Matrix models for quantifying competitive intransitivity from species abundance data

In a network of competing species, a competitive intransitivity occurs when the ranking of competitive abilities does not follow a linear hierarchy (A > B > C but C > A). A variety of mathematical models suggests that intransitive networks can prevent or slow down competitive exclusion and maintain biodiversity by enhancing species coexistence. However, it has been difficult to assess empirically the relative importance of intransitive competition because a large number of pairwise species competition experiments are needed to construct a competition matrix that is used to parameterize existing models. Here we introduce a statistical framework for evaluating the contribution of intransitivity to community structure using species abundance matrices that are commonly generated from replicated sampling of species assemblages. We provide metrics and analytical methods for using abundance matrices to estimate species competition and patch transition matrices by using reverse-engineering and a colonization-competition model. These matrices provide complementary metrics to estimate the degree of intransitivity in the competition network of the sampled communities. Benchmark tests reveal that the proposed methods could successfully detect intransitive competition networks, even in the absence of direct measures of pairwise competitive strength. To illustrate the approach, we analyzed patterns of abundance and biomass of five species of necrophagous Diptera and eight species of their hymenopteran parasitoids that co-occur in beech forests in Germany. We found evidence for a strong competitive hierarchy within communities of flies and parasitoids. However, for parasitoids, there was a tendency towards increasing intransitivity in higher weight classes, which represented larger resource patches. These tests provide novel methods for empirically estimating the degree of intransitivity in competitive networks from observational datasets. They can be applied to experimental measures of pairwise species interactions, as well as to spatio-temporal samples of assemblages in homogenous environments or environmental gradients.

Ground beetle (Carabidae) abundance data from north-eastern Australian rainforest, between June 2008 - January 2010, using pitfall traps

Understanding how the environment influences patterns of diversity is vital for effective conservation management, especially in a changing global climate. While assemblage structure and species richness patterns are often correlated with current environmental factors, historical influences may also be considerable, especially for taxa with poor dispersal abilities. Mountain-top regions throughout tropical rainforests can act as important refugia for taxa characterised by low dispersal capacities such as flightless ground beetles (Carabidae), an ecologically significant predatory group. We surveyed flightless ground beetles along elevational gradients in five different subregions within the Australian Wet Tropics World Heritage Area to investigate (1) whether the diversity and composition of flightless ground beetles are elevationally stratified, and, if so, (2) what environmental factors (other than elevation per se) are associated with these patterns. Generalised linear models and model averaging techniques were used to relate patterns of diversity to environmental factors. Unlike most taxonomic groups, flightless ground beetles increased in species richness and abundance with elevation. Additionally, each subregion consisted of distinct assemblages containing a high level of regional endemic species. Species richness was most strongly positively associated with the historical climatic conditions and negatively associated with severity of recent disturbance (treefalls) and current climatic conditions. Assemblage composition was associated with latitude and current and historical climatic conditions. Our results suggest that distributional patterns of flightless ground beetles are not only likely to be associated with factors that change with elevation (current climatic conditions), but also factors that are independent of elevation (recent disturbance and historical climatic conditions). Variation in historical vegetation stability explained both species richness and assemblage composition patterns, probably reflecting the significance of upland refugia at a geographic time scale. These findings are important for conservation management as upland habitats are under threat from climate change.

Mesozooplankton abundance data from the fjord branch Kapisigdlit located in the Godthaabsfjord system, West Greenland, 2010

