Coenocorrelation: gradient analysis of fossil communities and its applications in stratigraphy

Coenocorrelation is the correlation of positions in a stratigraphic sequence with corresponding positions along a paleoenvironmental gradient through gradient analysis of fossil communities. By ordination of community samples, the distribution and abundance of taxa along a depth gradient can be translated into a continuous scale that accurately measures the gradient and thus makes possible analysis of facies change on a continuum. The procedure is tested using data on marine benthic invertebrates along ten time-parallel transects down an Ordovician basin slope (Trenton Group, New York). Reversals between transgression and regression revealed in coenocorrelation curves served nearly as well as bentonite beds in time-correlation of sections. Ordinations gave estimates of depositional strike in close agreement with estimates from physical indicators. Walther's Law of Facies is evidently a more rigorous, quantifiable generalization than it has been revealed to be through classificatory approaches to communities and environments.     

Clinal variation, episodic evolution, and possible parapatric speciation: the trilobite Flexicalymene senaria along an Ordovician depth gradient

Pygidial segmentation in the trilobite Flexicalymene senaria changes with time (as measured in relation to bentonite beds) and with depth (as measured along downslopc transects along bentomtc beds and as measured by gradient analysis of fossil assemblages) along a 'fossil' depth gradient in the Middle Ordovician Trenton Group in central New York State. The pattern of strictly temporal change resembles a 'punctuation' between 'equilibria'. This evolutionary episode, a 10⁶ year interval of continuous change, may represent parapatric speciation. The clinal gradient is strongest during this episode and is weak or not apparent outside it. Strictly temporal change takes place by way of temporal change in the cline's shape. □ Evolution, geographic variation and dines, gradient analysis. Middle Ordovician, morphometry, speciation, Trilobita.     

Late Devonian marine ecology and zoogeography of the central Appalachians and New York

Ecological and statistical analyses of the Late Frasnian (Late Devonian) fauna of the Java Group of New York reveals the presence of three benthic marine communities. Using sedimentological and stratigraphic criteria, the Java Group may be divided into five physical-environmental zones: nearshore marine, platform, prodelta, open shelf, and slope and basin. The Ambocoelia- Cariniferella, Tylothyris-Schizophoria . and C yrtospirifer-Douvillina Communities inhabited open shelf, outer platform - prodelta, and inner platform - nearshore habitats. respectively. Java species gradient curves overlap each other continuously, and species optima replace each other in a regular fashion along the onshore-offshore environmental gradient. No evidence for competitive exclusion between species or sharp habitat discontinuities (ecotones) is seen for Java ecosystems. The same three-fold division of communities, in equivalent habitats, can be seen in central Appalachian marine environments located approximately 500 km away along the ancient shoreline. The distribution of communities in this region appears to be directly related to the position and relative size of fluviodeltaic systems along the 500 km of ancient coastline. Palacoecology. communities, Braschiopoda. palaegeography. Devonian. Frasnian, Appalachians, New. York .     

Evolution in a cline: the trilobite Triarthrus along an Ordovician depth gradient

Head proportions in the trilobite Triarthrus change by small but significant amounts with both lime and environment along a 'fossil' depth gradient in the Trenton Group of central New York. The putative zone species T. becki (Denmarkian Stage) and T. eatoni (post-Denmarkian; = T. becki , new synonymy) intermingle throughout the roughly two million year long sequence studied. The two represent extreme morphs in a graded spatio-temporal cline, not distinct species. Transgression as much as evolution accounts for their stratigraphic zonation. Clinal variation is expressed in the dependence of anatomical measures on the position of samples on a downslope transect parallel to bentonite beds, and on ordination score for community samples, a measure of environment that applies over time. Anatomy changes with environment in basically the same way it changes with time, and the amount of change over two million years in a given environment roughly equals the amount of change with environment at a given time. Compensated to a standard environment, anatomical measures follow a continuous trend that suggests an evolutionary random walk with non-random, directional drift. The case of Triarthrus illustrates that directional anatomical variation can have important components that are not strictly temporal and evolutionary in the normal sense.     

