Sampling design trade-offs in occupancy studies with imperfect detection: examples and software.

Researchers have used occupancy, or probability of occupancy, as a response or state variable in a variety of studies (e.g., habitat modeling), and occupancy is increasingly favored by numerous state, federal, and international agencies engaged in monitoring programs. Recent advances in estimation methods have emphasized that reliable inferences can be made from these types of studies if detection and occupancy probabilities are simultaneously estimated. The need for temporal replication at sampled sites to estimate detection probability creates a trade-off between spatial replication (number of sample sites distributed within the area of interest/inference) and temporal replication (number of repeated surveys at each site). Here, we discuss a suite of questions commonly encountered during the design phase of occupancy studies, and we describe software (program GENPRES) developed to allow investigators to easily explore design trade-offs focused on particularities of their study system and sampling limitations. We illustrate the utility of program GENPRES using an amphibian example from Greater Yellowstone National Park, U.S.A.     

Polonium-210 and lead-210 in Antarctic marine biota and sea water

Concentrations of the naturally-ocurring radionuclides ²¹⁰Po and ²¹⁰Pb were measured in krill (Euphausia superba), mesozooplankton, phytoplankton and sea water collected during the South African SIBEX cruise to the Antarctic in autumn 1984. The data reported constitute the first substantial measurements on ²¹⁰Po and ²¹⁰Pb in such samples in the Antarctic Ocean. The concentrations of ²¹⁰Po in mesozooplankton and phytoplankton are unexceptional in comparison with those from other oceans. The SIBEX E. superba, however, have higher levels of ²¹⁰Po than usually found in euphausiids. The ²¹⁰Po data, combined with reasonable estimates of biological quantitites such as the fractional assimilation, are used to obtain information about the diet of E. superba. It is suggested that the higher ²¹⁰Po in the SIBEX E. superba reflects a change from an almost entirely phytoplanktonic diet in summer to a more omnivorous diet as winter approaches. The data show that there are allometric relationships between the ²¹⁰Po content of euphausiids and animal size; these are discussed briefly. The limited sea-water data presented are characterized by unusually high ²¹⁰Po:²¹⁰Pb activity ratios and need further investigation.     

Metabolic constraints on burst-swimming in the Antarctic teleost Notothenia neglecta

Immediate-energy-supply pathways were analysed during burst-swimming in white muscle of the Antarctic fish Notothenia neglecta, (collected during Austral summer 1984 near Signy Island, South Orkneys). Selected enzyme activities were determined, and tissue metabolites were measured, before and after 3 min of burst-work. The activity of glycolytic enzymes is very low compared to that in other fish, while the activities of creatine phosphokinase, adenylate kinase and AMPdeaminase are relatively high. Concentrations of metabolites associated with cellular energy status declined with exercise. At the same time, there was no change in the concentrations of most of the glycolytic metabolites measured (including glycogen, glucose and lactate), while [creatine phosphate] declined and [inosine monophosphate] increased. These observations were used to propose that the potential for ATP generation via creatine-phosphate hydrolysis is enhanced in N. neglecta, while glycolytic capacity is relatively reduced. This is an unusual pattern of ATP production and places severe constraints upon the behavior of the fish. While the capacity for very short-term, high-power-output swimming is increased, the capacity for medium-duration swimming is reduced. Ecological repercussions are discussed.     

Incorporation of organic aggregates by marine mussels

Two marine mussels, Geukensia demissa (Dillwyn) and Mytilus edulis (L.) collected in 1990 in Old Silver Beach, Falmouth, Massachusetts, incorporated nitrogen when fed ¹⁵N-labelled organic aggregates produced from dissolved organic nitrogen released by the brown sea-weed Fucus vesiculosis. Uptake of ¹⁵N on the aggregate diet was linear over the course of 24 h, and unincorporated ¹⁵N was eliminated from the gut after 48 h. Both species of mussels incorporated approximately five times more N when they were fed organic aggregates than when they were fed either ¹⁵N-labelled dissolved organic material (DOM) or particulate detritus, both of which were also derived from the seaweed. Nitrogen uptake was greatest in controls fed the diatom Thalassiosira weissflogii; mussels fed phytoplankton incorporated seven times more nitrogen than those fed aggregates. However, aggregates could supply an estimated 7 to 14% of the N requirements for both mussels, whereas DOM or particulate detritus could only supply 1 to 3%. These data provide evidence that a food web pathway exists from primary producer to released dissolved organic nitrogen to microbial organic aggregate to metazoan consumer, and, further, that it can be more important in a detrital food web than either particulate detritus or DOM.     

