Diet and prey consumption of Antarctic petrels Thalassoica antarctica at Svarthamaren, Dronning Maud Land, and at sea outside the colony

The diet of the Antarctic petrel Thalassoica antarctica was studied during two seasons at Svarthamaren, an inland colony in Dronning Maud Land, Antarctica, and in the pack ice off the coast of Svarthamaren. The most important food (wet mass) at Svarthamaren was crustaceans (67%), fish (29%) and squid (5%); however, individuals collected in the pack ice took mostly fish (87%). The prey composition and lengths of prey are comparable to what has been documented in other studies on this species. Estimates of food consumption by birds breeding at Svarthamaren (ca. 250,000 pairs) suggest that approximately 6500 tonnes of crustaceans, 2800 tonnes of fish and 435 tonnes of squid are consumed during the breeding season. The annual consumptions of these birds are estimated to be 34,100 tonnes of crustaceans, 14,700 tonnes of fish, and 2300 tonnes of squid. Satellite telemetry data indicate that Antarctic petrels from Svarthamaren may fly more than 3000 km during one foraging trip, and thus may cover a huge ocean area to obtain their prey. Received: 1 September 1997 / Accepted: 3 February 1998     

DIETS OF FIN, SEI, AND SPERM WHALES IN BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS, 1963–1967

Diets of fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), and sperm whales ( Physeter macrocephalus ) were estimated from the stomach contents of individuals killed along the British Columbia coast from 1963 to 1967. The dominant prey types of fin whales were euphausiids, with minor contributions from copepods and fish. Sei whale stomachs contained primarily copepods in three years, whereas euphausiids or a variety of fish dominated the diet in the other two years. Sperm whales consumed primarily North Pacific giant squid ( Moroteuthis robusta ), but secondary prey differed between males and females. Female sperm whales frequently consumed ragfish ( Icosteus spp.) and other fish, whereas the male diet also contained rockfish ( Sebastes spp.). The high abundance of euphausiids along the British Columbia coast likely contributed to the presence of a summer resident population of fin whales. The high abundance of large copepods farther north probably influenced the migration of sei whales through the offshore waters of British Columbia. Sperm whale stomach contents differed by sex reflecting location and possibly breeding behaviors.     

Feeding ecology and interannual variations in diet of southern bluefin tuna, Thunnus maccoyii, in relation to coastal and oceanic waters off eastern Tasmania, Australia

The diets of 1219 southern bluefin tuna, Thunnus maccoyii, from inshore (shelf) and offshore (oceanic) waters off eastern Tasmania were examined between 1992 and 1994. Immature fish (< 155 cm fork length) made up 88% of those examined. In all, 92 prey taxa were identified. Inshore, the main prey were fish (Trachurus declivis and Emmelichthys nitidus) and juvenile squid (Nototodarus gouldi). Offshore, the diversity was greater, reflecting the diversity of micronekton in these waters. Interestingly, macrozooplankton prey (e.g. Phronima sedentaria) were prevalent in tuna > 150 cm. The offshore tuna, when in subantarctic waters, ate relatively more squid than when in the East Australia Current. In the latter, fish and crustacea were more important, although there were variations between years. No relationship was found between either prey type or size with size of tuna. Feeding was significantly higher in the morning than at other times of the day. The mean weight of prey was significantly higher in inshore-caught tuna than in those caught offshore. We estimated that the mean daily ration of southern bluefin tuna off eastern Tasmania was 0.97% of wet body weight day⁻¹. However, the daily ration of inshore-caught tuna was ∼ 3 times higher (2.7%) than for tuna caught offshore (0.8%) indicating that feeding conditions on the shelf were better than those offshore. Our results indicate that the inshore waters of eastern Tasmania are an important feeding area for, at least, immature southern bluefin tuna. This revised version was published online in July 2006 with corrections to the Cover Date.     