Spatial variation in leaf nutrient traits of dominant desert riparian plant species in an arid inland river basin of China

Understanding how patterns of leaf nutrient traits respond to groundwater depth is crucial for modeling the nutrient cycling of desert riparian ecosystems and forecasting the responses of ecosystems to global changes. In this study, we measured leaf nutrients along a transect across a groundwater depth gradient in the downstream Heihe River to explore the response of leaf nutrient traits to groundwater depth and soil properties. We found that leaf nutrient traits of dominant species showed different responses to groundwater depth gradient. Leaf C, leaf N, leaf P, and leaf K decreased significantly with groundwater depth, whereas patterns of leaf C/N and leaf N/P followed quadratic relationships with groundwater depth. Meanwhile, leaf C/P did not vary significantly along the groundwater depth gradient. Variations in leaf nutrient traits were associated with soil properties (e.g., soil bulk density, soil pH). Groundwater depth and soil pH jointly regulated the variation of leaf nutrient traits; however, groundwater depth explained the variation of leaf nutrient traits better than did soil pH. At the local scale in the typical desert riparian ecosystem, the dominant species was characterized by low leaf C, leaf N, and leaf P, but high leaf N/P and leaf C/P, indicating that desert riparian plants might be more limited by P than N in the growing season. Our observations will help to reveal specific adaptation patterns in relation to the groundwater depth gradient for dominant desert riparian species, provide insights into adaptive trends of leaf nutrient traits, and add information relevant to understanding the adaptive strategies of desert riparian forest vegetation to moisture gradients.     

Community and species-performance patterns along an alpine poor-rich mire gradient

Abstract. Vegetation, water table depth and water chemistry of 16 peatlands in the southern Alps, Italy, were analysed in 115 sample plots. A poor-rich gradient could be detected along the first axis of a partial Detrended Canonical Correspondence Analysis. Both vegetation and hydrochemistry vary gradually along the gradient. Vascular plants have much broader niches along the gradient than bryophytes. Mosses (except Sphangnum) and hepatics have narrower niches than Sphagnum. The various species of Sphagnum segregate more clearly from each other along the moisture gradient than along the poor-rich one. The positions of species optima along the latter gradient largely reflect the ecological preferences of mire plants in peatlands with respect to nutrient status.     

Continuous benthic community change along a depth gradient in Antarctic shallows: evidence of patchiness but not zonation

Many shallow water polar communities demonstrate considerable change along a bathymetric gradient. However, it is currently unclear whether community change is generally continuous or discrete. To determine the nature of community change with depth, extensive photographic surveys at three sites at Adelaide Island, West Antarctic Peninsula, were conducted along a bathymetric gradient of 5–35 m depth. Macroalgae were largely absent at the sites, so only distinguishable macrofauna were counted and analysed. Faunal abundance was greatest at the shallowest stations of the depth transects, whilst richness at both species and phylum level increased with depth. Variability in community structure between replicate transects decreased with depth, so that assemblages at >25 m depth were more homogenous. Depth had a highly significant effect on total abundance, species richness and community structure, and it is likely that the frequency of ice disturbance, which also decreases with depth, drives this pattern. Overall, high variability between transects at each site was recorded, which suggested considerable patchiness at the scale of tens of meters. Community change along the bathymetric gradient was continuous and no evidence of discrete zones of assemblages was recorded.     

Midge (Chironomidae,Chaoboridae, Ceratopogonidae) assemblages and their relationship with biological and physicochemical variables in shallow,polymictic lakes

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Latitudinal gradients of associations between beta and gamma diversity of trees in forest communities in the New World

Aims We analyze two continental data sets of forest communities from across the New World to examine the latitudinal gradients of beta diversity after accounting for gamma diversity and the latitudinal gradient of gamma diversity after accounting for beta diversity.Methods Correlation and regression analyses were used to relate beta and gamma diversity to latitude along two latitudinal gradients in the New World (one including 72 forest sites located south of the equator and the other including 79 forest sites located north of the equator).Important findings Beta diversity and gamma diversity were negatively correlated with latitude. Beta diversity was strongly and positively correlated with gamma diversity (Pearson's correlation coefficient: 0.783 for New World North and 0.848 for New World South). When beta diversity was regressed on latitude and gamma diversity, 69.8 and 85.7% of the variation in beta diversity were explained, respectively, for New World North and New World South. When gamma diversity was regressed on latitude and beta diversity, 81.8 and 84.3% of the variation in gamma diversity were explained, respectively, for New World North and New World South. After statistically removing the relationship between beta and gamma diversity, latitude has weak or no relationships with beta and gamma diversity. However, strong positive correlations between beta and gamma diversity may not be considered as evidence of one driving the other along a latitudinal gradient.     

