Minimum effective release rate of antifoulants. 2. Effects of TBTCI, 2-furyl-n-pentyl ketone and coumaric acid at Snug harbor,Hawaii

Minimum effective release rate (MERR) of three antifoulants was examined at Snug harbor, Hawaii using a dynamic diffusion system. Among the three antifoulants tested TBTCI was found to be effective in preventing the settlement of Hydroides elegans and Crisea sp at 0.5 microg cm(-2) d(-1) flux rate. At a maximum flux rate of 10 microg cm(-2) d(-1) of 2-furyl-n-pentyl ketone larval settlement of both the test species were 32-36% reduced. More or less similar effect was seen at 10 microg cm(-2) d(-1) of coumaric acid. Non-toxic antifoulants, 2-furyl-n-pentyl ketone and coumaric acid exhibit specific activity against target species. MERR obtainedfor the three antifoulants is discussed.     

Larval biology of six species of the genus Conus (Gastropoda: Toxoglossa) in Hawaii,USA

Among the Hawaiian species in the genus Conus, larval life histories range from long-term planktotrophy to nonpelagic lecithotrophy. Prehatching developmental time, hatching size, pelagic period and total prejuvenile developmental time are all significantly correlated with egg size, while settling size is correlated with none of these larval characteristics. Therefore, larval life history patterns in the genus Conus appear to be determined by 2 sets of selective forces: the first influencing egg size, and the second stipulating settling size. As egg sizes become larger, hatching sizes and prehatching developmental time increase while pelagic periods and total prejuvenile developmental times decrease. Settling size appears to be related to water depth, but food requirements of newly metamorphosed juveniles may also be important.     

Growth and mortality of subtidal red sea urchins (Strongylocentrotus franciscanus) at San Nicolas Island,California, USA: problems with models

Red sea urchins, Strongylocentrotus franciscanus, were tagged with tetracycline in 1990 at subtidal sites off San Nicolas Island, California, USA. After one year in the field, the sea urchins were collected and growth increments were measured based on tetracycline marks, which indicated initial slow growth, a maximum rate, and finally a prolonged period of very slow growth. Small red sea urchins (4 cm diam) were estimated to be 3 to 4 yr old, which is much older than has previously been reported. It is estimated that about 12 yr would be required to attain 10 cm diam. Survival has previously been modeled assuming a constant rate. If the population of red sea urchins is assumed to be stable and stationary, annual survival rate was between 71 and 77% yr^-1. Census data for the two years of study have permitted annual survival to be estimated without assuming stable and stationary population structure. Under these conditions, annual survival rate was between 79 and 86% yr^-1. Analysis of transitions in the size distributions from 1990 to 1991 suggested that annual survival may have been sizespecific: 91% yr^-1 for individuals 1.1 to 4.0 cm diam, 82% yr^-1 for individuals 4.1 to 7.0 cm diam, and 63% yr^-1 for those of 7.1 to 10.0 cm diam. An alternative explanation to size-specific survival in our study is sizespecific immigration.     

Effects of Polinices duplicatus (Gastropoda: Naticidae) on infaunal community structure at Barnstable Harbor,Massachusetts, USA

Effects of the predatory naticid snail Polinices duplicatus (Say) on species composition, diversity and density, of the infauna of intertidal sand-flats at Barnstable Harbor, Massachusetts, USA, were studied using field experiments. Responses of molluscs (prey of P. duplicatus) and nonmolluscs (nonprey) were considered separately to distinguish between the effects of feeding and sediment disturbance during foraging. The fauna of 0.25 m² predator-exclusion cages, coarse-mesh cages, sieved areas, and controls was followed for 1 yr. Species associations within cages from which predators were excluded were denser, more diverse, and richer in molluscs than those in other treatments. Larger areas (3×3 m) of natural bottom were fenced and maintained as snail exclosures and enclosures during two feeding seasons. For both molluscs and nonmolluscs, diversity (H'), number of species, evenness (SD), and density all decreased with increasing snail density. Intense predation pressure on molluscs and selective feeding on thin-shelled bivalves, which were rare, removed individuals and species of molluscs from the community. Comparison of samples taken inside and outside trails made by snails showed that disturbance of the surface sediment layers by snails decreased the abundance of spionid polychaetes and total nonmolluscs. The sipunculan Phascolopsis gouldi and the bivalve Gemma gemma dominated the community in all experimental treatments and were little affected by the activity of Polinices duplicatus. Predation and disturbance by snails lowered community diversity by removing individuals of the less abundant species, and generally maintained population densities below the level where strong competition would occur.     

