammonoid shells were adapted to withstand considerably higher hydrostatic pressures thanNautilus shells: (1) the corrugated and marginally fluted septa gave the shell wall efficient support against implosion; (2) the secondary connecting rings could grow a great deal in thickness; and (3) the last formed chambers remained full of liquid which supported the last septum. On the basis of the following characters it is concluded that ammonoids were incapable of swimming efficiently by jet-propulsion: (1) the retractor muscles were weakly developed; (2) the life position was unstable and highly variable; and (3) in animals with a ventral apertural rostrum the hyponome was probably absent. Ammonoids are considered here as having been pelagic cephalopods which lived in the upper 1000 m of the oceans, and which probably undertook considerable diurnal vertical migrations, similar to those inSpirula. Only some groups may have adopted a life in shallow epicontinental seas. In the late Mesozoic, ammonoids have been replaced by modern oceanic squids which are extremely numerous in the corresponding pelagic environment.     

Durophagous predation on scaphitid ammonoids in the Late Cretaceous Western Interior Seaway of North America

This study is the first to report a trend of predation intensity on scaphitid ammonoids from the Turonian to the Maastrichtian (Late Cretaceous) on the basis of analysis of ventral shell breakage in large samples from the US Western Interior Province. Analysis of 835 adult specimens revealed ventral shell breakage in 50 specimens. In most of the damaged specimens, the breakage occurred in a preferred position at the rear part of the body chamber. Ventral breakage is rare in the Turonian specimens, whereas it is common in the Campanian and Maastrichtian specimens. The shell diameter of adult scaphitid ammonoids tends to increase with time. The position of the breakage and the absence of repairs indicate that the ventral breakage resulted from lethal predation. Based on the incidence of breakage and the size and shape of the breaks, possible predators include fish, reptiles and cephalopods such as Placenticeras, Eutrephoceras and coleoids. Our statistical analysis of ventral shell breakage indicates that the incidence of lethal predation increased in conjunction with an increase in adult shell size, suggesting that the body size of the prey was an important factor in predator–prey interactions. In addition, the predatory damage is more extensive in larger adults.     

The A-mode sutural ontogeny in prolecanitid ammonoids

The generally accepted theory of a U-mode sutural ontogeny of prolecanitid ammonoids and their descendants is refuted. The basic suture formula of the order Prolecanitida is E A L U I (not E L U₂ U₁ I), resembling that of derived members of the suborder Tornoceratina (their phylogenetic ancestors), and of the suborder Goniatitina. During phylogeny of the prolecanitids, secondary umbilical lobes are introduced and lead to multilobate forms. In early ceratites, the original L lobe disappeared, and an increase in sutural elements took place by the introduction of supplementary U lobes. Consequently, sutural nomenclature of Permian ceratites and Mesozoic ammonoids has to be modified.     

末次盛冰期以来沙漠-黄土过渡带植被演替及其对气候变化的响应

末次盛冰期(LGM)以来的增温期和全新世高温期是人类古文化进步的重要时期。黄土高原北部沙漠-黄土过渡带两个自然剖面高分辨率花粉分析结果显示,该区LGM以来为草原植被景观,但优势群落和伴生植物随气候变化而有所不同,温度和降水变化对植被演替有着重要的控制作用。气候干冷时,以耐干旱的草本植物种属为主,群落成分单一,植被盖度低;增温期和高温期,出现少量暖温带乔木植物如胡桃科、栎属等,植被盖度高,群落发育良好。全新世暖期是植被发育良好、植被盖度高的适宜期。本研究为黄土高原地区未来增温下植被响应的分析提供了历史相似型。     

The genus Luppovia and the phylogeny of Cretaceous heteromorphic ammonoids

The unique preservation of heteromorphic shells of Luppovia Kakabadze, Bogdanova & Mikhailova 1978 from the Aptian deposits of Bolshoj Balkhan (western Turkmenia) made it possible to study the microstructure, internal shell characteristics and suture ontogeny of this genus. Microstructural investigations revealed that the dorsal shell wall in Luppovia is composed of three layers in contrast to that of monomorphic ammonoids, which is single-layered. This ability of secreting different numbers of layers in monomorphs and heteromorphs demonstrates plasticity of secreting activity of the mantle epithelium. Data obtained from the study of internal shell structure and suture ontogeny have been compared. Both these methods of investigation lead to the conclusion that the genus Luppovia belongs to the order Ammonitida. The results obtained confirm the viewpoint that the Cretaceous heteromorphic superfamilies Turrilitaceae and Ancylocerataceae have independent origins and belong to two different orders, Lytoceratida and Ammonitida, respectively. □ Ammonoidea , Luppovia, shell structure, ontogeny, phylogeny, Cretaceous.     

