Cape Bird Research Articles

Ornithogenic sedimentary profiles of n-alkanes and PAHs constrain breeding penguin population dynamics at Cape Bird, Ross Island, Antarctica, over the past 1,500 years

Ornithogenic sediment may contain important paleoecological information concerning past bird population dynamics. In this paper, we analyse the distribution of n-alkanes and polycyclic aromatic hydrocarbons (PAHs) in two sedimentary profiles collected from abandoned penguin colonies at Cape Bird, Ross Island, Antarctica. The geochronology of the sediment profiles was determined using 210Pb and AMS 14C dating techniques, and spans the past 1500 years. We observe low levels of n-alkanes (0.93–2.67 μg g−1) and PAHs (20.5–46.8 ng g−1) concentrations dominated by short-chain n-alkanes, low-molecular-weight and alkyl PAHs. Principal Component Analysis (PCA) shows that the PC 1 scores for n-alkanes and PAHs are significantly correlated with the input intensity of penguin guano and penguin bioelements. Using these proxies in combination with Generalized Additive Model (GAM), we reveal that breeding penguin population growth reaches a historical peak during the Medieval Climate Anomaly (MCA) prior to 1550 CE. This growth in population size corresponds to the extent of sea ice cover and the activity of the local atmospheric-ocean circulation. Warming induced by El Niño and the positive phase of the Southern Annular Mode, along with the expansion of ice-free zones and increased oceanic nutrient and food availability, are considered primary factors contributing to the growth of breeding penguin populations. Additionally, variations in sea ice extent and Amundsen Sea Low significantly influence penguin population dynamics. Our study suggests that n-alkanes and PAHs may be valuable organic proxies for reconstructing historical changes in breeding penguin populations, with the local sea ice extent and atmospheric-oceanic circulation exerting a major influence on Antarctic penguin population dynamics.

Historical population changes of Adélie penguins in the Ross Sea region, Antarctica, and its climatic forcings

The atmospheric-oceanic circulation patterns, especially for the Southern Annular Mode (SAM) and El Niño-Southern Oscillation (ENSO), two major atmospheric circulation patterns in the Ross Sea region, have been reported to greatly affect climate and marine ecosystems. However, from a historical perspective, the influence of atmospheric-oceanic circulation patterns on penguin populations remains unclear in this region. Here, we analyzed two lacustrine sediment cores collected from abandoned penguin colonies at Inexpressible Island, Ross Sea, Antarctica, and by applying alternative geochemical indices, successfully reconstructed the populations of the Adélie penguins (Pygoscelis adeliae) over the past ∼1500 years. We found that penguin population peaked during 750–1350 AD at Inexpressible Island, potentially due to habitat expansion in the warmer climate. After comparing with historical records of penguin populations at Cape Bird, Dunlop Island, and Cape Adare, all were found to have a common increase during the 750-1350 AD period in the Ross Sea. The population trend also coincided with extreme activities of El Niño and SAM (+). We inferred that the SAM-ENSO might promote influxes of Circumpolar and Modified Circumpolar Deep Waters into the Ross Sea. The enhanced influx of nutrient-rich deep water, together with a warmer climate may jointly enhance polynya efficiency and population increase of Adélie penguins. Our study indicates the potentially significant roles of ENSO and SAM in the regulation of Antarctic ecosystems.

Diet segregation in Adélie penguins: some individuals attempt to overcome colony-induced and annual foraging challenges

Intraspecific competition for food can be especially high in colonial breeding seabirds. To minimize colony-induced or annual foraging challenges, diet may vary among individuals, but few studies have simultaneously investigated the effects of both extrinsic conditions (e.g. colony or year effects) and parameters of an individual (e.g. sex, age or individual quality) on diet in seabirds. Using stable isotope analyses, we studied the diet of 214 Adélie penguins Pygoscelis adeliae of known sex, age and breeding quality, nesting in 2 colonies on Ross Island, Antarctica, over 3 breeding seasons. During the study, δ15N and δ13C isotope values were lower in penguins breeding at Cape Crozier compared to those at Cape Bird, revealing a difference in prey proportions. Cape Bird penguins were estimated to consistently consume more energy-rich silverfish Pleuragramma antarctica, while birds at Cape Crozier ate more crystal krill Euphausia crystallorophias. We also found inter-annual differences in diet, with a higher dietary fish proportion in both colonies during 2011. Males had significantly higher δ15N values, indicating a higher fish consumption than females. This sexual segregation in diet was particularly pronounced at Cape Bird, where the overall isotopic niche was wider than at Cape Crozier. Differences in diet among adults of varying ages only existed at Cape Bird, where middle-aged penguins consumed more fish than old and young penguins. This study provides evidence that Adélie penguin diet is largely driven by annual, seasonal and local abundances of prey, with only some individuals selectively foraging for more nutritional prey if prey choices are present.