Spatial variability in the distribution of dominant shallow-water benthos at Adelaide Island, Antarctica

Studies from temperate and tropical regions have shown that variability in the distribution of benthos exists at different spatial scales. There are very few similar studies from polar systems, the shallows of which represent some of the most intensely disturbed habitats on the planet. Variability in the abundances of the five most common macrofauna was examined at three spatial scales, metres, tens of metres and hundreds of metres, in the shallows (5-25 m depth) of Adelaide Island, West Antarctic Peninsula. Whilst significant community change occurs along a depth gradient at the study sites, not all of the common species studied showed clear depth-related patterns of distribution. Furthermore, although abundance patterns varied between the organisms, consistent depth-related trends in the spatial scale contributing most to the variability were observed for four of the five species. For four species the relative importance of large-scale variability (between sites) decreased from 5 to 25 m depth, whilst small-scale variability (between replicates) increased along the depth gradient. Variation between sites is probably largely driven by ice disturbance, which becomes less frequent with depth. Conversely, small-scale patchiness is promoted by biological interactions, which become relatively more influential as community complexity and species richness increase along the depth gradient.     

川西卧龙岷江冷杉林土壤有机碳组分与氮素关系随海拔梯度的变化特征

土壤碳氮沿海拔梯度变化及其耦合关系是山地生态系统碳氮循环研究的重要内容。为分析不同土层土壤有机碳,土壤全氮及有机碳活性组分在海拔梯度上的分布规律及相互之间的耦合关系,选取亚高山物种岷江冷杉(Abies faxoniana)原始林为研究对象,以卧龙邓生野牛沟岷江冷杉原始林2920—3700 m的样地调查数据为基础,分析不同土层土壤碳氮及活性组分沿海拔的变化规律,总结土壤有机碳稳定性沿海拔主要规律,从土壤有机碳活性组分和碳氮关系的角度揭示其对土壤有机碳沿海拔变化的影响。结果表明:1)腐殖质层土壤有机碳(SOC)随海拔升高逐渐增加,与温度显著负相关,轻组有机碳(LFOC)及颗粒态有机碳(POC)随海拔上升均表现先增加后降低的趋势,土壤全氮(TN)随海拔变化不显著,但林线处LOFC、POC和TN均显著增加;0—10 cm土壤有机碳及全氮则表现为双峰特征,峰值分别在3089 m和3260 m处,与年均温度无显著关系。2)LFOC及POC在腐殖质层和0—10 cm土层中所占比例较大,是表征土壤有机碳含量沿海拔变化规律的主要活性组分,腐殖质层LFOC/SOC和POC/SOC随海拔上升逐渐增高,0—10 cm层则逐渐降低,暗示腐殖质层有机碳稳定性沿海拔逐渐降低,0—10 cm有机碳稳定性逐渐升高。3)SOC与TN显著正相关,SOC是影响TN的主要因子,但腐殖质层TN与有机碳活性组分无显著相关关系。4)土壤C/N和微生物量C/N在3177 m大于25:1,是引起土壤有机碳含量显著降低的主要因素。     

Imperfect detection distorts depth‐related trends in marine macrofaunal species richness

Ecology and biodiversity research are underpinned by species richness patterns and their environmental drivers. However, a key topic in this discussion is the accuracy of these patterns which are greatly dependent on species detection probabilities. Due to variations in detection of species, true ecological patterns may be distorted. This is particularly true for subtidal macro‐infaunal communities. We tested three hypothesized relationships between marine benthic macrofaunal diversity and depth using species richness per site estimated with a capture–recapture heterogeneity model that accounts for variable detection probabilities. These metrics were based on data from 42 replicated sites across the continental shelf of the Southern Benguela. Average detection probability decreased with greater depth but species richness increased along the same depth gradient. The conflation of these trends in observed diversity data resulted in a positively near‐linear depth–diversity relationship, while accounting for variable species detection revealed a much stronger relationship. Ignoring species detection in ecosystems with imperfect detection could therefore distort species richness patterns, which has implications for ecological theory, management and conservation.     