Sampling was conducted from March 24 to August 5 2010, in the fjord branch Kapisigdlit located in the inner part of the Godthåbsfjord system, West Greenland. The vessel "Lille Masik" was used during all cruises except on June 17-18 where sampling was done from RV Dana (National Institute for Aquatic Resources, Denmark). A total of 15 cruises (of 1-2 days duration) 7-10 days apart was carried out along a transect composed of 6 stations (St.), spanning the length of the 26 km long fjord branch. St. 1 was located at the mouth of the fjord branch and St. 6 was located at the end of the fjord branch, in the middle of a shallower inner creek . St. 1-4 was covering deeper parts of the fjord, and St. 5 was located on the slope leading up to the shallow inner creek. Mesozooplankton was sampled by vertical net tows using a Hydrobios Multinet (type Mini) equipped with a flow meter and 50 µm mesh nets or a WP-2 net 50 µm mesh size equipped with a non-filtering cod-end. Sampling was conducted at various times of day at the different stations. The nets were hauled with a speed of 0.2-0.3 m s**-1 from 100, 75 and 50 m depth to the surface at St. 2 + 4, 5 and 6, respectively. The content was immediately preserved in buffered formalin (4% final concentration). All samples were analyzed in the Plankton sorting and identification center in Szczecin (www.nmfri.gdynia.pl). Samples containing high numbers of zooplankton were split into subsamples. All copepods and other zooplankton were identified down to lowest possible taxonomic level (approx. 400 per sample), length measured and counted. Copepods were sorted into development stages (nauplii stage 1 - copepodite stage 6) using morphological features and sizes, and up to 10 individuals of each stage was length measured.

Benthic foraminifera abundance data of sediment core GeoB7926-2

We present a high resolution, multiproxy study of the relationship between pelagic and benthic environments of a coastal upwelling system in the subtropical NE Atlantic Ocean. Marine sediments corresponding to late MIS3 to the Holocene in the radiocarbon dated core GeoB7926, retrieved off Mauritania (21°N) were analysed to reconstruct productivity in surface waters and its linkage to deep waters during the last 35 ka BP. High latitude cold events and changes in atmospheric and oceanographic dynamics influenced upwelling intensity over this time period. Subsequently, this caused changes in primary productivity off this low-latitude coastal upwelling locality. The benthic foraminiferal fauna displays four main community shifts corresponding to fundamental climatic events, first of all during late MIS3 (35-28 ka BP), secondly from 28 to 19 ka BP (including Heinrich event 2 and the LGM), thirdly within Heinrich event 1, the Bølling Allerød and the Younger Dryas (18-11.5 ka BP) and finally during the Holocene (11.5-0 ka BP). In particular, strong pelagic-benthic coupling is apparent in MIS 3, as demonstrated by increased primary productivity, indicated by moderate DAR and the dominance of benthic foraminiferal species which prefer fresh phytodetritus. A decline in upwelling intensity and nutrient availability follows, which resulted in a proportionately larger amount of older, degraded matter, provoking a shift in the benthic foraminifera fauna composition. This rapid response of the benthic environment continues with a progressive increase in upwelling intensity due to sea level and oceanographic changes and according high surface production during the LGM. During Heinrich event 1 and the Younger Dryas, extreme levels of primary production actually hindered benthic environment through the development of low oxygen conditions. After this period, a final change in benthic foraminiferal community composition occurs which indicates a return to more oxygenated conditions during the Holocene.

Mesozooplankton abundance data from Disko Bay, West Greenland, 2008

The study site was located in the Disko Bay off Qeqertarsuaq, western Greenland. Due to land-connected sea ice coverage during winter, 2 sampling sites were combined. At the first site in winter (21 February to 23 March 2008), sampling was conducted through a hole in the ice at ca. 65 to 160 m depth approximately 0.5 nautical mile (n mile) south of Qeqertarsuaq (69° 14' N, 53° 29' W). In spring and summer (9 April to 18 July), sampling was done at a monitoring station 1 n mile south from Qeqertarsuaq (69° 14' N, 53° 23' W) at 300 m depth. Sampling was carried out between 10:00 and 17:00 h. During sampling from the ice, mesozooplankton was collected using a modified WP-2 net (45 µm) equipped with a closing mechanism (Hydrobios). Samples were collected in 3 depth strata (0-50, 50-100, and 100-150 m). During ship-based sampling, mesozooplankton was collected with a multinet (50 µm) equipped with a flow meter (Multinet, Hydrobios type midi), and 2 additional depth strata (150-200m and 200-250 m) were included. In addition to the seasonal study one diurnal investigation with sampling every 6 h was conducted from 29 April at 12:00 h to 30 April 30 at 12:00 h. Samples were immediately preserved in buffered formalin (5% final concentration) for later analyses. Biomass values of the different copepod species were calculated based on measurements of prosome length, and length/weight relationships. Two regressions for Calanus spp. were established for biomass calculations: one applicable prior to and during the phytoplankton bloom until 4 May, and another from 9 May onwards.

