Cephalopod prey of the wandering albatross Diomedea exulans

Cephalopod beaks from the stomach contents of wandering albatross (Diomedea exulans L.) chicks from Bird Island, South Georgia, were sampled between May and September in 1983 and 1984. Lower beaks were identified and measured, and allometric data were used to calculated mantle length and biomass of the species consumed. A total of 3421 lower beaks were examined, representing 35 species in the 1983 sample and 45 species in the 1984 sample. Eight of the twenty families contributed over 95% of the biomass. In 1984 there were less Onychoteuthidae and more Ommastrephidae than in 1983 and a decrease in the number of species known to occur south of the Antarctic Polar Front. There was a difference in the size-frequency distribution of the cephalopod diet in the two years; in 1984 there was a higher frequency of intermediate-sized specimens, reflecting the greater importance of ommastrephids, especially Illex sp. The energy content of cephalopods in 1984 may have been greater than in 1983. Serial sampling of cephalopod beaks during the austral winter did not reveal evidence of growth. By the age of 200 d, wandering albatross chicks have consumed a total of approximately 100 kg wet weight of cephalopods each.     

Inter-annual variation in the cephalopod component of the diet of the wandering albatross, <Emphasis Type="Italic">Diomedea exulans</Emphasis>, breeding at Bird Island,South Georgia

Cephalopods play an important role in the diet of many predators of the Southern Ocean. We investigated the cephalopod component of the diet of the wandering albatross, during breeding at South Georgia, from 269 boluses containing more than 34,000 beaks (corresponding to 19,452 individual cephalopods), collected between 1989 and 1999, and assessed the inter-annual variability of the cephalopod species, the level of scavenging on cephalopods by wandering albatrosses and the relationships between cephalopod availability and wandering albatross breeding parameters (breeding period, breeding success, fledging period, fledging success, egg mass and chick mass). We also proposed possible foraging areas of wandering albatrosses based on the cephalopods eaten. The cephalopod component of the diet of wandering albatrosses was relatively stable over the 11 years of the study. By number of lower beaks, three species predominated in the cephalopod component of the diet: Kondakovia longimana (29.5% of the total number of lower beaks; family Onychoteuthidae), Taonius sp. (20.4%; family Cranchiidae) and Histioteuthis sp. B (19.5%; family Histioteuthidae). K. longimana was consistently the predominant cephalopod species by mass in every year of the study (range 60.2-88.7% of mass contribution to diet). A minimum estimate of 30% of the number of cephalopods scavenged corresponded to 85% of the total mass of the cephalopods that contributed to the diet. Wandering albatrosses fed consistently more on "Antarctic" cephalopods than on "sub-Antarctic" or "subtropical" cephalopods in all years of the study, suggesting that Antarctic waters are an important foraging area for wandering albatrosses. Although some significant correlations between cephalopod abundance and wandering albatross breeding parameters existed (e.g. correlation between Taonius sp. by number and fledging success), none included K. longimana. When comparing groups of variables (using canonical analysis), no correlations were found between the most important cephalopod species (by number and by mass), total mass of squid consumed, cephalopod diversity index (H) for each year, and wandering albatross breeding parameters. This may reflect the possibility that other components in the diet (e.g. fish and carrion) are more important or, more likely, that the consistency across years of the wandering albatross breeding performance indicates that it is well buffered against fluctuations in prey availability.     