Growth rates of summer nanoplankton (<10 μm) populations in lower Narragansett Bay,Rhode Island,USA

Growth rates of summer (June–September) phytoplankton assemblages and constituent species were measured in 30 diffusion culture experiments. Size-fractionated (<10 m) phytoplankton assemblages were incubated in situ or under simulated in-situ conditions in outdoor tanks connected to a running seawater system. Doubling rates of important species and groups (such as microflagellates) were compared to community biomass doubling rates estimated from ¹⁴C uptake and changes in chlorophyll a concentrations. Division rates of dominant diatom species generally equalled or exceeded community biomass doubling rates, while those of flagellates and non-motile ultraplankters were slower. Maximum division rates of sixteen common diatom species exceeded 2.1 divisions d^-1, while nine had maximum division rates in excess of 3 d^-1. Mean division rates of 12 diatom species exceeded 1 d^-1. Maximum division rates of flagellated species, uncharacterized microflagellates and non-motile ultraplankton assemblages were 2.1, 1.5 and 1.4 d^-1, respectively. Microflagellate and non-motile ultraplankton assemblage doubling rates were less than 0.5 d^-1 in over half of all growth experiments.     

Post-nesting migrations of green turtles (Chelonia mydas) at Wan-An Island, Penghu Archipelago, Taiwan

 During the summers of 1994 to 1997, eight green turtles (Chelonia mydas) nesting at Wan-An Island, PengHu Archipelago, Taiwan, were equipped with satellite-telemetry transmitters. Using the Argos-linked satellite system, the turtles' migration routes were tracked until the transmissions stopped. The turtles migrated widely on the continental shelf to the east of Mainland China. The migration distances ranged from 193 to 1909 km, and the migration speeds from 1.2 to 2.8 km h⁻¹. The turtles apparently utilize several coastal areas as temporal residential foraging sites, and their migrations consist of both trans-oceanic and coastal legs. The wide distribution of the foraging sites of the turtles comprising this rookery reflects the extent to which the green turtle migrates in northeast Asia; regional and international cooperation will therefore be needed to conserve this declining population. Received: 14 December 1998 / Accepted: 8 May 2000     

Sex differences in mass loss rate and growth efficiency in Antarctic fur seal (Arctocephalus gazella) pups at Macquarie Island

We investigated the relationship between the mass gained by a pup during a period of maternal attendance (as an index of milk intake) and the duration of the preceding foraging trip in relation to the mass-specific rate of mass loss during fasting periods and the growth rate of Antarctic fur seal (Arctocephalus gazella) pups at Macquarie Island. We found that (1) serially weighed male pups grew significantly faster than females pups and that (2) fasting female pups lost mass at a significantly higher rate (2.55% day⁻¹) than male pups (2.12% day⁻¹) of the same mass; (3) during periods of maternal attendance, there were no intersexual differences in the amount of mass gained by pups of the same size, hence (4) female pups required a higher daily mass gain to grow at the same rate as male pups. Our results show that intersexual differences in growth rate may be accounted for by intersexual differences in mass-specific rate of mass loss, because females lost 0.42% more of their total mass per day (i.e. 4.2 g kg⁻¹ day⁻¹) compared with male pups of the same body mass. Despite intersexual differences in growth rates, our results indicate equality of maternal expenditure between the sexes. Intersexual differences in the rate of mass loss may be due to differences in the metabolic rate, activity level and/or body composition of male and female pups. Received: 19 August 1998 / Received in revised form: 19 April 1999 / Accepted: 19 April 1999     

Dietary overlap among coral-feeding butterflyfishes (Chaetodontidae) at Lizard Island,northern Great Barrier Reef