Presence-absence versus abundance data for monitoring threatened species

Effective detection of population trend is crucial for managing threatened species. Little theory exists, however, to assist managers in choosing the most cost-effective monitoring techniques for diagnosing trend. We present a framework for determining the optimal monitoring strategy by simulating a manager collecting data on a declining species, the Chestnut-rumped Hylacola (Hylacola pyrrhopygia parkeri), to determine whether the species should be listed under the IUCN (World Conservation Union) Red List. We compared the efficiencies of two strategies for detecting trend, abundance, and presence-absence surveys, underfinancial constraints. One might expect the abundance surveys to be superior under all circumstances because more information is collected at each site. Nevertheless, the presence-absence data can be collected at more sites because the surveyor is not obliged to spend a fixed amount of time at each site. The optimal strategy for monitoring was very dependent on the budget available. Under some circumstances, presence-absence surveys outperformed abundance surveys for diagnosing the IUCN Red List categories cost-effectively. Abundance surveys were best if the species was expected to be recorded more than 16 times/year; otherwise, presence-absence surveys were best. The relationship between the strategies we investigated is likely to be relevant for many comparisons of presence-absence or abundance data. Managers of any cryptic or low-density species who hope to maximize their success of estimating trend should find an application for our results.

Global distributions of picophytoplankton abundance and biomass - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

The smallest marine phytoplankton, collectively termed picophytoplankton, have been routinely enumerated by flow cytometry since the late 1980s, during cruises throughout most of the world ocean. We compiled a database of 40,946 data points, with separate abundance entries for Prochlorococcus, Synechococcus and picoeukaryotes. We use average conversion factors for each of the three groups to convert the abundance data to carbon biomass. After gridding with 1° spacing, the database covers 2.4% of the ocean surface area, with the best data coverage in the North Atlantic, the South Pacific and North Indian basins. The average picophytoplankton biomass is 12 ± 22 µg C L-1 or 1.9 g C m-2. We estimate a total global picophytoplankton biomass, excluding N2-fixers, of 0.53 - 0.74 Pg C (17 - 39 % Prochlorococcus, 12 - 15 % Synechococcus and 49 - 69 % picoeukaryotes). Future efforts in this area of research should focus on reporting calibrated cell size, and collecting data in undersampled regions.

Global distributions of diatoms abundance, biovolume and biomass - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

This study is a first effort to compile the largest possible body of data available from different plankton databases as well as from individual published or unpublished datasets regarding diatom distribution in the world ocean. The data obtained originate from time series studies as well as spatial studies. This effort is supported by the Marine Ecosystem Data (MAREDAT) project, which aims at building consistent data sets for the main PFTs (Plankton Functional Types) in order to help validate biogeochemical ocean models by using converted C biomass from abundance data. Diatom abundance data were obtained from various research programs with the associated geolocation and date of collection, as well as with a taxonomic information ranging from group down to species. Minimum, maximum and average cell size information were mined from the literature for each taxonomic entry, and all abundance data were subsequently converted to biovolume and C biomass using the same methodology.