Genetic evidence of population heterogeneity and cryptic speciation in the ommastrephid squid Martialia hyadesi from the Patagonian Shelf and Antarctic Polar Frontal Zone

Horizontal starch gel electrophoresis was used to investigate levels of genetic differentiation between four samples of the nominate squid species Martialia hyadesi Rochbrune and Mabille, 1889, obtained from regions of the Patagonian Shelf and Antarctic Polar Fron-tal Zone over 1000 km apart. M. hyadesi is an ecologically important South Atlantic ommastrephid squid and it is probable that, in the future, fishing effort will be increasingly directed towards this species. Details regarding the population structure of the species are therefore required. In comparison with the other three samples of M. hyadesi, one of the samples from the Patagonian Shelf (PAT 89II) exhibited fixed allelic differences at 16 of the 39 enzyme loci which were resolved (genetic identity, I=0.51). This high level of genetic differentiation contradicts the apparent morphological similarity between samples, indicating the presence of a cryptic or sibling congeneric species. Deviations from Hardy-Weinberg equilibrium and significant differences in allele distribution were also detected within and between the other three putative M. hyadesi samples, suggesting that the species fails to maintain effective panmixia across its geographical range. The occurrence of both temporal (1986 cf. 1989) and geographic structuring within the species complex is consequently indicated, caused possibly by an overlap of reproductively isolated stocks (stock mixing) outside their respective breeding areas. Low levels of genetic variability were detected throughout the samples examined, estimates of average heterozygosity per locus within the two species detected being in the order of 0.01 and 0.002. These values are discussed in relation to levels of genetic variability reported for other squid species, and in comparison with values typically expected for marine invertebrates.     

Extra-pair fertilizations in mountain white-crowned sparrows

Summary A population of white-crowned sparrows (Zonotrichia leucophrys oriantha) located near Tioga Pass, in the central Sierra Nevada of California, was studied during the 1985 and 1986 breeding seasons to determine how often chicks were not the offspring of both adults who reared them. Blood samples were obtained from 35 nesting pairs and feather pulp specimens were collected from all 110 young they fledged. Electrophoretic analyses of 4 polymorphic loci revealed that a minimum of 9 nests (26%) contained a chick that could not have been conceived in a mating between the attending pair, and that at least 15 chicks (14%) were not offspring of both attending adults. Using two recently developed probability indices, it was estimated that 34%–38% of chicks hatched at the study site were not offspring of one of their putative parents. Because there was no evidence that intraspecific brood parasitism occurred, all the mismatched young apparently resulted from extrapair copulations. Our data, along with similar results from other species, raise new questions about sexual selection and patterns of parental care in monogamous birds.     

Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

Cooperative breeding in purple gallinules: the role of helpers in feeding chicks

Summary The participation of breeders and helpers in the feeding of 21 broods of chicks was studied in a population of cooperatively breeding purple gallinules (Porphyrula martinica). In the breeding group, all birds over the age of 2 months fed chicks. Female breeders fed chicks at the highest rate, followed by male breeders and adult helpers, old juvenile helpers, and young juvenile helpers. The amount of food breeders fed chicks was independent of the number of helpers in the breeding group. However, breeders made fewer feeding visits when they had helpers. Male and female breeders spent similar amounts of time feeding chicks. Helpers spent significantly less time feeding chicks than did breeders. As helpers grew older they fed chicks at a faster rate, made more feeding visits and spent more time feeding chicks. Analysis of variance was used to determine which variables explained the variation in the brood feeding rate (amount of food delivered to an entire brood during one observation). Age of chicks had a significant nonlinear effect, and size of brood and number of helpers had significant linear effects on the brood feeding rate. Chicks in groups with helpers received more food and were accompanied for longer periods of time than chicks in groups without helpers; either or both of these factors may have led to increased chick survival.     

Abundances of surgeonfishes on patch-reefs in Caribbean Panamá: due to settlement,or post-settlement events?