Experimental study on the effect of diet on fatty acid and stable isotope profiles of the squid Lolliguncula brevis

Fatty acid and stable isotope analyses have previously been used to investigate foraging patterns of fish, birds, marine mammals and most recently cephalopod species. To evaluate the application of these methods for dietary studies in squid, it is important to understand the degree to which fatty acid and stable isotope signatures of prey species are reflected in the squids' tissue. Four groups of Lolliguncula brevis were fed on prey species with distinctly different fatty acid and stable isotope profiles over 30 consecutive days. One group of squid were fed fish for fifteen days, followed by crustaceans for a further fifteen days. A second and third group were fed exclusively on fish or crustaceans for thirty days. And a fourth group was fed on a mixture of fish and crustaceans for thirty days. Analysis of squid tissue showed that, after 10 days of feeding, fatty acid profiles of squid tended to reflect those of their prey. Squid that fed on a single prey type, i.e. fish or crustacean, showed only minor modifications in fatty acid proportions after the initial change and fatty acid profiles were clearly distinguishable between the two feeding groups. Shifts in fatty acid proportions towards respective prey profiles could clearly be observed in squid the diet of which was swapped after 15 days. Clear differences could also be seen in fatty acid profiles of squid feeding on a mixed diet with trends towards either fish or crustacean fatty acid signatures. Stable isotope signatures of squid tissues clearly distinguished between animals feeding on different diets and supported findings from fatty acid analysis, thus indicating both methods to be viable tools in feeding studies on squid species.     

The feeding habits of six pelagic and predatory teleosts in eastern Cape coastal waters (South Africa)

A four-year study of the feeding habits of six pelagic teleost species in eastern Cape coastal waters of South Africa is analysed. These predators are Lichia amia, Pomatomus sallatrix, Seriola lalandi, Atractoscion aequidens, Katsuwonus pelamis and Thunnus albacares. All species occur in inshore coastal water (< 1 -8 km from shore) for at least part of the year. The tunas were also sampled from the edge of the continental shelf (40–50 km from shore). The use of squid 'beaks' and fish otoliths allowed accurate prey identification and size estimates. Variations in prey taken are attributed to differences in locality, predator species and size. Three pelagic fish species are important to all the predators: Sardinops ocettata, Etrumeus teres and Engraulis capensis. The inshore squid, Loligo reynaudi, is another major prey item. Offshore tuna samples revealed a different prey spectrum, viz. oceanic squid and saury, Scomberesox saurus, being taken frequently.     

Relationships between bird morphology and prey selection in two sympatric Great Cormorant Phalacrocorax carbo subspecies during winter

Variation in bird morphology (notably sex size dimorphism) has been suggested to contribute to differences in food use between individuals. We explore the hypothesis of food partitioning (diet overlap and prey size selection) in two sympatric subspecies of the Great Cormorant Phalacrocorax carbo with respect to bird morphology (subspecies and sex) in inland French waters. These areas represent a recent and increasing contact zone used as a common non-territorial winter feeding area by the continental P. c. sinensis and the marine P. c. carbo subspecies. A high dietary overlap between subspecies and sexes was found at the major sites studied. Prey size selection was found to be site-specific and generally related to a gradient of structural size and body mass (male P. c. carbo  > male P. c. sinensis  > female P. c. carbo  > female P. c. sinensis ). With respect to bird morphology, differences in prey size consumed by Great Cormorants were more pronounced between sexes in freshwater habitats than between subspecies. This was reinforced by the fact that P. c. carbo birds entering inland areas were smaller than birds on the coast. These results also suggest that bird morphology is an important determinant of dietary differences. Our study demonstrated that P. c. carbo is an efficient generalist forager in continental areas, and food partitioning in type and size of prey between the two subspecies is low.     

Variation in the diet of Arctic Cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) in the Pacific Arctic and Bering Sea

Arctic cod, Boreogadus saida (Lepechin, 1774), is a nodal species in Arctic marine foodwebs as an important prey of many birds, marine mammals, and other fishes, as well as an abundant predator of zooplankton and epibenthic fauna. We examined the summer diet of Arctic cod across a latitudinal gradient extending from the southern limit of their distribution in the eastern Bering Sea to the northern margins of the eastern Chukchi Sea (ECS) continental shelf. Specimens were collected from demersal and pelagic trawls conducted between 1999 and 2012, and across a range of predator sizes (3–26 cm). Arctic cod diets vary with body size and between regions within the study area, and appear to vary between years in the eastern Bering Sea, indicating opportunistic feeding habits. Constrained Analysis of Principal Coordinates was conducted on ECS demersal samples and revealed consumption of fish and decapod crustacea were positively correlated with Arctic cod length while consumption of euphausiids and copepods had the opposite relationship. The demersal Arctic cod diet in the northern latitudes of the ECS was dominated by copepod consumption (47% by weight, %W), but copepods were less important (12–26%W) in the central and southern latitudes of the ECS and in the northern and eastern Bering Sea—areas where diets were more varied in their composition. High levels of variation in the diet of Arctic cod highlights the need to monitor Arctic cod diets to identify consistently dominant prey types and potential future changes to trophic relationships related to climate change or increasing anthropogenic activity.     