Inter-individual differences in the foraging behavior of breeding Adélie penguins are driven by individual quality and sex

Inter-individual differences in demographic traits of iteroparous species can arise through learning and maturation, as well as from permanent differences in individual ‘quality’ and sex-specific constraints. As the ability to acquire energy determines the resources an individual can allocate to reproduction and self-maintenance, foraging behavior is a key trait to study to better understand the mechanisms underlying these differences. So far, most seabird studies have focused on the effect of maturation and learning processes on foraging performance, while only a few have included measures of individual quality. Here, we investigated the effects of age, breeding experience, sex, and individual breeding quality on the foraging behavior and location of 83 known-age Adélie penguins at Cape Bird, Ross Sea, Antarctica. Over a 2 yr period, we showed that (1) high-quality birds dived deeper than lower quality ones, apparently catching a higher number of prey per dive and targeting different foraging locations; (2) females performed longer foraging trips and a higher number of dives compared to males; (3) there were no significant age-related differences in foraging behavior; and (4) breeding experience had a weak influence on foraging behavior. We suggest that high-quality individuals have higher physiological ability, enabling them to dive deeper and forage more effectively. Further inquiry should focus on determining the physiological differences among penguins of different quality.

Oceanographic mechanisms and penguin population increases during the Little Ice Age in the southern Ross Sea, Antarctica

The Adélie penguin is a well-known indicator for climate and environmental changes. Exploring how large-scale climate variability affects penguin ecology in the past is essential for understanding the responses of Southern Ocean ecosystems to future global change. Using ornithogenic sediments at Cape Bird, Ross Island, Antarctica, we inferred relative population changes of Adélie penguins in the southern Ross Sea over the past 500 yr, and observed an increase in penguin populations during the Little Ice Age (LIA; 1500–1850 AD). We used cadmium content in ancient penguin guano as a proxy of ocean upwelling and identified a close linkage between penguin dynamics and atmospheric circulation and oceanic conditions. During the cold period of ∼1600–1825 AD, a deepened Amundsen Sea Low (ASL) led to stronger winds, intensified ocean upwelling, enlarged Ross Sea and McMurdo Sound polynyas, and thus higher food abundance and penguin populations. We propose a mechanism linking Antarctic marine ecology and atmospheric/oceanic dynamics which can help explain and predict responses of Antarctic high latitudes ecosystems to climate change.

South Polar Skua breeding populations in the Ross Sea assessed from demonstrated relationship with Adélie Penguin numbers

In the Ross Sea region, most South Polar Skuas (Stercorarius maccormicki) nest near Adelie Penguin (Pygoscelis adeliae) colonies, preying and scavenging on fish, penguins, and other carrion. To derive a relationship to predict skua numbers from better-quantified penguin numbers, we used distance sampling to estimate breeding skua numbers within 1000 m of 5 penguin nesting locations (Cape Crozier, Cape Royds, and 3 Cape Bird locations) on Ross Island in 3 consecutive years. Estimated numbers of skua breeding pairs were highest at Cape Crozier (270,000 penguin pairs; 1099 and 1347 skua pairs in 2 respective years) and lowest at Cape Royds (3000 penguin pairs; 45 skua pairs). The log–log linear relationship (R 2 = 0.98) between pairs of skuas and penguins was highly significant, and most historical estimates of skua and penguin numbers in the Ross Sea were within 95 % prediction intervals of the regression. Applying our regression model to current Adelie Penguin colony sizes at 23 western Ross Sea locations predicted that 4635 pairs of skuas now breed within 1000 m of penguin colonies in the Ross Island metapopulation (including Beaufort Island) and northern Victoria Land. We estimate, using published skua estimates for elsewhere in Antarctica, that the Ross Sea South Polar Skua population comprises ~50 % of the world total, although this may be an overestimate because of incomplete data elsewhere. To improve predictions and enable measurement of future skua population change, we recommend additional South Polar Skua surveys using consistent distance-sampling methods at penguin colonies of a range of sizes.