Seasonal succession in the pelagic ecosystem of the North Atlantic and the utilization of nitrogen

Observations during the Joint Global Ocean Flux Study (JGOFS)North Atlantic Bloom Experiment in 1989 are compared with amixed-layer model of phytoplankton seasonal succession in whichthe latitudinal variation of the succession is driven by physicalforcing. In the model, the first phytoplankton groups to growat the end of the winter are those with the fastest intrinsicgrowth rates: the phytoflagellates and picophytoplankton. Theincreases of these groups are soon halted by the rapid growthof microzooplankton and heteroflagellates whereupon they aresucceeded by diatoms, the next fastest growers. With depletionof silicate, the diatom bloom ends and is briefly replaced byphytoflagellates and picophytoplankton, before these in turnare replaced by slower growing dinoflagellates. Differencesin the physical forcing cause the timings and magnitudes ofthese changes to vary with latitude. By sampling the model resultsat the times and places of the JGOFS observations, it is shownthat the major changes of populations and nutrients are reproduced,as are many production and grazing rates. The model resultssuggest the nature of nutrient utilization in the region. Whilenitrate and silicate are both reduced to low values at 47N,only silicate is depleted at 60N. Nitrate is not depleted atnorthern latitudes due to the greater depth of the mixed layer,more intense vertical mixing and the shorter season, so thatphytoplankton have more nutrients to utilize and a shorter timein which to do it. Phytoflagellates and picophytoplankton areunable to utilize all the inorganic nitrogen because of thegrazing by the micrograzers, and diatoms because of silicatedepletion. Dinoflagellates are slow growing and only have sufficienttime to deplete the nitrogen at low latitudes. There is no needto invoke limitation by a trace nutrient, such as iron, to reproducethe events in the NE Atlantic during 1989.     

中国与北极—北极纪行(一)

中国极地考察起步是不平衡的,曾经重南极而轻北极。不仅在南极建立了长城站和中山站,而且自1984年以来,每年都派考察队赴南极考察,取得了一系列重大科学成果。对比而言,我国的北极考察要弱一些,至今没有设考察站,也很少组队前往北极。我国作为北半球国家,相对来说北极列我国的影响要大于南极,仅气候影响就非常明显。为了扭转这一局面,近年来我国的北极考察被提上日程,有的组队到达北极点,有的科学家乘船赴北极冰区考察。现在,北极斯瓦尔巴德群岛又进入中国考察队视野,已经成为中国北极考察基地。2001年10月,新华社记者张继民随科学探险队到达了斯瓦尔巴德群岛,感于北极独特的风情,写出了他的所见所闻,特刊出。     

Quantification of ontogenetic allometry in ammonoids

Ammonoids are well‐known objects used for studies on ontogeny and phylogeny, but a quantification of ontogenetic change has not yet been carried out. Their planispirally coiled conchs allow for a study of “longitudinal” ontogenetic data, that is data of ontogenetic trajectories that can be obtained from a single specimen. Therefore, they provide a good model for ontogenetic studies of geometry in other shelled organisms. Using modifications of three cardinal conch dimensions, computer simulations can model artificial conchs. The trajectories of ontogenetic allometry of these simulations can be analyzed in great detail in a theoretical morphospace. A method for the classification of conch ontogeny and quantification of the degree of allometry is proposed. Using high‐precision cross‐sections, the allometric conch growth of real ammonoids can be documented and compared. The members of the Ammonoidea show a wide variety of allometric growth, ranging from near isometry to monophasic, biphasic, or polyphasic allometry. Selected examples of Palaeozoic and Mesozoic ammonoids are shown with respect to their degree of change during ontogeny of the conch.     

Biogeographic synthesis of secondary succession rates in eastern North America

Aim Mechanistic models of old‐field plant succession have generally lacked a broader biogeographic context in which climate, soils and species pools could play a significant role in mediating succession rates. We examine broad‐scale patterns in old‐field succession and introduce a hierarchical conceptual model to incorporate potential mechanisms operating at multiple spatial scales. Location We reviewed secondary succession studies across the Eastern Deciduous Forest (EDF) of North America. Methods We collected data from all published studies that reported the time necessary for woody species to reach either 10% or 50% cover in old‐field systems throughout the EDF. We used regression approaches to determine whether rates of succession are controlled by climatic and edaphic factors that vary at broad spatial scales. Results We found that the rate at which woody species colonize and dominate old fields decreases significantly with latitude. Rates of woody succession were highly correlated with both annual temperature (growing degree‐days for years to 10% and 50% cover) and measures of soil fertility (cation exchange capacity and reported pre‐agroindustrial (1930) maize yields for years to 10% cover), all of which show a significant latitudinal trend across the EDF. Main conclusions We suggest that the driver of this pattern is geographic variation in the intensity of competition between herbaceous and woody species, which we relate to the effects of (1) temperature regime, (2) edaphic factors related to soil fertility, and (3) plant traits, which may vary latitudinally in response to climate and edaphic factors. Although insufficient data exist to evaluate the relative importance of these factors, we argue that research in this area is necessary to gain an understanding of how future landscapes will be affected by global climate and land use change.     