Geographic and stratigraphic change in the morphology of Triarthrus beckii (Green) (Trilobita): a test of the Plus ça change model of evolution

The Plus ça change model predicts that deepwater trilobite species such as Triarthrus should exhibit gradual phyletic evolution. A detailed stratigraphic sequence of Triarthrus beckii specimens considered together with geographically separated samples from a single time interval provide a test of the Plus ça change model. We examined geographic patterns of variation in cranidial shape based on specimens from four approximately synchronous levels within the Upper Ordovician (lower Edenian) strata of New York, Québec, Kentucky and Pennsylvania. All geographic populations differ in mean size, except for New York and Pennsylvania. Because allometry is present in both meraspid and holaspid phases of the trilobite, size effects on shape were removed by ontogenetic standardization. Cranidial shapes were then compared among the four study sites and eight stratophenetic samples from New York, obtained by subdividing our Mohawkian Composite Standard Section into 20-m-thick intervals. Goodall's F-test of pairwise comparisons of cranidial shape between eight subintervals in New York and the other three populations are all significant. Geographic variation in cranidial size and shape in T . beckii appears to display a gradient or cline-like pattern that is related to paleogeography. Although cranidial shape is not entirely static within the New York stratophenetic series, it is in all cases different from that sampled elsewhere in the region. Thus, there is no evidence of wholesale immigration and emigration among the geographic areas. Instead, the geographic gradient of T . beckii shape variation may have remained fairly stable over ca. 3 million years during the mid-Chatfieldian to early Edenian interval. This result is indicative of stasis. Consequently, the evolutionary history of T . beckii contradicts the Plus ça change model.     

Topography and tectonics of the Taconic outer trench slope as revealed through gradient analysis of fossil assemblages

Similarities in stratigraphic sequence, syndepositional topography and tectonics, and regional tectonic setting between the Mohawk Valley, New York, in the Middle Ordovician and the Australian flank of the Timor Trough today support the idea that the Taconic Foreland Basin orginated in an arc-continent collision. The Trenton Group strata studied - a roughly four million year long sequence correlated by bentonite beds along an 83 km downslope transect in the central Mohawk Valley - represent the continental shelf and outer trench slope. Bank limestones pass upward and downslope into deep water black shales in a manner reminiscent of the lateral and vertical sedimentary sequence on the Sahul Shelf and Australian flank of the Timor Trough. In studying topography, relative depth was measured through reciprocal averaging ordination of benthic macroinvertebrate fossil assemblages. Downslope bathymetric profiles show the development of a horst and graben topography coincident with regional tilting and transgression - a pattern associated with downward flexure of the Australian Plate approaching Timor. The net rate of transgression corresponds to an arc-continent convergence rate on the order of 2 cm per year.     

Plant biomass and species composition along an environmental gradient in montane riparian meadows

In riparian meadows, narrow zonation of the dominant vegetation frequently occurs along the elevational gradient from the stream edge to the floodplain terrace. We measured plant species composition and above- and belowground biomass in three riparian plant communities—a priori defined as wet, moist, and dry meadow—along short streamside topographic gradients in two montane meadows in northeast Oregon. The objectives were to: (1) compare above- and belowground biomass in the three meadow communities; (2) examine relations among plant species richness, biomass distribution, water table depth, and soil redox potential along the streamside elevational gradients. We installed wells and platinum electrodes along transects (perpendicular to the stream; n=5 per site) through the three plant communities, and monitored water table depth and soil redox potential (10 and 25 cm depth) from July 1997 to August 1999. Mean water table depth and soil redox potential differed significantly along the transects, and characterized a strong environmental gradient. Community differences in plant species composition were reflected in biomass distribution. Highest total biomass (live+dead) occurred in the sedge-dominated wet meadows (4,311±289 g/m²), intermediate biomass (2,236±221 g/m²) was seen in the moist meadow communities, dominated by grasses and sedges, and lowest biomass (1,403±113 g/m²) was observed in the more diverse dry meadows, dominated by grasses and forbs. In the wet and moist communities, belowground biomass (live+dead) comprised 68–81% of the totals. Rhizome-to-root ratios and distinctive vertical profiles of belowground biomass reflected characteristics of the dominant graminoid species within each community. Total biomass was positively correlated with mean water table depth, and negatively correlated with mean redox potential (10 cm and 25 cm depths; P <0.01) and species richness (P <0.05), indicating that the distribution of biomass coincided with the streamside edaphic gradient in these riparian meadows.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Epeiric sedimentation and sea level: synthetic ecostratigraphy