This study documented the range of corals, and other prey types, consumed by 20 species of butterflyfishes, which co-occur at Lizard Island, northern Great Barrier Reef, Australia. Six species (Chaetodon aureofasciatus, C. baronessa, C. lunulatus, C. plebius, C. rainfordi and C. trifascialis) fed almost exclusively on scleractinian corals, and a further eight species (C. citrinellus, C. kleinii, C. lunula, C. melannotus, C. rafflesi, C. speculum, C. ulietensis, and C. unimaculatus) took a significant proportion of their bites from corals. The other six species (C. auriga, C. ephippium, C. lineolatus, C. semeion, C. vagabundus, and Chelmon rostratus) rarely consumed coral, but fed on small discrete prey items from non-coral substrates. Coral-feeding butterflyfishes consumed a wide range of corals. Chaetodon lunulatus, for example, consumed 51 coral species from 24 different genera. However, there was up to 72% dietary overlap between coral-feeding butterflyfishes, with 11/14 species feeding predominantly on Acropora hyacinthus or Pocillopora damicornis. The most specialised corallivore, C. trifascialis, took 88% of bites from A. hyacinthus. Chaetodon trifascialis defend territories encompassing one or more colonies of A. hyacinthus, and may have prevented other species such as C. lunulatus from feeding even more extensively on this coral. This study has shown that coexistence of coral-feeding butterflyfishes occurs despite an apparent lack of partitioning of prey resources. While different coral-feeding butterflyfishes were more or less selective in their use of different coral prey, virtually all species fed predominantly on A. hyacinthus or P. damicornis.Electronic Supplementary Material Supplementary material is available for this article at     

Metal survey of the marine clam Pitar morrhuana collected near a Rhode Island (USA) electroplating plant

Benthic fauna were collected from 17 stations in mid-Narragansett Bay, Rhode Island, during September 1973 from the vicinity of the recently-closed Quonset Point electroplating facility. Despite repeated sampling, most of the 14 species of molluscs taken, including the widgeon clam Pitar morrhuana, were absent from stations in the immediate vicinity of plant outfalls. In general, P. morrhuana captured near outfall sites exhibited high moisture content and elevated body burdens of silver, cadmium, cobalt, chromium, copper, iron, manganese, nickel, lead, and zinc compared to clams of similar size from more distant stations. Observed changes in metal concentrations and moisture content of P. morrhuana were probably attributable to plant operations during the preceding 30 years.     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Ecology of juvenile hawksbills (Eretmochelys imbricata) at Buck Island Reef National Monument, US Virgin Islands

Surveys of juvenile hawksbills around Buck Island Reef National Monument, US Virgin Islands from 1994 to 1999 revealed distributional patterns and resulted in a total of 75 individual hawksbill captures from all years; turtles ranged from 23.2 to 77.7 cm curved carapace length (CCL; mean 42.1 ± 12.3 cm SD). Juveniles concentrated where Zoanthid cover was highest. Length of time between recaptures, or presumed minimum site residency, ranged from 59 to 1,396 days (mean 620.8 ± 402.4 days SD). Growth rates for 23 juveniles ranged from 0.0 to 9.5 cm year^?1 (mean 4.1 ± 2.4 cm year^?1SD). Annual mean growth rates were non-monotonic, with the largest mean growth rate occurring in the 30–39 cm CCL size class. Gastric lavages indicated that Zoanthids were the primary food source for hawksbills. These results contribute to our understanding of juvenile hawksbill ecology and serve as a baseline for future studies or inventories of hawksbills in the Caribbean.     