I examined whether patterns of settlement of planktonic juveniles determined the size and composition of adult populations of three Caribbean surgeonfishes (A canthurus bahianus, A. chirurgus and A. coeruleus) on six isolated patch-reefs in Panamá. I monitored settlement monthly on six reefs (0.14 to 0.67 ha) for 8 yr (1978–1986), and censused adults on those reefs annually. Year-to-year fluctuations in the absolute and relative abundances of settlers arriving on each reef were not extreme. Densities of adults in 1986 were not correlated with the densities of settlers that arrived between 1979 and 1984. Survival of settlers, although apparently unrelated to the density of adults, was negatively related to the density of settlers per unit of adult habitat (but not per unit of settlement habitat) for each species. Changes in adult populations of each species between 1978 and 1986 were not correlated with the abundance of settlers relative to the initial number of adults. The relative abundances of adults of each species in 1986 did not reflect the relative abundances of settlers (1979–1984); these abundances were due to both spatial variation and interspecific variation in survival. In some cases, relocation of adults between isolated patch-reefs (as much as 100 m from other reefs) substantially altered the size and composition of surgeonfish populations. Thus, either post-settlement mortality or relocation overrode effects of settlement patterns in determining adult abundances on the six reefs. Relocation is important in organizing reef-fish communities and must be further examined, particularly in large habitat mosaics.     

Trophic relations of the cephalopod Martialia hyadesi (Teuthoidea: Ommastrephidae) at the Antarctic Polar Front,Scotia Sea

Samples of the squid Martialia hyadesi were collected aboard two Japanese squid-jigging vessels carrying out commercial fishing trials at the Antarctic Polar Frontal Zone, north Scotia Sea, in February 1989. The dissected stomachs of 61 specimens were classified according to fullness and the contents were examined visually. Identifiable food items included fish sagittal otoliths, crustacean eyes, the lappets on euphausiid first antennule segments and cephalopod sucker rings. The most frequent items in the squid's diet were the myctophid fishes Krefftichthys anderssoni and Electrona carlsbergi, the euphausiid Euphausia superba and a hyperiid amphipod, probably Themisto gaudichaudi. A small proportion of the sample had been feeding cannibalistically. Total lengths of the fish prey were estimated from sagittal otolith size using published relationships. All fish were relatively small; 7 to 35% of squid mantle-length. However, it is possible that some heads of larger fish are discarded by the squid and so are not represented by otoliths in the stomach contents. Over the size range of squid in the sample there was no relationship between size of fish prey and size of squid. Similarly, when the squid sample was divided into groups according to prey categories: crustaceans, crustaceans+fish, fish, cephalopod, there was no evidence that dietary preference was related to squid size. The prevalence of copepod-feeding myctophids in the diet of this squid, which is itself a major prey item of some higher predators in the Scotia Sea, suggests that a previously unrecognised food chain: copepod-myctophid-M. hyadesi-higher predator, may be an important component of the Antarctic oceanic ecosystem.     

Replacement males in the western bluebird: opportunity for paternity, chick-feeding rules, and fitness consequences of male parental care

In western bluebirds (Sialia mexicana), most pairs remain together for life and share equally in post-hatching parental care. We removed resident males of socially monogamous pairs during laying and after clutch completion to examine chick-feeding rules used by replacement males and current and future fitness consequences of paternal care. Replacement males were not infanticidal and 7 (47%) fed nestlings. Feeding replacement males and the females they joined fed at rates similar to controls. Females without a feeding replacement male compensated by feeding more themselves so that overall feeding rates were not compromised, but they reduced their brooding time. Unlike assisted females, unassisted females exhibited reduced nesting success and their 14-day-old chicks weighed less than controls. Field metabolic rates of unassisted females were 17% higher than those of control females, but the difference was not statistically significant. Older females were better able to raise young without the male's help than were yearling females. Female condition was not affected by male provisioning and unassisted females were as likely to survive to the next breeding season as assisted females. We found no future benefits of provisioning by replacement males; those that fed were no more likely to breed with the female on her subsequent attempt than were males that did not feed, and subsequent clutch sizes were not reduced for females rearing young without the male's help. These experiments suggest that male parental care increases nesting success in western bluebirds and that replacement males use an all-or-none rule to determine whether or not to feed chicks: if they are present during the fertile period they feed at typical rates; if they are not, they usually do not feed at all. Because chick-feeding by males is tied to opportunity for paternity, influences success in the current nest, and does not affect the male's future breeding success, it appears to be parental rather than mating effort. Received: 8 May 1998 / Accepted after revision: 23 July 1998     