From warm to cold: migration of Adélie penguins within Cape Bird, Ross Island

Due to their sensitivity to environmental change, penguins in Antarctica are widely used as bio-indicators in paleoclimatic research. On the basis of bio-element assemblages identified in four ornithogenic sediment profiles, we reconstructed the historical penguin population change at Cape Bird, Ross Island, for the past 1600 years. Clear succession of penguin population peaks were observed in different profiles at about 1400 AD, which suggested a high probability of migration within this region. The succession was most obviously marked by a sand layer lasting from 1400 to 1900 AD in one of the analyzed profiles. Multiple physical/chemical parameters indicated this sand layer was not formed in a lacustrine environment, but was marine-derived. Both isostatic subsidence and frequent storms under the colder climatic condition of the Little Ice Age were presumed to have caused the abandonment of the colonies, and we believe the penguins migrated from the coastal area of mid Cape Bird northward and to higher ground as recorded in the other sediment profiles. This migration was an ecological response to global climate change and possible subsequent geological effects in Antarctica.

Adélie penguin dietary remains reveal Holocene environmental changes in the western Ross Sea (Antarctica)

Carbon and nitrogen stable isotope analyses were performed on modern and Holocene Adélie penguin guano samples collected from ornithogenic soils along the Scott Coast (Southern Victoria Land, Antarctica), from Cape Irizar to Dunlop Island, and at Cape Bird (Ross Island). Guano samples also were sieved and sorted under stereomicroscope in order to select penguin dietary remains, such as fish bones and otoliths. Carbon and nitrogen stable isotope composition, coupled with the taxonomic identification of fish otoliths from Scott Coast Holocene samples, indicated a mainly fish-based diet for this area, with Pleuragramma antarcticum as the most eaten prey throughout the investigated period (from 390cal BP to ca 7300cal BP). The isotopic values of Ross Island samples (from modern to 3850cal BP) showed a krill consumption increase in the samples younger than 2000cal BP, with the maximum in modern samples. Scott Coast and Ross Island Holocene samples showed δ13C and δ15N trends similar to those previously published from Terra Nova Bay (northern Victoria Land), whereas modern samples from Ross Island have similar δ15N composition but different δ13C values. This δ13C divergence started at ca 2000 BP and follows the abandonment of the Scott Coast colonies.The δ13C trend observed in Ross Island and Terra Nova Bay samples and the abandonment of the Scott Coast colonies could suggest the stability and the persistence of the previous oceanographic conditions (i.e. polynya) for the Terra Nova Bay area and the establishment of new conditions for water circulation in the Southern Ross Sea since ~2000 BP when persistent sea-ice sealed the Scott Coast.

Increase in penguin populations during the Little Ice Age in the Ross Sea, Antarctica

Penguins are an important seabird species in Antarctica and are sensitive to climate and environmental changes. Previous studies indicated that penguin populations increased when the climate became warmer and decreased when it became colder in the maritime Antarctic. Here we determined organic markers in a sediment profile collected at Cape Bird, Ross Island, high Antarctic, and reconstructed the history of Adélie penguin colonies at this location over the past 700 years. The region transformed from a seal to a penguin habitat when the Little Ice Age (LIA; 1500–1800 AD) began. Penguins then became the dominant species. Penguin populations were the highest during ca. 1490 to 1670 AD, a cold period, which is contrary to previous results in other regions much farther north. Different responses to climate change may occur at low latitudes and high latitudes in the Antarctic, even if for same species.

Phil Hockey (1956-2013)

Phil Hockey (1956-2013)

Environmental influences on Adelie penguin breeding schedules, endocrinology, and chick survival