Temporal stability of gene frequencies within genetically heterogeneous populations of the queen scallop Aequipecten (Chlamys) opercularis

The widely held assumption that gene frequencies within samples of multi-aged organisms are temporally stable has rarely been tested. In order to test this assumption in the commercially exploited bivalve scallop Aequipecten (Chlamys) opercularis (L.), 12 populations were sampled from around the UK over the period 1988–1990 and each was assayed at four putative polymorphic enzyme loci. Most populations were comprised of 3 to 4 year-classes, distinguishable by annual rings laid down on the shell. Three alleles from each locus were used as discriminant-function variables. Discriminant-function analysis based on allele frequencies averaged for each inividual year-class (45 demes) revealed highly significant inter-site heterogeneity. A posteriori classification of individual year-classes based on the discriminant analysis resulted in assignment of year-classes to the correct home site in 80% of cases. If allozyme neutrality is assumed, the phenomenon of predominant local allele frequency-stability through time within populations which are genetically heterogeneous from one site to another raises the possibility that queen scallop populations may be self-recruiting. Despite a planktonic larval phase lasting up to several weeks, this possibility is not excluded by the available hydrographic data.     

Spatial segregation amongst goby species within an Australian estuary,with a comparison of the diets and salinity tolerance of the two most abundant species

The present study was undertaken to determine whether the various species of gobies that are found within the large Swan Estuary in south-western Australia are segregated within that system, and to attempt to determine the basis for any differences in their spatial distributions. The Swan Estuary comprises a long entrance channel (lower estuary), two wide basins (middle estuary) and the saline reaches of the tributary rivers (upper estuary). A total of 26232 gobies, representing seven species, was collected using a 3 mm-mesh seine net at 15 sites throughout this estuary on at least one occasion monthly over seven consecutive seasons between September 1983 and March 1985. Favonigobius lateralis and Pseudogobius olorum contributed 47.0 and 47.8%, respectively, to the total catch of gobies at all sites. The densities of each species at each site were used to determine the relative contribution of each species to the gobiid fauna at each of the sites in the lower, middle and upper estuary. Comparisons of these data with those published on the distribution and abundance of gobiid larvae confirmed that F. lateralis, which was found predominantly in the lower estuary, is a marine species that spawns in high salinities near the estuary mouth or in inshore coastal waters. In contrast, the life cycle of P. olorum and Papillogobius punctatus are typically completed within the saline reaches of the upper estuary, and that of Arenigobius bifrenatus within both this region and parts of the middle estuary where the substrate is particularly soft. Afurcagobius suppositus also spawns in this area, as well as in fresh water. Tridentiger trigonocephalus, represented by only eight individuals, is an introduced, marine species that was found mainly in the lower estuary. A single representative of the marine species Callogobius depressus was caught. The relatively low numbers of gobies caught in the middle estuary, where they contributed only about 3.5% to the total number of all gobies at all sites, may represent an aversion to the presence of rougher waters in the large basins. Circumstantial evidence suggests that the sandy substrate and consistently high salinities found in the lower estuary are preferred by F. lateralis, whereas the silty surface to the substrate and lower salinities of the upper estuary are preferred by Pseudogobius olorum. Densities of three of the four most abundant species were higher in either spring or summer than in winter, reflecting the influx of 0 + recruits, and possibly also the tendency for species in estuaries to congregate in the shallows during the warmer periods of the year. F. lateralis fed mainly on polychaetes and crustaceans, whereas P. olorum ingested predominantly algae, reflecting differences in mouth morphology and feeding behaviour, rather than the type of food available.     