Global distributions of epipelagic macrozooplankton abundance and biomass - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

Macrozooplankton are an important link between higher and lower trophic levels in the oceans. They serve as the primary food for fish, reptiles, birds and mammals in some regions, and play a role in the export of carbon from the surface to the intermediate and deep ocean. Little, however, is known of their global distribution and biomass. Here we compiled a dataset of macrozooplankton abundance and biomass observations for the global ocean from a collection of four datasets. We harmonise the data to common units, calculate additional carbon biomass where possible, and bin the dataset in a global 1 x 1 degree grid. This dataset is part of a wider effort to provide a global picture of carbon biomass data for key plankton functional types, in particular to support the development of marine ecosystem models. Over 387 700 abundance data and 1330 carbon biomass data have been collected from pre-existing datasets. A further 34 938 abundance data were converted to carbon biomass data using species-specific length frequencies or using species-specific abundance to carbon biomass data. Depth-integrated values are used to calculate known epipelagic macrozooplankton biomass concentrations and global biomass. Global macrozooplankton biomass has a mean of 8.4 µg C l-1, median of 0.15 µg C l-1 and a standard deviation of 63.46 µg C l-1. The global annual average estimate of epipelagic macrozooplankton, based on the median value, is 0.02 Pg C. Biomass is highest in the tropics, decreasing in the sub-tropics and increasing slightly towards the poles. There are, however, limitations on the dataset; abundance observations have good coverage except in the South Pacific mid latitudes, but biomass observation coverage is only good at high latitudes. Biomass is restricted to data that is originally given in carbon or to data that can be converted from abundance to carbon. Carbon conversions from abundance are restricted in the most part by the lack of information on the size of the organism and/or the absence of taxonomic information. Distribution patterns of global macrozooplankton biomass and statistical information about biomass concentrations may be used to validate biogeochemical models and Plankton Functional Type models.

Global distributions of Phaeocystis sp. abundance and biomass - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

The planktonic haptophyte Phaeocystis has been suggested to play a fundamental role in the global biogeochemical cycling of carbon and sulphur, but little is known about its global biomass distribution. We have collected global microscopy data of the genus Phaeocystis and converted abundance data to carbon biomass using species-specific carbon conversion factors. Microscopic counts of single-celled and colonial Phaeocystis were obtained both through the mining of online databases and by accepting direct submissions (both published and unpublished) from Phaeocystis specialists. We recorded abundance data from a total of 1595 depth-resolved stations sampled between 1955-2009. The quality-controlled dataset includes 5057 counts of individual Phaeocystis cells resolved to species level and information regarding life-stages from 3526 samples. 83% of stations were located in the Northern Hemisphere while 17% were located in the Southern Hemisphere. Most data were located in the latitude range of 50-70° N. While the seasonal distribution of Northern Hemisphere data was well-balanced, Southern Hemisphere data was biased towards summer months. Mean species- and form-specific cell diameters were determined from previously published studies. Cell diameters were used to calculate the cellular biovolume of Phaeocystis cells, assuming spherical geometry. Cell biomass was calculated using a carbon conversion factor for Prymnesiophytes (Menden-Deuer and Lessard, 2000). For colonies, the number of cells per colony was derived from the colony volume. Cell numbers were then converted to carbon concentrations. An estimation of colonial mucus carbon was included a posteriori, assuming a mean colony size for each species. Carbon content per cell ranged from 9 pg (single-celled Phaeocystis antarctica) to 29 pg (colonial Phaeocystis globosa). Non-zero Phaeocystis cell biomasses (without mucus carbon) range from 2.9 - 10?5 µg l-1 to 5.4 - 103 µg l-1, with a mean of 45.7 µg l-1 and a median of 3.0 µg l-1. Highest biomasses occur in the Southern Ocean below 70° S (up to 783.9 µg l-1), and in the North Atlantic around 50° N (up to 5.4 - 103 µg l-1).

Global distributions of diazotrophs abundance, biomass and nitrogen fixation rates - Gridded data product (NetCDF) - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