Islands at bay: rising seas,eroding islands,and waterbird habitat loss in Chesapeake Bay (USA)

Like many resources in the Chesapeake Bay region of the U.S., many waterbird nesting populations have suffered over the past three to four decades. In this study, historic information for the entire Bay and recent results from the Tangier Sound region were evaluated to illustrate patterns of island erosion and habitat loss for 19 breeding species of waterbirds. Aerial imagery and field data collected in the nesting season were the primary sources of data. From 1993/1994 to 2007/2008, a group of 15 islands in Tangier Sound, Virginia were reduced by 21% in area, as most of their small dunes and associated vegetation and forest cover were lost to increased washovers. Concurrently, nesting American black ducks (Anas rubripes) declined by 66% , wading birds (herons-egrets) by 51%, gulls by 72%, common terns (Sterna hirundo) by 96% and black skimmers (Rynchops niger) by about 70% in this complex. The declines noted at the larger Bay-wide scale suggest that this study area maybe symptomatic of a systemic limitation of nesting habitat for these species. The island losses noted in the Chesapeake have also been noted in other Atlantic U.S. coastal states. Stabilization and/or restoration of at least some of the rapidly eroding islands at key coastal areas are critical to help sustain waterbird communities.     

Diel patterns of emergence of crinoids (Echinodermata) from within a reef at Lizard Island,Great Barrier Reef,Australia

Periods of emergence of nine species (88 individuals) of crinoids (Comanthus parvicirrus, Clarkcomanthus albinotus, Comaster multifidus, Oxycomanthus comanthipinna, Oligometra serripinna, Comanthus gisleni, Comanthus wahlbergi, Comatula purpurea, and Oxycomanthus exilis) were monitored at Lizard Island, Queensland, over seven days in March 1983. One species (O. serripinna) was fully exposed and the others partially exposed. Two patterns of emergence were species-specific: emergent both day and night, and emergent only at night. Intraspecific exposure patterns were generally synchronous in six species and asynchronous in three species. The number of species visible was relatively constant during all dives, while at least twice as many individuals were visible at night compared to day. Degree of emergence was determined as the number and length of arms extended into the water column. Except for species fully exposed, degree of crinoid emergence was generally minimal at midday and maximal at twilight and night. Increases in both number of crinoids visible, and the amount of an individual's emergence, corresponded to decreasing light intensities, even during daytime. It is suggested that the patterns of exposure are a response to increased prey abundance at twilight and night, and also a means of avoiding diurnal predators. Gonads on the longest arms of some partially exposed crinoids were not extended into the water, thus protecting the gonads from predators.     

Effects of kraft-mill effluents on benthic macrophyte assemblages in a shallow-bay system (Apalachee Bay,North Florida,USA)

A 14-month study was carried out to determine the impact of kraft-mill effluents on the offshore benthic macrophyte distribution in a shallow north Florida Bay (USA). A polluted river drainage system was compared to an adjoining (unpolluted) one. The affected area was characterized by elevated levels of color and turbidity. Inshore areas associated with high levels of KME were characterized by severely reduced benthic macrophyte biomass, reduced numbers of species/unit area, and altered species composition when compared to control stations. Areas of chronic impact also had reduced levels of biomass, although the total number of species taken were not significantly different from the controls. In polluted areas, red and brown algae were proportionately more abundant than chlorophytes and spermatophytes. Dominance generally was reduced in areas of the Bay affected by KME. Except for areas of acute impact, there was no significant difference in species diversity (H) between polluted and unpolluted portions of the Bay. There were uniform increases in the total number of species on either side of the affected drainage system. Such areas were considered to be transition zones between the polluted and unpolluted parts of Apalachee Bay. Benthic macrophyte distribution, in terms of biomass and species (community) composition, was considered an important indicator of the impact of KME on the shallow coastal systems in question. The pattern of macrophyte species composition reflected various water-quality parameters. Species normally inhabiting deeper water were found in areas of increased turbidity and color. It was postulated that reduction of normal dominants such as Thalassia testudinum and Halimeda incrassata allowed the colonization of such areas by opportunistic species. This would explain the maintenance of comparable (total) numbers of species in polluted areas and the lack of effect of KME on species diversity even though biomass was severely restricted. These parameters were analogous to the epibenthic fish distribution in the respective drainage areas. Near-shore coastal systems in Apalachee Bay thus were affected by gradients in water quality in addition to natural (seasonal) fluctuations in key physical and chemical parameters. The benthic plant assemblages reflected variations in dominance, the occurrence of opportunistic species, and ecological diversity that resulted in a continuum of disturbance phenomena ranging from sparsely distributed groups in grossly polluted systems to well developed plant assemblages in areas that remain unaffected by KME.     