Occurrence and significance of nitrogen transport in the brown alga Laminaria digitata

Nitrogen transport in a Laminaria digitata (Huds.) Lamour population growing at Arbroath, Scotland has been investigated (during the period 3 June 1981 to 3 July 1981), using ¹⁵N as tracer. NO ₃ ^- was assimilated both by the blade meristem and by the mature blade. NO ₃ ^- uptake by the blade meristem alone was insufficient to supply the nitrogen demand for growth. This additional demand was met by nitrogen transport from the mature blade to the meristem. It was estimated that 70% of the nitrogen demand of the meristematic region was supplied by nitrogen transport from the mature blade. Although transport occurred, the size of the endogenous pool of stored nitrogen in the mature blade did not change, the quantity of nitrogen transported being equivalent to the amount of NO ₃ ^- assimilated during the experimental period. The possible co-transport of nitrogen and carbon is discussed.     

Age-and sex-dependent patterns of aggregation in the tropical gastropod Strombus luhuanus

The spatial dispersion of the tropical coral sand-dwelling mesogastropod Strombus luhuanus Linné, 1758 was studied at several sites in two localities (Australian Great Barrier Reef and southern Papua New Guinea), from September 1980 to February 1983. S. luhuanus usually occurred in local aggregations, many of which were relatively dense with discrete boundaries: Four types of aggregation are described: mixed age-class, juvenile, mating, and clusters. The former two are termed colonies, because they persisted over time and exhibited some coordinated movements. Mixed age-class colonies contain individuals of all ages, but within them the different ageclasses are frequently spatially segregated from each other. This was sometimes due to the younger age-classes' preference for shallower areas, but also occurred in areas of constant depth. Juvenile colonies are relatively smaller in area but higher in density, and are dominated by a single size-class. Mating aggregations are concentrations of copulating individuals, in which competition between males for access to females occurs. Clusters are concentrations of inactive individuals (juvenile and adult) piled together in close contact. All types of aggregation were concentrated in only a part of what appeared to be the suitable habitat. Most aggregations' boundaries shifted over time within the habitat. We conclude that intraspecific attraction must be an important factor which maintains the spatial structure of all aggregations, although habitat specificity sets the larger area in which they move. Processes which may underlie this pattern, and some of its implications, are discussed.     

Unstable processes in the metabolic rate measurements in flow-through systems

Measurements of metabolic rate within a flow-through system generally begin only after the difference between input and output concentration has become constant. This limits the use of such a system to objects with high metabolic rates. The processes in the flow-through system were described mathematically and the differential equations obtained are analytically resolved for different situations. They can be used for the calculation of metabolic rates for any time after the onset of measurements. The flexibility of a flow-through system and its suitability for physiological and ecological measurements are significantly increased.     

The role of isopods and amphipods in the initial fragmentation of eelgrass detritus in Nova Scotia,Canada

Daytime observations on the isopods Idotea phosphorea and I. baltica and the amphipod Gammarus oceanicus held in laboratory microcosms showed that I. phosphorea and G. oceanicus spent 45% and 30% respectively, of their active time feeding on dead, intact eelgrass leaves which had been recently released from plants. I. baltica spent 41% of its active time consuming intact green leaves. The shredding of intact dead leaves by I. phosphorea and G. oceanicus resulted in production of small detrital particles which were liberated from the faeces of the invertebrates and this type of feeding led to the breakdown of whole leaves. Field experiments which separated the effects of shredding by invertebrates and grinding by waves and ice on the loss of weight from leaf packs showed that relative to controls isopods significantly increased weight loss from dead leaves. Loss of weight from leaf packs exposed to both biotic and physical shredding forces was not significantly different from that found on those exposed only to shredding by isopods. However, trends in the data indicated that fragmentation of whole, dead leaves in the field probably is a result of the synergistic effects of shredding by invertebrates and physical factors, particularly ice grinding. The role played by invertebrates in fragmenting intact, dead leaves is discussed in the light of energy flow and nutrient cycling within seagrass systems.