Fish and squid in the diet of king penguin chicks,Aptenodytes patagonicus,during winter at sub-antarctic Crozet Islands

The diet of king penguins, Aptenodytes patagonicus, rearing chicks was studied during three consecutive austral winters (1990, 1991 and 1992) at Crozet Islands. The mean stomach content mass of the 47 samples was 503 g. Percentages of wet and reconstituted masses showed that both fishes (66 and 36%, respectively) and squid (34 and 64%) are important components of the winter diet. Juveniles of the demersal onychoteuthid squid Moroteuthis ingens form the bulk of the cephalopod diet, and this was the main prey by reconstituted mass (57%). Myctophid fish (lantern-fishes) accounted for most of the fish diet, constituting together 32% by mass. The three main species of myctophids eaten in summer by king penguins were either very rare in winter (Electrona carlsbergi) or accounted for a smaller proportion of the diet (Krefftichthys anderssoni = 1.5% by mass and Protomyctophum tenisoni = 4.6%). Five other myctophids, which are rarely consumed in summer, contributed 24% of the diet by mass in winter (Gymnoscopelus piabilis = 18.1%, Lampichthys procerus = 2.4%, G. nicholsi = 1.3%, and Metelectrona ventralis and Electrona subaspera = 1.0%). The greater diversity of prey in winter suggests a more opportunistic feeding behaviour at a time probably marked by a change in prey availability. Both the known ecology of the fish and squid prey and the barely digested state of some items suggest that in winter breeding adults forage in the outer shelf, upper slope and oceanic areas in the close vicinity of the Crozet Islands to feed their chicks. Finally, using king penguins as biological samplers, the present work provides novel data on the previously unstudied mesopelagic/epibenthic marine community in waters surrounding the Crozet Islands. Seventeen myctophid fish have been identified to species level. These include several poorly known species in the southern Indian Ocean. The occurrence of small, nearly intact, cephalopods in the diet of king penguins suggests that spawning grounds of four squid species may be located near the Crozet Archipelago.     

Positive correlation between helpers at nest and nestling immune response in a cooperative breeding bird

In cooperatively breeding species, helpers may contribute to the success of the brood by increasing the number of independent offspring, but also, they may affect offspring condition and, hence, their survival and recruitment into the breeding population. This second type of benefits is rarely included in theoretical models or assessed in field studies. Immune response is a good proxy of individual quality and fitness, and there is good evidence that the performance of the immune system of chicks during the nestling phase is related to their chances of survival and future reproduction. However, no study has so far explored whether helpers at the nest might contribute to enhance immune functioning of nestlings in species with a cooperative breeding system. Here we investigate this issue in the azure-winged magpie (Cyanopica cyanus) and found for the first time a positive correlation between the number of helpers at nest and the cell-mediated immune response of nestlings. This effect was not explained by a general improvement of body condition of chicks because it was independent of individual variation in body mass or tarsus length. Our results suggest that helping can have subtle effects on the quality of offspring that may influence their survival and future reproduction.     

The effects of helpers in cooperatively breeding purple gallinules

Summary The effects of helpers in a population of cooperatively breeding purple gallinules (Porphyrula martinica) were examined. All young birds past the age of 2 months helped feed and protect subsequent broods of chicks and participated in territorial defense. Most helpers remained on their natal territories for approximately 1 year. The number of helpers varied both among and within breeding groups. Clutch production and chick survival were related positively to the number of helpers in the group. The increase in chick survival was independent of several measures of territory quality. Helpers possibly aided chick survival by provicing extra food for the chicks and decreasing predation risk. Helpers were necessary in order for a breeding pair to keep a territory long enough to produce more than one clutch of eggs. A change in the number of helpers (increase or decrease) often was followed by a similar change in territory size. These results suggest that purple gallnule helpers can increase the reproductive success of the breeding group and may be vital for the continued maintenance of a breeding territory.     