Niche dynamics of sympatric non-breeding shearwaters under varying prey availability

Variation in prey availability can cause changes in species interactions among marine predators. Foraging theory predicts that niche breadth will expand when resources become limited, possibly leading to higher niche overlap among sympatric species; however, a species’ niche can become constrained by interactions with other similar species, resulting in an inability to shift niche breadth or position. In coastal Newfoundland, Capelin Mallotus villosus is the main forage fish species and its availability (i.e. biomass) during the summer has varied considerably following a population collapse in the 1990s. During the summer, non-breeding Great and Sooty Shearwaters Ardenna gravis and A. grisea migrate and aggregate at annually persistent Capelin spawning sites. We used stable isotope ratios (δ¹³C, δ¹⁵N) of blood components (plasma, red blood cells) to investigate variation in isotopic niche breadth (95% prediction ellipse areas) and overlap of the two shearwater species during 2014, 2015 and 2016. Capelin availability varied among years, illustrated by lower peak biomass in 2015 (0.126 g/m²) and 2016 (0.027 g/m²) relative to 2014 (0.254 g/m²). The isotopic niche breadth (plasma) of both shearwater species expanded similarly from 2014 (0.65–0.66‰²) to 2015 (2.22–2.57‰²) and 2016 (1.15–1.42‰²), suggesting the incorporation of alternative prey types into their diet during years of lower prey availability. Isotopic niche overlap between Great and Sooty Shearwaters remained high across years (44–63%), however, providing little evidence for dietary niche partitioning during years of lower prey availability. Findings suggest that both shearwater species are flexible foragers and can modify their diet during the non-breeding season to accommodate fluctuations in prey availability.     

An effective method for trapping scavenging seabirds at sea

ABSTRACT Most studies of seabirds that involve trapping and marking birds are carried out at breeding colonies. This bias toward the breeding period and colony‐based research is partially caused by difficulties in capturing birds at sea. From 2005 to 2007, we used a cast net thrown by hand from a fishing boat to capture albatrosses and petrels at sea in the southwestern Atlantic Ocean. About 500 birds of 13 species were captured, ranging in size from the 30‐g Wilson's Storm‐petrel (Oceanites oceanicus) to the 10‐kg Wandering Albatross (Diomedea exulans). Cast nets are potentially useful for capturing any seabird that can be attracted close to fishing vessels by bait, such as sardines, squid, offal, or shark liver, thrown into the water. Our method was most effective for capturing bold species, such as Cape (Daption capense), Spectacled (Procellaria conspicillata), and White‐chinned (Procellaria aequinoctialis) petrels and Great Shearwaters (Puffinus gravis), but was not effective for capturing shy species, such as Cory's (Calonectris diomedea), Cape Verde (Calonectris edwardsii) and Manx (Puffinus puffinus) shearwaters, species that rarely sit on the water, such as Wilson's Storm‐petrels, Thin‐billed (Pachyptila belcheri) and Antarctic (Pachyptila desolata) prions and Atlantic Petrels (Pterodroma incerta), and species with excellent diving abilities, such as Sooty Shearwaters (Puffinus griseus). For many species of seabirds, cast nets would likely be more efficient for capturing large numbers of birds than other methods, such as hoop nets.     

OBSERVATIONS ON THE NESTING SITES AND NESTING BEHAVIOUR OF THE KITTLITZ'S SANDPLOVER CHARADRIUS PECUARIUS

Jackson, S. 1984. Predation by Pied Kingfishers and Whitebreasted Cormorants on fish in the Kosi estuary system. Ostrich 55:113-132.