Recovery of benthic communities at Macquarie Island (sub-Antarctic) following a small oil spill

Intertidal communities at Macquarie Island (sub-Antarctic) were re-evaluated seven years after a small oil spill resulting from the grounding of the “Nella Dan”, and six years after the first assessment of biological impact. Sampling was conducted to evaluate community structure in three zones of the exposed rocky shore (upper red, kelp, and lower red) and in samples of Durvillaea antarctica holdfasts. There were no significant differences between the community structure in oiled and control locations in any of the three shore zones, but holdfast macrofaunal communities at oiled sites still showed evidence of impact. Holdfast community-structure in samples from heavily oiled sites showed moderate levels of recovery, with increased abundances of species which were considered sensitive to the oiling in the first post-impact studies and decreased abundances of opportunistic polychaete and oligochaete worms. In contrast, samples collected from the moderately oiled location at Secluded Bay showed little evidence of recovery. Holdfasts at this site were filled with sediment containing traces of diesel oil, and the macrofaunal community was dominated by opportunistic worms. This study clearly indicates that even small incidents of anthropogenic perturbation can have long-lasting consequences for marine communities at Macquarie Island. Received: 23 July 1997 / Accepted: 5 February 1998     

A multiple instrument approach to quantifying the movement patterns and habitat use of tiger (Galeocerdo cuvier) and Galapagos sharks (Carcharhinus galapagensis) at French Frigate Shoals, Hawaii

We equipped individual tiger (Galeocerdo cuvier Péron and Lesueur, 1822) and Galapagos (Carcharhinus galapagensis Snodgrass and Heller, 1905) sharks with both acoustic and satellite transmitters to quantify their long-term movements in the Papahanaumokuakea Marine National Monument (Northwestern Hawaiian Islands). Tiger sharks exhibited two broad patterns of behavior. Some individuals were detected at French Frigate Shoals (FFS) year round, whereas others visited FFS atoll in summer to forage on fledging albatross, then swam thousands of kilometers along the Hawaiian chain, or out into open ocean to the North Pacific transition zone chlorophyll front, before returning to FFS in subsequent years. These patterns suggest tiger sharks may use cognitive maps to navigate between distant foraging areas. Different patterns of spatial behavior may arise because cognitive maps are built up through individual exploration, and each tiger shark learns a unique combination of foraging sites. Galapagos shark detections were all associated with FFS, suggesting these sharks may be more resident around oceanic islands. Both Galapagos and tiger sharks primarily used the mixed layer (<100 m depth) and made occasional deeper dives through the thermocline down to 680 m. Results show reef-associated sharks utilize a wide variety of habitats ranging from shallow atoll lagoons to deep reefs and open ocean and may provide important trophic links between these habitats.     

Karyology of the toadfish Porichthys plectrodon (Jordan and Gilbert, 1882) (Batrachoididae) from Margarita Island,Venezuela

This paper reports the results of cytogenetic analyses carried out on Porichthys plectrodon using conventional Giemsa staining, C-banding and silver staining techniques. A diploid chromosome count of 2n=44 was observed, consisting of 8 metacentric, 10 submetacentric, 6 subtelocentric and 20 acrocentric chromosomes. Differences in length made it possible to identify homologous chromosomes within the metacentric group. Constitutive heterochromatin was distributed as large pericentromeric blocks in pairs 1 and 2, while the rest of the chromosomes were marked in centromeric regions, some more conspicuously than others. One pair of small-sized acrocentric NOR-bearing chromosomes (21) was identified by the nucleolar regions located terminally on their short arms.Communicated by P.W. Sammarco, Chauvin     

Diving behaviour of female northern rockhopper penguins, Eudyptes chrysocome moseleyi, during the brooding period at Amsterdam Island (Southern Indian Ocean)