To understand how the social and physical environment influences behaviour, reproduction and survival, studies of underlying hormonal processes are crucial; in particular, interactions between stress and reproductive responses may have critical influences on breeding schedules. Several authors have examined the timing of breeding in relation to environmental stimuli, while others have independently described endocrine profiles. However, few studies have simultaneously measured endocrine profiles, breeding behaviour, and offspring survival across seasons. We measured sex and stress hormone concentrations (oestrogens, testosterone, and corticosterone), timing of breeding, and chick survival, in Adelie penguins ( Pygoscelis adeliae) at two colonies in two different years. Clutch initiation at Cape Bird South (CBS; year 1, ∼14,000 pairs) occurred later than at Cape Crozier East (CCE; year 2, ∼25,000 pairs); however, breeding was more synchronous at CBS. This pattern was probably generated by the persistence of extensive sea ice at CBS (year 1). Higher corticosterone metabolite and lower sex hormone concentrations at CBS correlated with later breeding and lower chick survival compared to at CCE – again, a likely consequence of sea ice conditions. Within colonies, sub-colony size (S, 50–100; M, 200–300; L, 500–600; XL, >1000 pairs) did not influence the onset or synchrony of breeding, chick survival, or hormone concentrations. We showed that the endocrine profiles of breeding Adelie penguins can differ markedly between years and/or colonies, and that combining measures of endocrinology, behaviour, and offspring survival can reveal the mechanisms and consequences that different environmental conditions can have on breeding ecology.

Extended ecophysiological analysis of Gomphiocephalus hodgsoni (Collembola): flexibility in life history strategy and population response

The springtail Gomphiocephalus hodgsoni (Arthropoda: Collembola) has been the focus of extensive ecophysiological and molecular genetic work and is now arguably the most well-studied of the continental Antarctic springtails. Here, we further the ecophysiological catalogue of this species. First, we provide experimental data on G. hodgsoni from one summer season at Cape Bird (Ross Sea Region) examining dispersal ability and desiccation tolerance. Next, we expand an existing metabolic rate dataset that encompasses individual metabolic rate measurements across both temporal and spatial scales in southern Victoria Land, adding an additional season of metabolic rate measurements taken at a cooler, drier continental location (Garwood Valley). Our data show that some G. hodgsoni individuals can survive at least ten days of suspension on the surface of both fresh and sea water. This, coupled with the presence of G. hodgsoni specimens in air and pitfall traps suggests that dispersal over local scales (i.e. metres) is possible for this species. Our metabolic data show that different populations within the same Antarctic region have different average metabolic rates at both temporal and spatial scales, indicating that distinct populations may respond differently to environmental variables. We suggest that G. hodgsoni maintains a flexible life history strategy that allows its ecophysiological response(s) to be dependent on local environmental conditions. Accordingly, there may be no ‘typical’ response to environmental changes—a factor that should be considered in both future ecophysiological work and conservation approaches.

Temporal and spatial metabolic rate variation in the Antarctic springtail Gomphiocephalus hodgsoni

Spatial and temporal environmental variation in terrestrial Antarctic ecosystems are known to impact species strongly at a local scale, but the ways in which organisms respond (e.g. physiologically, behaviourally) to such variation are poorly understood. Further, very few studies have attempted to assess inter-annual variability of such responses.Building on previous work demonstrating intra-seasonal variation in standard metabolic rate in the springtail Gomphiocephalushodgsoni, we investigated variation in metabolic activity of G. hodgsoni across two austral summer periods at Cape Bird, Ross Island. We also examined the influence of spatial variation by comparing metabolic rates of G. hodgsoni at Cape Bird with those from two other isolated continental locations within Victoria Land (Garwood and Taylor Valleys).We found significant differences between metabolic rates across the 2 years of measurement at Cape Bird. In addition, standard metabolic rates of G. hodgsoni obtained from Garwood and Taylor Valleys were significantly higher than those at Cape Bird where habitats are comparable, but environmental characteristics differ (e.g. microclimatic temperatures are higher).We discuss potential underlying causes of these metabolic rate variation patterns, including those related to differences among individuals (e.g. physiological and genetic differences), locations (e.g. habitat quality and microclimatic regime differences) and populations (e.g. acclimation differences among G. hodgsoni populations in the form of metabolic cold adaptation (MCA)).

Corticosterone responses in birds: Individual variation and repeatability in Adelie penguins ( Pygoscelisadeliae) and other species, and the use of power analysis to determine sample sizes