Ventral bite marks in Mesozoic ammonoids

Reports on the predators of ammonoids are rare, although ammonoids were abundant and diverse invertebrates in many Paleozoic and Mesozoic marine ecosystems. Most previous work on lethal ammonoid predation has focused on (sub)circular tooth marks which resulted from fish and mosasaur attacks. In the present study we discuss a relatively common type of bite mark in ammonoid shells, the ‘ventral bite mark’. This typically occurs in a restricted position on the ventral side of the outer body chamber whorl and does not affect either the aperture or the phragmocone. Ammonoid specimens revealing ventral bite marks used in this study were collected from a wide range of strata which range in age from the Lower Jurassic to the uppermost Cretaceous (close to the Cretaceous–Paleogene boundary). These ventral bite marks are absent in the Paleozoic collections studied. The vast majority of ventral bite marks are situated at the end of the body chamber, close to the phragmocone. This is interpreted as the result of predatory attacks on the back or blind side of ammonoids in their living position. The predators aimed for the vital parts and muscle attachments to obtain the edible soft tissues. The agents for most of the ventral bite marks to ammonoids are probably coleoid cephalopods (especially teuthoids) and predatory fishes to a lesser extent.     

Life-cycles of some Devonian ammonoids

Whorl expansion rates of six representative ammonoid genera from late Emsian and Eifelian strata of Morocco were calculated for each whorl. The corresponding body chamber lengths and the orientations of the apertures were computed based on these values. The resulting body chamber length and orientation of the aperture graphs were compared with other conch features, ecology of Recent cephalopods, and sedimentological data of the host rocks in the Tafilalt (eastern Anti-Atlas, Morocco). A subdivision of the ontogeny of these ammonoids was achieved comprising the early and late embryonic periods, the juvenile period, the preadult, and the adult growth period. All growth periods are defined by specific changes in growth, conch morphology, and mode of life. According to this reconstruction, hatchlings were probably already capable of active movements. Differentiation in two main modes of life of the examined taxa occurred in the late juvenile or early preadult period. As preadult animals, most of the Mimagoniatitoidea and Agoniatitoidea became active swimmers (Nektonic), whereas the representatives of the Anarcestoidea were capable of slow movements only (Planktonic). As adults, most representatives of the three superfamilies had an approximately horizontally oriented aperture, allowing active swimming and possibly active choice of spawning sites. Additionally, the new ammonoid taxon Rherisites tuba gen. nov., sp. nov. from the late Emsian is introduced.     

Role of heterochronies in the morphogenesis of paleozoic ammonoids

Various types of evolutionary changes related to shifts in the appearance or order of formation of characters, i.e., heterochronies are shown. The most common such changes in Paleozoic ammonoids was the appearance of mosaic forms combining conservative and advanced characters resulting from recapitulation and paedomorphosis. These phenomena are related to certain phylogenetic stages in each group: the growth stage predominantly showed recapitulations; the diversification stage showed the appearance of many mosaic forms; the decline stage commonly shows paedomorphosis connected with retardation.     

Simulated dynamics of succession in a North American subtropical Prosopis savanna

Abstract. A transition matrix model was used to explore the dynamics, rate and potential extent of changes in landscape vegetation patterns on a southern Texas Prosopis savanna. Transitions between seven vegetation classes were determined for the periods 1941–1960 and 1960–1983 on aerial photographs of three sites. During these periods, the sites were heavily grazed by cattle and were fire-free. Vegetation states assessed in grids of 20 m x 20 m cells superimposed on photographs ranged from grass-dominated to woody plant-dominated. The 1941–1960 period (denoted DRY) was characterized by prolonged drought, whereas annual rainfall during the 1960–1983 period (denoted WET) was typically normal to above-normal. The 1941 landscape consisted of herbaceous zones (6% of cells), woodland (50% of cells) and savanna parkland (44% of cells with grass/woody plant mixtures). The woodland state was the most stable, with probabilities of no change being 0.970 and 0.873 in WET and DRY periods, respectively. The herbaceous state was least stable, with corresponding values of 0.074 and 0.353. Past and future landscape structure was modelled by randomly selecting DRY or WET transitions at 20 year time steps. The model was run under a series of rainfall scenarios where the probability of selecting the WET transition matrix (P[WET]) ranged from 0 (DRY always chosen) to 1 (WET always chosen). Historical records indicate P[WET] has approximated 0.3 to 0.4 in the region. The rate of succession to states of greater woody cover increased as P[WET] increased. Forward simulations based on P[WET] > 0.2 suggest the present landscape is unstable and will develop into a closed-canopy woodland within the next 180 years, assuming the processes operating between 1941 and 1983 continue (e.g. grazing by cattle and lack of fire). Reverse simulations concur with historical observations and projections derived from woody plant growth rates in other studies and suggest that 200 to 300 yr BP these landscapes contained a substantially greater proportion of cells dominated by grassland or grassland with scattered woody plants (43 to 74%) than was present in 1983 (19%). Based upon elapsed time between predicted past and future steady states, succession from open savanna to closed-canopy woodland may occur in ca. 400 to 500 yr for P(WET) ≥ 0.33. Arresting or reversing the projected trend may require changes in climate and/or changes in livestock grazing and land management practices. The approaches employed in this study illustrate how time series maps, aerial photographs and satellite imagery can be analyzed and used to interpret, project and reconstruct local and regional changes in ecosystem structure. Difficulties and limitations associated with the use of Markov chains to model succession are identified and discussed.     