Identification of otoliths from the regurgitated pellets of Pied Kingfishers Ceryle rudis and Whitebreasted Cormorants Phalacrocorax carbo from the Kosi estuary system provides information on the relative proportions of fish species in the diets of the birds. This information can be related to the feeding habits, distribution and abundance of their prey. It is also an indication of the feeding range of the birds. There is little overlap between both the size classes and the species of fish taken by the two predators. This is because of the difference in size and fishing techniques of C. rudis and P. carbo, and of differences in their feeding ranges. Competition for food between the two populations of birds studied is minimized by these differences.     

Comparison of the feeding apparatus in the muttonbird, Puffinus tenuirostris (Temminck) and the fairy prion, Pachyptila turtur (Kuhl) in relation to the capture of the krill, Nyctiphanes australis sars

The short-tailed shearwater (muttonbird) Puffinus tenuirostris (Temminck) and the fairy prion Pachyptila turtur (Kuhl) are procellariiform birds which feed extensively on the euphausiid Nyctiphanes australis Sars in south eastern Tasmania. Data from a hydrodynamic study of the heads of the two birds were used to develop a simple qualitative model of food capture. This model predicts that as the bird moves through the water the open gape, aided by the bill architecture, produces and maintains a low pressure volume into which the prey is sucked. Once inside the bill euphausiids are prevented from escaping by filtering systems which differ in the two birds. In the muttonbird, the filtering mesh consists of retroverted papillae of the upper palate which overlap with opposing papillae on the tongue. In the fairy prion a combination of papillae in the oral cavity and a series of closely-set lamellae on the margins of the upper palate serve to retain the krill.     

Food habits of the sandpaper skate, Bathyraja kincaidii (Garman, 1908) off central California: seasonal variation in diet linked to oceanographic conditions

The stomachs of 130 sandpaper skates, Bathyraja kincaidii (Garman, 1908), were sampled from off central California to determine their diet composition. The overall diet was dominated by euphausiids, but shrimps, polychaetes and squids were also important secondary prey. A three-factor MANOVA demonstrated significant differences in the diet by sex, maturity status and oceanographic season using numeric and gravimetric measures of importance for the major prey categories. These three main factors explained more variation in diet than interactions between the factors, and season explained the most variance overall. A detailed analysis of the seasonal variation among the prey categories indicated that abundance changes in the most important prey, euphausiids, were coupled with seasonal changes in the importance of other prey. When upwelling occurred and productivity was great (Upwelling and Oceanic seasons), euphausiids were likely highly abundant in the study area and were the most important prey for B. kincaidii. As productivity declined (Davidson Current season), euphausiids appeared to decrease in abundance and B. kincaidii switched to secondary prey. At that time, gammarid amphipods and shrimps became the most important prey items and polychaetes, mysids and euphausiids were secondary.     

Sexual differences in the diet of great cormorants <Emphasis Type="Italic">Phalacrocorax carbo sinensis</Emphasis> wintering in Greece

Sexual differences in the diet of the great cormorant, Phalacrocorax carbo sinensis, were studied in four Greek wintering areas, the Amvrakikos Gulf, the Axios and Evros Deltas and the Messolonghi Lagoon, through the analysis of stomach contents. Great cormorants are birds sexually dimorphic in size, with males being generally larger than females. Although similar prey species were found in the stomachs of both sexes in all the studied areas, significant differences were observed with respect to the proportion of species taken. Male birds ate higher proportions of large fish species such as grey mullets, European sea bass, Dicentrarchus labrax, and Prussian carp, Carassius gibelio, while female birds took higher proportions of smaller species such as big-scale sand smelt, Atherina boyeri, and black goby, Gobius niger. As a consequence, male great cormorants were found to feed on significantly larger prey than did females by means of fish standard length and body mass. There was no significant difference between the sexes in the mass of food found in stomachs.     