Plasma corticosterone concentrations increase when birds experience a stressor, and in this study we quantified variation in corticosterone responses for the first time in a species of free-living bird. Adelie penguins ( Pygoscelis adeliae) nesting at Cape Bird on Ross Island in Antarctica were sampled on three occasions. Penguins with relatively low or high corticosterone responses on the first occasion had consistently low or high responses, as previously found for great tits and chickens. A model for birds is proposed in which birds with low corticosterone responses and proactive personalities are likely to be more successful (have greater fitness) in constant or predictable conditions, whilst birds with reactive personalities and high corticosterone responses will be more successful in changing or unpredictable conditions. There is thus no linear relationship between the size of a corticosterone response and fitness. Whilst the absolute magnitude of corticosterone responses varies markedly across species of birds, coefficients of variation are similar. Individual corticosterone responses are generally repeatable, with significant statistical repeatabilities for 30 min corticosterone concentrations and integrated corticosterone concentrations in the Adelie penguin, great tit and chicken. Coefficients of variation in corticosterone responses between birds and power analyses were used to provide a rule of thumb for determining differences between groups of birds in mean corticosterone concentrations to enable statistical analyses to have acceptable levels of statistical power for given sample sizes. It is suggested that power analyses and this rule of thumb be adopted in future investigations of corticosterone responses in birds.

Bacterial diversity associated with ornithogenic soil of the Ross Sea region, AntarcticaThis article is one of a selection of papers in the Special Issue on Polar and Alpine Microbiology.

In the Ross Sea region of Antarctica, ornithogenic soils form on land under Adélie Penguin rookeries. Compared with mineral soils of the Ross Sea region, ornithogenic soils are generally high in microbial biomass, organic carbon, and total nitrogen and phosphorus, with high electrical conductivity and large variations in pH. The objective of this study was to assess the bacterial composition of ornithogenic soils from Cape Hallett and Cape Bird in the Ross Sea region using culture-independent methods. Soil clone libraries were constructed and those clones that occurred > or = 3 times were sequenced. The bacterial diversity of the soils was dependent on the presence of penguins. Firmicutes most closely related to the endospore-formers (e.g., Oceanobacillus profundus and Clostridium acidurici) and (or) Gammaproteobacteria belonging to the genus Psychrobacter dominated soils currently occupied with penguins. In contrast, Gammaproteobacteria, closely related to cultured members of the genera Rhodanobacter, Psychrobacter, Dokdonella, and Lysobacter, dominated the soils previously colonized by penguins. Results of this study indicate that despite relatively high nutrient levels and microbial biomass, bacterial communities of ornithogenic soils were not more diverse than those of mineral soils of the Ross Sea region of Antarctica.

Corticosterone Responses to Capture and Restraint in Emperor and Adelie Penguins in Antarctica

Birds respond to capture, handling and restraint with increased secretion of corticosterone, a glucocorticoid hormone that helps birds adjust to stressful situations. Hoods are reported to calm birds, but possible effects of hoods on corticosterone responses have not been reported for any bird. Corticosterone responses to restraint in Adelie penguins held by their legs with their head covered by a hood were markedly lower than responses of penguins restrained in a mesh bag inside a cardboard box (corticosterone at 30 min 15.69+/-1.72 cf. 28.32+/-2.75 ng/ml). The birds restrained by the two methods were sampled at the same location but in different years, so the differences in corticosterone responses cannot unequivocally be ascribed to an effect of hoods to reduce corticosterone responses. Corticosterone responses have been measured in some penguins, but not in the largest, the emperor penguin (Aptenodytes forsteri). The relationship between body mass and corticosterone responses to capture and restraint in penguins was examined in emperor penguins captured on sea ice in McMurdo Sound and Adelie penguins (Pygoscelis adeliae) captured at Cape Bird, Ross Island, Antarctica. Total integrated corticosterone responses were higher in the emperor than the Adelie penguins, but corrected integrated corticosterone responses, which represent the increase in corticosterone from initial concentrations and hence the corticosterone response to restraint, were the same. The results for the emperor and Adelie penguins, together with data from other penguin species, suggest that there is no relationship between the size of corticosterone responses and body mass in penguins.

Corticosterone in relation to body mass in Adelie penguins ( Pygoscelis adeliae) affected by unusual sea ice conditions at Ross Island, Antarctica