The MAREDAT atlas covers 11 types of plankton, ranging in size from bacteria to jellyfish. Together, these plankton groups determine the health and productivity of the global ocean and play a vital role in the global carbon cycle. Working within a uniform and consistent spatial and depth grid (map) of the global ocean, the researchers compiled thousands and tens of thousands of data points to identify regions of plankton abundance and scarcity as well as areas of data abundance and scarcity. At many of the grid points, the MAREDAT team accomplished the difficult conversion from abundance (numbers of organisms) to biomass (carbon mass of organisms). The MAREDAT atlas provides an unprecedented global data set for ecological and biochemical analysis and modeling as well as a clear mandate for compiling additional existing data and for focusing future data gathering efforts on key groups in key areas of the ocean. This is a gridded data product about diazotrophic organisms . There are 6 variables. Each variable is gridded on a dimension of 360 (longitude) * 180 (latitude) * 33 (depth) * 12 (month). The first group of 3 variables are: (1) number of biomass observations, (2) biomass, and (3) special nifH-gene-based biomass. The second group of 3 variables is same as the first group except that it only grids non-zero data. We have constructed a database on diazotrophic organisms in the global pelagic upper ocean by compiling more than 11,000 direct field measurements including 3 sub-databases: (1) nitrogen fixation rates, (2) cyanobacterial diazotroph abundances from cell counts and (3) cyanobacterial diazotroph abundances from qPCR assays targeting nifH genes. Biomass conversion factors are estimated based on cell sizes to convert abundance data to diazotrophic biomass. Data are assigned to 3 groups including Trichodesmium, unicellular diazotrophic cyanobacteria (group A, B and C when applicable) and heterocystous cyanobacteria (Richelia and Calothrix). Total nitrogen fixation rates and diazotrophic biomass are calculated by summing the values from all the groups. Some of nitrogen fixation rates are whole seawater measurements and are used as total nitrogen fixation rates. Both volumetric and depth-integrated values were reported. Depth-integrated values are also calculated for those vertical profiles with values at 3 or more depths.

Bias-corrected Pearson estimating functions for Taylor`s power law applied to benthic macrofauna data

Estimation of Taylor`s power law for species abundance data may be performed by linear regression of the log empirical variances on the log means, but this method suffers from a problem of bias for sparse data. We show that the bias may be reduced by using a bias-corrected Pearson estimating function. Furthermore, we investigate a more general regression model allowing for site-specific covariates. This method may be efficiently implemented using a Newton scoring algorithm, with standard errors calculated from the inverse Godambe information matrix. The method is applied to a set of biomass data for benthic macrofauna from two Danish estuaries. (C) 2011 Elsevier B.V. All rights reserved.

Correspondence analysis of raw data

Correspondence analysis has found extensive use in ecology, archeology, linguisticsand the social sciences as a method for visualizing the patterns of association in a table offrequencies or nonnegative ratio-scale data. Inherent to the method is the expression of the datain each row or each column relative to their respective totals, and it is these sets of relativevalues (called profiles) that are visualized. This relativization of the data makes perfect sensewhen the margins of the table represent samples from sub-populations of inherently differentsizes. But in some ecological applications sampling is performed on equal areas or equalvolumes so that the absolute levels of the observed occurrences may be of relevance, in whichcase relativization may not be required. In this paper we define the correspondence analysis ofthe raw unrelativized data and discuss its properties, comparing this new method to regularcorrespondence analysis and to a related variant of non-symmetric correspondence analysis.

Do phytoplankton communities evolve through a self-regulatory abundance-diversity relationship?

A small group of phytoplankton species that produce toxic or allelopathic chemicals has a significant effect on plankton dynamics in marine ecosystems. The species of non-toxic phytoplankton, which are large in number, are affected by the toxin-allelopathy of those species. By analysis of the abundance data of marine phytoplankton collected from the North-West coast of the Bay of Bengal, an empirical relationship between the abundance of the potential toxin-producing species and the species diversity of the non-toxic phytoplankton is formulated. A change-point analysis demonstrates that the diversity of non-toxic phytoplankton increases with the increase of toxic species up to a certain level. However, for a massive increase of the toxin-producing species the diversity of phytoplankton at species level reduces gradually. Following the results, a deterministic relationship between the abundance of toxic phytoplankton and the diversity of non-toxic phytoplankton is developed. The abundance–diversity relationship develops a unimodal pathway through which the abundance of toxic species regulates the diversity of phytoplankton. These results contribute to the current understanding of the coexistence and biodiversity of phytoplankton, the top-down vs. bottom-up debate, and to that of abundance–diversity relationship in marine ecosystems.