The pattern and characteristics of diving in 14 female northern rockhopper penguins, Eudyptes chrysocome moseleyi, were studied at Amsterdam Island (37°50′S; 77°31′E) during the guard stage, using electronic time–depth recorders. Twenty-nine foraging trips (27 daily foraging trips and two longer trips including one night) with a total of 16 572 dives of ≥3 m were recorded. Females typically left the colony at dawn and returned in the late afternoon, spending an average of 12 h at sea, during which they performed ∼550 dives. They were essentially inshore foragers (mean estimated foraging range 6 km), and mainly preyed upon the pelagic euphausiid Thysanoessa gregaria, fishes and squid being only minor components of the diet. Mean dive depth, dive duration, and post-dive intervals were 18.4 m (max. depth 109 m), 57 s (max. dive duration 168 s), and 21 s (37% of dive duration), respectively. Descent and ascent rates averaged 1.2 and 1.0 ms⁻¹ and were, together with dive duration, significantly correlated with dive depth. Birds spent 18% of their total diving time in dives reaching 15 to 20 m, and the mean maximum diving efficiency (bottom time:dive cycle duration) occurred for dives reaching 15 to 35 m. The most remarkable feature of diving behaviour in northern rockhopper penguins was the high percentage of time spent diving during daily foraging trips (on average, 69% of their time at sea); this was mainly due to a high dive frequency (∼44 dives per hour), which explained the high total vertical distance travelled during one trip (18 km on average). Diving activity at night was greatly reduced, suggesting that, as other penguins, E. chrysocome moseleyi are essentially diurnal, and locate prey using visual cues. Received: 9 December 1998 / Accepted: 3 March 1999     

The delineation of two plankton communities from one sampling site (Fire Island Inlet,Long Island,N.Y.)

Zooplankton and phytoplankton communities were observed and recorded at Fire Island Inlet from 1971–1974. Species typical of a temperate, neritic environment were found. Sampling by net and bottle at several stations in the area revealed two populations, one representative of the bay water and the other representative of the ocean water. It was further found, by comparing frequency of occurrence relative abundance, rank order and indicator species, that these two populations could be monitored at one sampling station (Oak Beach) by sampling at the appropriate tidal intervals, i.e., midtide after slack current, on both the ebb and flow tides. Observations from one site of plankton communities representative of bay water and of ocean water, as they move into and out of the Inlet, could be of significance in monitoring the effects of proposed man-made changes in the environment. It is possible that other coastal areas could benefit by such a combined approach.Based on a thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, New York University, New York, N.Y.     

Response of Ecklonia radiata (Laminariales) to light at 15°C with reference to the field light budget at Goat Island Bay,New Zealand

Gametophytes of Ecklonia radiata (C.Ag.)J.Ag. grew in culture at 15°C under daily quantum doses ranging from 0.86 to 360 cE m^-2. Growth rates increased with quantum doses up to 40 cE m^-2 d^-1, then reproduction began and the relative growth constant declined while ovum release came earlier with increasing light up to about 100 cE m^-2 d^-1. Above 100 cE m^-2 d^-1 there were no consistent trends with increasing light, except that at the higher quantum doses, fertile female plants had fewer and larger cells and therefore fewer potential ova. Reproduction varied with daily quantum dose rather than with daylength. Given the same daily dose, plants grew fastest in low photon flux density, long daylength conditions. Gametophytes grown in the field developed at similar rates to those in culture. Gametophytes survived seven months of darkness at 10°C but died after one week of darkness at 20° to 23°C. Sunlight of 1 000 E m^-2 s^-1 was fatal to gametophytes and to sporophytes under 2 mm long after 10 to 15 min. Light budgets were prepared for plants growing at 7-and 15-m depths from 1976 to May 1980 in Goat Island Bay, New Zealand (Lat.36° 16S, Long. 174° 48E). Underwater light was measured under various environmental conditions. Relationships between transmission of light through the sea, data from diving visibility records and continuous surface meteorological records were studied. Approximations were made of the average percentage of surface light transmitted to 7 and 15 m over half-monthly periods. By applying these average transmittance values to the records of surface incident light, the average daily quantum doses were calculated. Light on open bottom in Goat Island Bay may sometimes be limiting for gametophyte reproduction in winter at 15-m depth. At depths less than 7 m, summer photon flux densities may reach damaging levels.