SUMMER DISTRIBUTION OF PELAGIC BIRDS OFF THE COAST OF ARGENTINA

Eighteen species of sea–birds were observed from CSS 'Hudson' in the South Atlantic between the Rio de la Plata and Tierra del Fuego in late January and late February 1970. Major feeding areas were noted off Mar del Plata, the Valdes Peninsula and Tierra del Fuego, Argentina, the first two in rich modified or unmodified water of the subantarctic Falkland Current, associated with upwelling. The southernmost area had most birds, the northernmost fewest. A temperate–latitude assemblage including Yellow–nosed Albatross and Cory's Shearwater occurred off Mar del Plata in warmed water of subantarctic (Falkland Current) origin. South of 39°S these two species were replaced by Black–browed Albatross and Great Shearwater in cold offshore water of the Falkland Current. Off the Valdes Peninsula Black–browed Albatross, Great and Sooty Shearwaters, White–chinned Petrels and family groups of Magellanic Penguins occurred near the edge of highly–productive water of the Falkland Current. The largest feeding area, east of Tierra del Fuego, had Magellanic and Rockhopper Penguins, Great Shearwaters, King Cormorants, South American Terns and jaegers, but no oceanographic information is available. At 50°S prions and diving petrels appeared for the first time, marking a possible oceanographic discontinuity or proximity to colonies on the Falkland Islands. Several common species of the South Atlantic were not seen on the transect including Cape Pigeon, Southern Fulmar, Soft–plumaged Petrel, Schlegel's Petrel and Pediunker. Kelp Gull, South American Tern, Manx Shearwater and jaegers were rare or absent far from land. New southern limits for the western South Atlantic Ocean were established for Cory's and Manx Shearwaters, and the Great Shearwater was observed for the first time in Chilean waters.     

PREY SELECTION AND TEMPORAL VARIATION IN THE DIET OF SUBANTARCTIC SKUAS AT INACCESSIBLE ISLAND,TRISTAN DA CUNHA

Ryan, P.G., & Moloney, C.L. 1991. Prey selection and temporal variation in the diet of Subantarctic Skuas at Inaccessible Island, Tristan da Cunha. Ostrich 62:52-58.

More than 2 500 prey items of Subantarctic Skuas Cuthuructa anturcricu were identified from regurgitated pellets collected at a roost of non-breeding skuas during summer 1989–90 at West Point, Inaccessible'Is-land. Most prey items (96,5%) were birds, primarily burrow-nesting procellariiforms. Sixteen bird species were recorfed in the diet, but five secies accounted for 94,6% of identifiable prey remains: Whitebellied Storm Petrel Fieettu grullariu (53,5%), Whitefaced Storm Petrel Pelugodrorna marina 15,1%), Broad-billed Prion Pacaptilu vittutu (14,0%), Great Shearwater Puffinus gruvis (7,3%), and Iommon Diving Petrel Pelecunoids urinutrix (4,7%). Petrel chicks were important in the diet, particularly Great Shearwaters. The main non-avian prey were fish and goose barnacles Lepas sp., but their importance may have been underestimated. The remains of a rat Rattus sp. presumably were derived from the neighbouring island of Tristan da Cunha; rats are not known to occur on Inaccessible Island. Prey size affected seabird availability to skuas, and dietary composition vaned throughout the summer in relation to seabird breeding cycles. Both these factors reduce the value of skua diet as an indicator of the relative abundances of burrow-nesting petrels. There was no correlation between skua diet and estimates of breeding bird abundance, but this may be a consequence of studying non-breeding skuas. Subantarctic Skua diet in 1989–90 was similar to that recorded in previous years, with consistent seasonal trends between years.     

The food of Atlantic salmon, Salmo salar L., caught by long–line in northern Norwegian waters

A total of 1145 stomachs from Atlantic salmon caught over the shelf off Helgeland/Trøndelag, and in the oceanic waters off Andenes, northern Norway during late winter–spring, 1969–1972 were examined. Food was found in 52.9% of the stomachs examined. The most important food items found in fish caught in the Helgeland/Trøndelag area were euphausids and hyperid amphipods while the myctophid Benthosema glaciale , the squid Gonatus fabricil and euphausids were found most frequently in the salmon caught off Andenes. Most salmon had preyed upon only one species, and few stomachs contained three or more prey species. The type of food did not appear to be related to the length of the fish. It is suggested that some mesopelagic feeding occurred.     