Penguins naturally fast each year during breeding and again whilst moulting, and may lose more than 40% of body mass during a fast. Fasting in emperor ( Aptenodytes forsteri) and king ( Aptenodytes patagonicus) penguins has been divided into three phases, with phase III characterised by an increased rate of body mass loss, increased plasma corticosterone concentrations, and a change in behaviour leading to abandonment of the breeding attempt and return to sea to feed. Initial corticosterone concentrations and corticosterone responses to a handling stressor were measured in the current study to determine if they increase during phase III of fasting in Adelie penguins ( Pygoscelis adeliae). The study was conducted in 2001 at the northern Cape Bird colony on Ross Island, Antarctica. Penguin breeding on Ross Island was disrupted in the 2001–2002 summer by a large iceberg (B15A) which stopped the normal movement of sea ice in the Ross Sea. Penguins departing from the Cape Bird colony were lighter than returning or incubating birds (3.39 ± 0.10 cf. 4.16 ± 0.06 and 4.07 ± 0.08 kg). It is likely that the departing birds were males that had been lighter than normal when they arrived at the colony. Initial plasma corticosterone concentrations were higher in departing than returning or incubating penguins (6.89 ± 1.69 cf. 2.36 ± 0.42 and 1.08 ± 0.19 ng/ml). Corticosterone responses to handling were also greater in departing penguins. Initial plasma corticosterone, concentrations at 30 min and total and corrected integrated corticosterone responses were inversely related to body mass in departing penguins, whereas there were no relationships in arriving penguins. β-hydroxybutyrate and uric acid concentrations were consistent with departing birds having entered phase III of fasting. The results indicate that corticosterone and corticosterone responses are elevated in phase III of fasting in the Adelie penguin.

Richard Liversidge, 1926–2003

Richard Liversidge, 1926–2003

Long‐term isolation and recent range expansion from glacial refugia revealed for the endemic springtail Gomphiocephalus hodgsoni from Victoria Land, Antarctica

We examined the phylogeography of the endemic Antarctic collembolan Gomphiocephalus hodgsoni using allozymes and mitochondrial DNA (mtDNA; COI) to determine if potentially limited dispersal and long-term habitat fragmentation have promoted regional genetic differentiation. Allozyme analyses showed that differentiation among 21 populations within the Ross Dependency was high (FST = 0.55) with two main groups each representing a distinct geographical region: (1) Ross Island and Beaufort Island; and (2) all continental sites. Ross Island populations showed low levels of differentiation (FST = 0.05) and no correlation with geographical distance, suggesting their derivation from a single glacial refuge. By contrast, continental regions revealed moderate levels of differentiation (FST = 0.27) and a strong correlation with geographical distance, indicating a much older history with several refugia likely. Two sympatric allozyme genotypes were found at three continental sites from Taylor Valley and were congruent with two mtDNA haplotypes, implying nonrandom breeding groups. Although haplotype sharing between one Ross Island site (Cape Bird) and one continental site (Granite Harbour) was identified, the clades showed mostly fragmented allopatric distributions. The extensive Pleistocene glaciations, in conjunction with limited dispersal opportunities, appear to have promoted isolation and divergence among the fragmented habitats. Furthermore, the McMurdo Sound appears to be an effective isolating barrier to dispersal. However, we suggest that the unaided dispersal capacity of G. hodgsoni is unlikely to account for the limited mixing of haplotypes across the McMurdo Sound and recent human- or bird-mediated dispersal is highly probable.

Invertebrates in ornithogenic soils on Ross Island, Antarctica

A habitat suitability model developed for soil biotic communities in the McMurdo Sound region, Antarctica predicts that soil moisture, organic carbon, and salinity exert control on the abundance and complexity of soil food chains. The model has been intensively tested in dry and carbon-poor soils of the Dry Valleys. To determine the influence of moisture and soil organic mater in wetter soils with high C content, invertebrates (nematodes, rotifers, and tardigrades) from soil samples collected in and near penguin rookeries on Ross Island (Cape Bird, Cape Crozier, and Cape Royds) were examined. Invertebrates were present in less than 50% of all collected soil samples. Although four nematode species were identified (Eudorylaimus antarcticus, Panagrolaimus davidi, Plectus antarcticus, Scottnema lindsayae), only populations of Panagrolaimus davidi were observed in soils collected from within penguin rookeries. Abundances of Panagrolaimus davidi and rotifers differed among rookeries, and year of sampling had a significant effect only on the populations of Panagrolaimus davidi. There were no temporal differences in soil moisture and soil chlorophyll a concentrations within each rookery or across rookeries. No invertebrates were correlated with soil moisture or chlorophyll a at the time of collection. Counter to our expectations, higher nutrient, organic matter, and moisture levels did not result in more abundant and diverse invertebrate communities in the rookery soils. It appears that excessive accumulations of nutrients, creating high soil salinity, may limit soil invertebrate presence within active rookeries.