Major trends in the evolution of Permian ammonoids

The phylogeny of major families of Permian ammonoids is analyzed. The evolution of most families followed a typical scenario with distinct stages of early evolution, diversification, and decline. A smaller group followed a different evolutionary narrative, with indistinct stages. The former group includes families with both simple and complex morphology and a wide range of variation. The nature and trends in the evolution of the families may change depending on their phylogenetic stage. The Early Permian (Asselian), the second half of the Artinskian, and the beginning of the Middle Permian were marked by the most significant evolutionary changes. The Late Permian was the time of the decline of Paleozoic ammonoid orders and of the onset of the evolution of the Mesozoic order Ceratitida.     

Soft-part anatomy of the siphuncle in Permian prolecanitid ammonoids

Exceptionally well-preserved remains of phosphatized siphuncles were discovered in four specimens of a Permian prolecanitid ammonoid Akmilleria electraensis (Plummer and Scott) from Buck Mountain, Nevada. These structures occur as truncated segments within the siphuncular tube. The outer surface of the siphuncle is sculptured by numerous equally spaced longitudinal ridges and furrows; the ridges represent an infolded basement membrane of epithelial cells which corresponds to the distal ends of individual canaliculi between epithelial cells. In cross-section, the siphuncle of A. electraensis consists of a large central vein, possibly two pairs of arteries, porous connective tissue with reticulate hemocoelic spaces, and a thin epithelium. In the presence of two pairs of arteries and porous connective tissue, the siphuncle of A. electraensis is more like that of Nautilus pompilius than that of Spirula spirula, which has nine arteries and dense connective tissue. However, Nautilus possesses relatively smaller and more numerous epithelial cells around the siphuncle than does Akmilleria. These observations strongly suggest that the siphuncular epithelium of Akmilleria served as the salt-­concentrating organ for buoyancy regulation of the living animal, just as in Nautilus and Spirula.     

Origin of intracameral sheets in ammonoids

The distribution, morphology and mutual relationships of cameral sheets in ammonoids are revised and re-evaluated. Taking into account recent models of ammonoid septum and chamber formation, three different origins can be attributed to the morphological types of sheets: (1) membranes replicated by the rear mantle (pseudosepta and septal linings), (2) membranes secreted sequentially and/or stretched across the chamber (horizontal membranes and chamber linings) and (3) products of desiccation of the cameral liquid (transverse and siphuncular sheets), presumably a cameral hydrogel. Sheets are always preserved near the siphuncular area, because as the cameral liquid was pumped out from the chamber it became progressively richer in dissolved mucus. In the last-formed drops, or menisci, this mucus adhered to the surface of the previously secreted sheets and, on dehydration, it also replicated the surface of the residual reservoirs, producing desiccation sheets. On the basis of the new evidence, changes in the shape of the rear mantle in Triassic ammonoids can be reconstructed. In general, deformations affected the rounded or bottle-neck saddles, which deflated after detachment of the last-formed septum and reinflated when the position of the next septum was reached. The rest of the elements of the septal epithelium were affected to a much lesser extent. One of the functions of those cameral sheets secreted by the rear body was related to a more efficient transport of the cameral liquid upon decoupling from the siphuncular tube. Ammonoids, Triassic, septum, chamber growth, cameral sheets.     

Soil formation under progressive accelerated succession of forest ecosystems in the North of Russia

The paper presents experimental data on restoration of soil, an ecosystem component. Each trial plot was treated differently. It is shown that the use of a combination of techniques, namely grass sowing and mineral and organic fertilization of soil, aimed at restoration of biological rotation of plants, radically intensifies the soil restoration. The developed organic-accumulating layer is the main indicator of successful soil restoration. The importance of soil as a structural element of an ecosystem is determined by its capacity to retain and accumulate nutrients thus ensuring stable conditions for self-restoration of ecosystem components.