A comparative study of diet in three sympatric populations of<Emphasis Type="Italic"> Patagonotothen</Emphasis> species (Pisces: Nototheniidae)

The diet of three Patagonotothen species was investigated from the bycatch of near-bottom trawls for squid on the southern Falkland Island shelf. Patagonotothen ramsayi had a moderately diverse diet that was mostly zooplankton, especially amphipods, but included benthos and discards from the fishery, especially squid. P. guntheri also had a moderately diverse zooplankton diet, especially copepods, and benthos and squid damaged by the fishery. The diet of P. tessellata was the most diverse and was mainly benthos, including damaged scallops, with some euphausiids and amphipods. Feeding niche overlap of fish between 10 and 20 cm TL for the three species was between 5.8 and 9.2% for each pair of species, indicating relatively little overlap even though all species scavenged on organisms made available by the fishery. Thus the diet of these three Patagonotothen species indicates substantial feeding niche separation, and that the three species are moderately specialised feeders.     

INVESTIGATING THE BIASES IN THE USE OF HARD PREY REMAINS TO IDENTIFY DIET COMPOSITION USING ANTARCTIC FUR SEALS (ARCTOCEPHALUS GAZELLA) IN CAPTIVE FEEDING TRIALS

The analysis of pinniped scats has been used to quantify their diet, using prey remains to identify species and to estimate the numbers and sizes of prey consumed. There are, however, potential biases involved with scat analysis and, for this method to be used successfully, these biases need to be quantified. Thirty-six Antarctic fur seals ( Arctocephalus gazella ) were fed meals of exclusively either fish, squid, or krill and their scats were collected and analyzed. The major sources of error in the analysis of prey remains from scats were the differential erosion and passage rate of items in relation to their size. However, using simple correction functions, such as those which model otolith erosion, it is possible to reduce these errors. Using plastic beads as dietary markers showed recovery rates were negatively related to their size. Larger squid beaks had lower recovery rates than smaller beaks, but there was no size-related bias in the recovery rates of krill carapaces. Only 33% of the squid beaks and 27% of the otoliths originally fed were recovered from the scats. Recovery rates were greater for squid (77%) and fish (50%) eye lenses and these structures gave a better estimate of numbers eaten. Differences found between experimentally derived and published regression equations (used to calculate prey sizes eaten from prey remains) highlights the need for regression equations based on local prey characteristics, if these are to be used with any success to describe the prey eaten.     

FOOD PREFERENCE AND THE EXTENT OF OVERLAP IN SOME INSECTIVOROUS BIRDS, WITH SPECIAL REFERENCE TO THE TYRANNIDAE

Analysis of published records of the food of flycatchers (Tyrannidae), swallows (Hirundinidae) and vireos (Vireonidae) of North America, and terns (Laridae) of the Pacific Ocean showed that size, rather than taxonomic, differences in food appear to be the most important ones for these birds. Although the distribution of insect sizes in nature approaches a two-parameter log-normal distribution, the distributions of the sizes of food items taken by birds show no significant differences from log-normal. Birds of a given feeding type (e.g. flycatchers) show a strong correlation of average prey size with bird body weight, and a significantly less strong correlation with bill characteristics, indicating that body size is a better predictor of prey size than any single bill character. The slopes and/or intercepts of regressions of food size against body weight are different for birds of different foraging type. Values of overlap in food preference are proportional to the similarity of the two species compared, expressed as the ratio of their weights. Because insect taxa differ in size and because the amount of overlap in taxonomic composition of foods is therefore very roughly proportional to the amount of overlap in size, taxonomic differences in food may merely reflect differences in size preferences. Values for overlap in food are greater than for most cases involving spatial niche parameters, indicating space is more easily divided than food, a conclusion supported by the relative rarity of large flycatchers.     

The relationship between intra–guild diet overlap and breeding in owls in Israel

Even though intra-guild predators frequently prey on the same species, it is unclear whether diet overlap between two predators is a source of interspecific competition or whether predators simply use the same abundant prey resource. We measured the extent to which the diets of barn owls (Tyto alba) and long-eared owls (Asio otus) in Israel overlap and examined whether yearly differences in diet overlap correlate with barn owl breeding success. Pianka’s index of niche overlap was positively related to barn owl population size but not to its breeding success. The number of breeding barn owls was higher when long-eared owls consumed more rodents, suggesting that diet overlap most likely increased when rodents became more abundant. Therefore, in Israel, when these two owl species prey more often on rodents, their diets are more similar and interspecific competition is reduced. Unlike sympatric populations in Europe, in years when rodents are less abundant in Israel long-eared owls switch to hunting alternative prey (e.g., birds), perhaps to avoid competition with barn owls.