Effects of paternal care on reproductive success in the polygynous spotless starling Sturnus unicolor

For males of socially polygynous avian species like the spotless starling, there may exist a trade-off between investing in paternal care and controlling several nests. To determine how the intensity of paternal care affects reproductive success per brood sired or expressed as the total number of young raised in all nests controlled by the same male, it is necessary to manipulate paternal care. Testosterone (T) has been shown to depress the tendency for males to care for their young, and induces them to acquire more mates. The effects of paternal care on reproductive success were studied by treating certain male starlings with exogenous T and others with the antiandrogen cyproterone acetate (CA), and comparing the parental behavior of T- and CA-males throughout the breeding season with that of controls. CA-males fed their chicks more during the first week after hatching than T-males, with controls feeding at intermediate rates, both on a per nest basis and as total effort for all nests controlled by the same male. Paternal feeding rates during the first week of chick life had a significant positive effect on the number of fledged young. The hormone treatment significantly affected the number of chicks raised per nest, CA-males having a higher breeding success per nest than T-males, and controls showing intermediate levels of success. There was no significant effect of treatment on total reproductive success attained by males throughout the season. In the polygonous spotless starling, the intensity of paternal care of young affects reproductive success per nest positively but not on a seasonal basis. Received: 6 February 1999 / Received in revised form: 30 June 1999 / Accepted: 11 July 1999     

Stable isotopes identify age- and sex-specific dietary partitioning and foraging habitat segregation in southern giant petrels breeding in Antarctica and southern Patagonia

We examined the isotopic signatures (δ¹³C, δ¹⁵N) of adult body feathers from southern giant petrels Macronectes giganteus collected at two breeding colonies in Antarctica (Potter Peninsula and Cape Geddes) and one in southern Patagonia (Observatorio Island), as well as in whole blood collected from adults of both sexes at each Antarctic colonies and from chicks at Potter Peninsula. As body feather moult is a continuous process in giant petrels, feathers provide an integrated annual signal of an adult’s diets and foraging habitats. In contrast, the stable isotope values of adult and chick blood are reflective of their diets during the breeding season. We found that sex-specific dietary segregation in adults breeding in Antarctica was notable during the breeding season (blood samples) but absent when examined across the entire year (feather samples). In addition, blood stable isotope values differed between chicks and adults, indicating that adults provision their offspring with a relatively higher amount of penguin and seal prey that what they consume themselves. This finding confirms previous work that suggests that chicks are preferentially fed with prey of presumably higher nutritional value such as carrion. Finally, based on isotopic differences between major oceanographic zones in the Southern Ocean, our data indicate population-specific differences in foraging distribution, with Antarctic populations move seasonally between Antarctic and subantarctic zones, while Patagonian populations likely forage in subtropical waters and in continental shelf habitats year-round.     

Cephalopod remains from the stomach of a sperm whale stranded off Patagonia

The stomach of a male sperm whale (Physeter macrocephalus) stranded near Comodoro Rivadavia, Argentina, on 21 March 1986, contained 247 cephalopod beaks (128 upper and 119 lower). The lower beaks were identified, their rostral lengths were measured and used to estimate the mantle lengths and wet weight of the representatives of each cephalopod taxon. Fifteen species from nine families were represented. The most numerous species, in order of decreasing abundance, were the large cranchiidMesonychoteuthis hamiltoni (32.8% of total numbers),Mastigoteuthis sp. A (16.0%), andKondakovia longimana andHistioteuthis atlantica (both 13.5%). Almost 85% of the estimated total weight was contributed by two species,Mesonychoteuthis hamiltoni (61.6%) andK. longimana (23.1%). The presence of beaks fromK. longimana, Alluroteuthis antarcticus andM. hamiltoni, which are all thought to be restricted to the Antarctic, suggests that the whale had migrated North prior to stranding.     