Carbonate strata from the central parts of epicontinental seas are ideal for detailed biostratigraphic study of eustatic sea level change. Using a model for epeiric seas in which carbonate accumulation rate is depth-dependent, we derive synthetic stratigraphies for sea level histories simulating post-glacial transgression and constant and sinusoidally fluctuating ridge volume increase. These sea level histories give distinctively different trends for water depth as a function of stratigraphic position in sections' bathymetric curves. In general, depth is proportional to the rate of sea level rise. Depth-dependent sedimentation leads to a time lag between sea level fluctuation and corresponding depth fluctuation which, as examples show, can approach 10⁶ years for depth fluctuations of only a few meters – a fundamental consideration in reconstructing sea level curves, time-correlating sections by their bathymetric curves, and attempting to relate bathymetric history on continents to mechanisms driving sea level change. Bathymetric curves based on gradient analysis of fossil assemblages (coenocorrelation curves) for Middle Ordovician sections in New York and the American Midwest approximate patterns for sinusoidally increasing sea level. The model's predictions are tested in an ‘artificial experiment’ that takes advantage of differential subsidence between the craton's middle and its edge to make a difference in the bathymetric histories of sections that otherwise record the same sea level history. Depth fluctuations of no more than a few meters over million year spans are potentially useful in time-correlation to within fractions of a cycle's period. The depth-dependence in sedimentation was that, above wave base, net accumulation per year was very roughly three-millionths the water depth. Association of volcanic ash layers with transitory sea level minima on the craton, and with onset of more rapid subduction in the Taconic are - continent collision zone, suggests interrelationship on a million year scale between sea level and large-scale tectonic phenomena.     

Depth-related variation in gross chemical composition of several seaweeds

The percentages of dry weigth, ash, carbon and nitrogen in Chondrus crispus Stackh. and Furcellaria lumbricalis (Huds.) Lamour. were determined for plants distributed along a depth gradient. Analyses were made of samples collected from June throug August. Dry weights and ash were also determined for other major perennial species, but on fewer samples. In all instances the trend was an increase in ash as a percentage of the dry weight and a decrease in the percentage dry weight and organic matter, with depth. The percentage of carbon also decreased with depth, reflecting the changes in organic matter. However, the percentage of nitrogen increased with depth, thus the C/N ratio decreased with depth. These changes in gross chemical composition are assumed to be analogous to seasonal changes in composition at a given depth or level in the littoral zone.     

Changes in scleractinian coral <Emphasis Type="Italic">Seriatopora hystrix</Emphasis> morphology and its endocellular <Emphasis Type="Italic">Symbiodinium</Emphasis> characteristics along a bathymetric gradient from shallow to mesophotic reef

The algae living endosymbiotically within coral are thought to increase algal pigmentation with increasing depth to capture the diminishing light. Here, we follow distribution of the hermatypic coral Seriatopora hystrix along a 60-m bathymetric gradient in the Gulf of Eilat, Red Sea, to study coral ecophysiology and response to light regimes. Combining work on coral morphology, pigment content and genotyping of the photosymbiont, we found that total chlorophyll concentration per zooxanthellae cell and the dark- and light-acclimated quantum yield of photosystem II did not vary significantly along the 60-m gradient. However, the chlorophyll a/c ratio increased with depth. This suggests that the symbiotic algae in S. hystrix possess a mechanism for acclimatization or adaptation that differs from previously described pathways. The accepted photoacclimatory process involves an increase in chlorophyll content per alga as light intensity decreases. Based on corallite and branch morphology, this research suggests that S. hystrix has two depth-dependent ecophenotypes. Above 10 m depth, S. hystrix exhibits sturdier colony configurations with thick branches, while below 30 m depth, colonies are characterized by thin branches and the presence of a larger polyp area. Between 10 and 30 m depth, both ecophenotypes are present, suggesting that corallite morphology may act as another axis of photoacclimation with depth.     

Distribution of arbuscular mycorrhizal fungi in stands of the wetland grass Panicum hemitomon along a wide hydrologic gradient

Although wetland plant species usually aggregate into zones that correspond with their water depth/dryness tolerances, it is not known whether associated arbuscular mycorrhizal (AM) fungi show a similar zonation. We assessed the distribution of AM fungi in two similar depressional wetlands dominated by the semi-aquatic grass Panicum hemitomon by sampling soil in plots along dry-to-wet gradients that spanned 80 cm in relative elevation, and identifying/counting viable AM fungal spores. We found that eight of nine AM fungal species were common to both of the wetlands. Within each wetland, there were significant differences in species composition related to relative water depth. The zonation patterns were not identical between wetlands but revealed that certain species were relegated to the drier portions of the gradient in both. No species were relegated to only the wet portions of the wetlands; those that dominated there were also present in the drier areas. Our data show that water depth is an important factor determining the distribution of the AM fungi, even when, as in our study wetlands, the host plant remains constant along a dry-to-wet gradient. This suggests that the fungi are not physiologically equivalent in their tolerance to wetland conditions. Received: 9 October 1998 / Accepted: 24 February 1999