Phenology of mixed woody-herbaceous ecosystems following extreme events: net and differential responses

Ecosystem responses to key climate drivers are reflected in phenological dynamics such as the timing and degree of "green-up" that integrate responses over spatial scales from individual plants to ecosystems. This integration is clearest in ecosystems dominated by a single species or life form, such as seasonally dynamic grasslands or more temporally constant evergreen forests. Yet many ecosystems have substantial contribution of cover from both herbaceous and woody evergreen plants. Responses of mixed woody-herbaceous ecosystems to climate are of increasing concern due to their extensive nature, the potential for such systems to yield more complex responses than those dominated by a single life form, and projections that extreme climate and weather events will increase in frequency and intensity with global warming. We present responses of a mixed woody-herbaceous ecosystem type to an extreme event: regional-scale pi?on pine mortality following an extended drought and the subsequent herbaceous green-up following the first wet period after the drought. This example highlights how reductions in greenness of the slower, more stable evergreen woody component can rapidly be offset by increases associated with resources made available to the relatively more responsive herbaceous component. We hypothesize that such two-phase phenological responses to extreme events are characteristic of many mixed woody-herbaceous ecosystems.     

Four-decade responses of soil trace elements to an aggrading old-field forest: B, Mn, Zn, Cu, and Fe

In the ancient and acidic Ultisol soils of the Southern Piedmont, USA, we studied changes in trace element biogeochemistry over four decades, a period during which formerly cultivated cotton fields were planted with pine seedlings that grew into mature forest stands. In 16 permanent plots, we estimated 40-year accumulations of trace elements in forest biomass and O horizons (between 1957 and 1997), and changes in bioavailable soil fractions indexed by extractions of 0.05 mol/L HCl and 0.2 mol/L acid ammonium oxalate (AAO). Element accumulations in 40-year tree biomass plus O horizons totaled 0.9, 2.9, 4.8, 49.6, and 501.3 kg/ha for Cu, B, Zn, Mn, and Fe, respectively. In response to this forest development, samples of the upper 0.6-m of mineral soil archived in 1962 and 1997 followed one of three patterns. (1) Extractable B and Mn were significantly depleted, by -4.1 and -57.7 kg/ha with AAO, depletions comparable to accumulations in biomass plus O horizons, 2.9 and 49.6 kg/ha, respectively. Tree uptake of B and Mn from mineral soil greatly outpaced resupplies from atmospheric deposition, mineral weathering, and deep-root uptake. (2) Extractable Zn and Cu changed little during forest growth, indicating that nutrient resupplies kept pace with accumulations by the aggrading forest. (3) Oxalate-extractable Fe increased substantially during forest growth, by 275.8 kg/ha, about 10-fold more than accumulations in tree biomass (28.7 kg/ha). The large increases in AAO-extractable Fe in surficial 0.35-m mineral soils were accompanied by substantial accretions of Fe in the forest's O horizon, by 473 kg/ha, amounts that dwarfed inputs via litterfall and canopy throughfall, indicating that forest Fe cycling is qualitatively different from that of other macro- and micronutrients. Bioturbation of surficial forest soil layers cannot account for these fractions and transformations of Fe, and we hypothesize that the secondary forest's large inputs of organic additions over four decades has fundamentally altered soil Fe oxides, potentially altering the bioavailability and retention of macro- and micronutrients, contaminants, and organic matter itself. The wide range of responses among the ecosystem's trace elements illustrates the great dynamics of the soil system over time scales of decades.     

UVR-induced photoinhibition of summer marine phytoplankton communities from Patagonia

During austral summer 2006, experiments were carried out to evaluate the effects of ultraviolet radiation (UVR, 280–400 nm) on carbon fixation of natural phytoplankton assemblages from Patagonia (Argentina). Surface water samples were collected (ca. 100 m offshore) at mid morning using an acid-cleaned (1 N HCl) dark container. The short-term impact of UVR (measured as radiocarbon incorporation) was immediately assessed by exposing samples to three artificial illumination treatments: PAR (400–700 nm), PAR + UV-A (320–700 nm), and PAR + UV-A + UV-B (280–700 nm). Pico-nanoplankton characterized the assemblages, and taxon-specific pigment fingerprinting combined with CHEMTAX and supplemented with microscopic observations showed varied proportions of diatoms, chlorophytes, and cyanobacteria throughout January–February 2006. Photosynthetic efficiency, as assessed through assimilation numbers, was high [between 4.4 and 10.4 μg C (μg chl-a)⁻¹ h⁻¹], and it was probably favored by the supply of inorganic nutrients from the Chubut River. UVR-induced photoinhibition appeared to be related to the taxonomic composition: in general, higher photoinhibition was observed when diatoms dominated, whereas this was lower when samples were dominated by chlorophytes. Our data suggest that xanthophyll pigments might have provided only limited protection in these already highlighted acclimated assemblages.     