Benefits of large broods by higher chick survival and better territories in a precocial shorebird

When reproductive success is constant in one breeding phase, different tactics that increase variation in reproductive success among individuals may evolve in other phases. For instance, in shorebirds, which usually have a limited clutch size of four eggs, variation in reproductive tactics among individuals is expected either before egg-laying (e.g. diverse mating systems) or after hatching of the young (e.g. diverse parental care). In this paper, I studied the pied avocet (Recurvirostra avosetta), a shorebird with a modal clutch size of four eggs, to test whether post-hatch chick adoption as an alternative tactic can be linked to increased variation in annual reproductive success. When predation was high, naturally adopting pairs produced more filial fledglings than did pairs not adopting chicks and not losing chicks to adoption. The number of filial fledglings increased with the number of adopted young, possibly through diluting the chances of predation on filial young. Experimental chick addition did not lead to more fledged young due to low brood integrity as shown by the frequent loss of chicks from some experimental broods. When predation was low, larger broods occupied feeding territories with higher prey abundance than smaller broods, possibly due to their dominance over smaller ones. Pairs that lost chicks to adoption (donors) fledged as many filial young in their broods as did non-adopters/non-donors, whereas the total number of donors’ filial fledglings, including those raised in adopting broods, approached that of adopters. These findings show, for the first time, that post-hatch alternative reproductive tactics can lead to variation in annual reproductive success and to higher success for some pairs even in species where past adaptations limit variation in reproductive success in a certain phase of reproduction.     

Diet and nest attendance of incubating and chick-rearing northern fulmars (Fulmarus glacialis) in Shetland

We examined diets and nest attendance patterns of northern fulmars (Fulmarus glacialis) on Foula and Unst, Shetland, UK, during the breeding seasons of 1998 and 1999. Mean foraging trip duration, derived from nest attendance patterns, differed between incubation (32.5 h) and chick-rearing (early stage: 11.2 h, mid-stage: 20.4 h). It was influenced by chick age and obviously also by prey availability. The numbers of fulmars attending the colonies were influenced by wind direction, wind speed, time of day and state of reproduction. Diet samples were collected by regurgitations of adult and young fulmars. Diet was analysed by identifying prey items on the basis of otoliths, vertebrae, premaxillae (fish), cephalopod beaks and fragments of arthropod exoskeletons. Adult fulmars and chicks fed upon a wide range of prey types. Gadoid fish, including Norway pout (Trisopterus esmarkii), were the most common prey in the diet. Fish offal was found in 32% of regurgitates, clupeids in 15%. In contrast, the proportions of sandeels were very low (1%). This is different from previous studies where sandeels formed a large part of fulmar diet at Shetland in summer. Differences in food composition of incubating and chick-rearing fulmars were found for fish, but not for offal, crustaceans or squid. The percentage of non-discard fish (Clupeidae, Ammodytidae, Isospondylae) was significantly different between regurgitates from incubating and chick-rearing fulmars. Samples from chick-rearing adults contained non-discard items more frequently than regurgitates from incubating birds. The results strongly indicate that fulmars select to feed their chicks on energy-rich clupeids.     

Grazing as a measure against grass encroachment in Dutch dry dune grassland: effects on vegetation and soil

A grazing experiment was started in 1984 and 1989 respectively, in two parts of a dune grassland in the nature reserve ‘Zwanenwater’, North Holland; a third part with similar geology and topography was used as a control area and not grazed. An evaluation of the effects of grazing on vegetation patterns, species composition, vegetation structure and humus form was made with the help of vegetation maps from 1986 and 1992 as well as field surveys. Dense tall-grass communities dominated byAmmophila arenaria increased over the period 1986–1992 in the grazed areas, and especially in the non-grazed area (increase in area to 20%, 22% and 50%, respectively). Open communities decreased in the grazed areas, but are still prevalent, while in the ungrazed area they virtually disappeared, with the result that the present percentage areas are 53%, 38% and 17%. Field survey data were classified by TWINSPAN producing four vegetation types. These occur more or less equally in grazed and ungrazed areas, albeit with different percentage areas: (1) open vegetation dominated byCorynephorus canescens; (2) open vegetation characteized, byKoeleria macrantha; (3) heathland dominated byEmpetrum nigrum; and (4) tall-grass communities dominated byAmmophila arenaria. Within a vegetation type, species composition was only marginally affected by grazing regime. Within the open communities the number of species, vegetation height, vegetation cover and soil organic horizons were not affected by grazing. In the tall-grass communities the number of species was significantly larger and the height of the vegetation significantly lower in the area grazed since 1984. In the heathland community the number of species and cover of the moss layer were significantly higher in the 1984 area and ectorganic and endorganic horizons significantly thicker in the ungrazed area. It is suggested that these effects are the result of an increased availability of light, but possibly also of a decreased stock of organic matter and nutrients, due to a decreased input of litter and accelerated rates of decomposition.     

Distribution patterns,abundance and population dynamics of the euphausiidsNyctiphanes capensis andEuphausia hanseni in the northern Benguela upwelling system

Distribution patterns, population structure and biomass of the euphausiidsNyctiphanes capensis andEuphausia hanseni were examined off the coast of Namibia, southwest Africa, in relation to temperature, depth and season, from data collected on nine surveys from September 1982 to March 1984. High densities ofN. capensis were found in the shallow coastal waters (<200 m), with the biomass of adults ranging from 675 to 5 706 mg dry wt m^–2. For adultE. hanseni, the biomass was an order of magnitude lower, ranging from 65 to 505 mg dry wt m^–2, with most specimens occurring over the shelf break at depths of 200 to 1000 m. These distribution patterns remained relatively constant throughout the year, despite seasonal differences in upwelling events. Both species displayed continuous breeding, with 43 to 82% of the adult femaleE. hanseni being fertilized, while a much lower proportion ofN. capensis females were reproductively active (0.5 to 26%). Different breeding strategies were adopted by these two euphausiid species, withE. hanseni producing frequent broods (14.8 broods in 6 mo) consisting of relatively large eggs which are released into the sea, andN. capensis exhibiting a lower frequency of spawning, with broods consisting of large numbers of relatively small eggs, protected by a brood pouch. These strategies enable both species to maintain high densities throughout the year in a fluctuating physical environment. Growth rate estimated from size-frequency distributions were 0.003 to 0.063 mm d^–1 forN. capensis and 0.077 to 0.083 mm d^–1 forE. hanseni, suggesting an adult lifespan of approximately 6 mo for both species. Maximum sizes were attained in September, withN. capensis reaching a total length of 21 mm (in contrast to all previous studies onN. capensis, where the maximum size recorded was only 13 mm total length) andE. hanseni a total length of 33 mm.     

Assuming a 50% cell occupancy of the lorica overestimates tintinnine ciliate biomass

To estimate biomass and production, the cell volume of tintinnine ciliates has been assumed to be 50% of lorica volume. This assumption was tested using a high-precision digitizing system which measured individuals of 17 tintinnine species in samples taken in two annual studies of biomass and production in both tropical (Kingston, Jamaica in 1986) and temperate (Passamaquoddy Bay, Canada in 1983–1984) regions. Using this assumption, percent occupancies and hence, biovolume estimates, were overestimated by as much as 317% for tropical species and 178% for temperate species. Daily production estimates of the tropical tintinnine assemblage were overestimated by as much as 147%. In studies attempting to estimate biomass and production accurately, it is concluded that cell volume of tintinnines should be measured directly rather than assumed to be 50% of lorica volume.This research was financed by an NSERC Canada operating grant awarded to Dr D. H. Lynn