Simplicity and complexity in ecological data analysis

I argue that ecological data analyses are often needlessly complicated, and I present two examples of published analyses for which simpler alternatives are available. Unnecessary complexity is often introduced when analysts focus on subunits of the key experimental or observational units in a study, or use a very general framework to present an analysis that is a simple special case. Simpler analyses are easier to explain and understand; they clarify what the key units in a study are; they reduce the chances for computational mistakes; and they are more likely to lead to the same conclusions when applied by different analysts to the same data.     

Adjusting age and stage distributions for misclassification errors

Ecologists often use samples from the age or stage structure of a population to make inferences about population-level processes and to parameterize matrix models. Typically, researchers make a simplifying assumption that age and stage classes are determined without error, when in fact some level of misclassification often can be expected. If unaccounted for, misclassification will lead to overly optimistic levels of precision and can cause biased estimates of age or stage structure. Although several studies have used information from known-age individuals to quantify errors in age or stage distribution, the problem of estimating the age or stage structure in face of such errors has received comparably little attention. In this paper, we describe a general statistical framework for estimating the true stage distribution of a sample when misclassification rates can be estimated. The estimation process requires auxiliary information on misclassification rates, such as data from individuals of known age. We analyze age-structured harvest records from black bears in Pennsylvania to illustrate how incorporating misclassification errors leads to changes in point estimates and provides a measure of precision.     

Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling

Killer whale (Orcinus orca Linnaeus, 1758) abundance in the North Pacific is known only for a few populations for which extensive longitudinal data are available, with little quantitative data from more remote regions. Line-transect ship surveys were conducted in July and August of 2001–2003 in coastal waters of the western Gulf of Alaska and the Aleutian Islands. Conventional and Multiple Covariate Distance Sampling methods were used to estimate the abundance of different killer whale ecotypes, which were distinguished based upon morphological and genetic data. Abundance was calculated separately for two data sets that differed in the method by which killer whale group size data were obtained. Initial group size (IGS) data corresponded to estimates of group size at the time of first sighting, and post-encounter group size (PEGS) corresponded to estimates made after closely approaching sighted groups. ‘Resident’-type (fish-eating) killer whales were more abundant than the ‘transient’-type (mammal-eating). Abundance estimates of resident killer whales (991 [95% CI = 379–2,585] [IGS] and 1,587 [95% CI = 608–4,140] [PEGS]), were at least four times greater than those of the transient killer whales (200 [95% CI = 81–488] [IGS] and 251 [95% CI = 97–644] whales [PEGS]). The IGS estimate of abundance is preferred for resident killer whales because the estimate based on PEGS data may show an upward bias. The PEGS estimate of abundance is likely more accurate for transients. Residents were most abundant near Kodiak Island in the northern Gulf of Alaska, around Umnak and Unalaska Islands in the eastern Aleutians, and in Seguam Pass in the central Aleutians. This ecotype was not observed between 156 and 164°W, south of the Alaska Peninsula. In contrast, transient killer whale sightings were found at higher densities south of the Alaska Peninsula between the Shumagin Islands and the eastern Aleutians. Only two sightings of ‘offshore’-type killer whales were recorded during the surveys, one northeast of Unalaska Island and the other south of Kodiak Island. These are the first estimates of abundance of killer whale ecotypes in the Aleutian Islands and Alaska Peninsula area and provide a baseline for quantifying the role of these top predators in their ecosystem. Electronic Supplementary Material  Supplementary material is available in the online version of this article at and is accessible for authorized users.

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Spatial patterns of tungsten and cobalt in surface dust of Fallon,Nevada

Spatial patterns of tungsten and cobalt are described for surface dust of Fallon, Nevada, where a cluster of childhood leukemia has been ongoing since 1997. In earlier research, airborne tungsten and cobalt was shown to be elevated in total suspended particulates in Fallon. To fine-tune the spatial patterns of tungsten and cobalt deposition in Fallon, surface dust was collected in a grid pattern within as well as outside of Fallon to establish background concentrations of metals. In surface dust, tungsten and cobalt show sharp peaks (934 ppm and 98 ppm, respectively) within Fallon just north of highway 50 and west of highway 95. These two peaks overlap spatially, and given the grid pattern used for collecting surface dust, the source area of these two airborne metals can be pinpointed to the vicinity of hard-metal industry located north of highway 50 and west of highway 95. Fallon is distinctive in west central Nevada because of high airborne tungsten and cobalt particulates, and given its cluster of childhood leukemia, it stands to reason that additional biomedical research is in order to test directly the leukogenicity of combined airborne tungsten and cobalt particulates.