年龄和性别因素对奇台沙蜥食性的影响

采用胃容物分析法研究年龄和性别因素对奇台沙蜥(Phrynocephalus grumgrzimailoi)食性的影响.共解剖220号浸制标本,取出整胃分离胃容物,鉴别胃容物中的食物种类,统计各种食物组分的数量频率、重量比例、出现频率和相对重要性指数(index of relative importance,IRI).结...     

Food and feeding habits of pearly razorfish, <Emphasis Type="Italic">Xyrichtys novacula </Emphasis>(Linnaeus, 1758), in the southern Tyrrhenian Sea: variation by sex and size

We collected pearly razorfish, Xyrichtys novacula (Labridae), from a sandy bottom of Capo dOrlando (Tyrrhenian Sea) to study their feeding habits. We caught fish by hand-lines and seine nets and identified, counted and weighed food items in their stomachs. We evaluated the importance of the different prey types by calculating the frequency of occurrence, abundance and mass. We used these values to calculate the index of relative importance (IRI) for each taxonomic category and a modified index (MI) which did not incorporate %N into the formula. Gammarideans were dominant food items in terms of %F but showed a very low MI value; the bivalve Acanthocardia tubercolata was the dominant species in terms of %W and MI value. The prey items were mostly benthic organisms belonging to the assemblage of fine, well-sorted sands showing that X. novacula is a benthic feeder. Despite the large number of prey taxa found, few species accounted for most of the prey consumed, indicating specialist feeding, as confirmed by the low value of the Levins standardised index. We found significant differences in prey distribution among size classes, with the highest prey diversity recorded in 120–140 mm TL size classes. Correspondence analysis showed a trend of increasing predator size classes among prey items, with small prey, such as copepods and amphipods, in the smallest predator size classes and bigger prey in the larger ones. Prey caught by larger predators differed significantly in weight from those taken by smaller specimens. There was a significant positive correlation of fish length with bivalves, decapods and echinoids, and a negative one with copepods, ostracods, caprellids, gammarideans. There were no significant differences between males and females of overlapping sizes. Juveniles are able to exploit only small, vulnerable prey, while adults take few, large prey, thereby avoiding competition with juveniles.     

Ontogenetic diet shifts and diet overlap among three closely related neotropical armoured catfishes

Analysis of stomach contents of larvae, juveniles and adults of three closely related armoured catfishes in coastal plain swamps of Suriname, South America, revealed major ontogenetic diet shifts. Larvae of Hoplosternum littorale, H. thoracatum and Callichthys callichthys showed a restricted diet of rotifers and microcrustacea. Diets of juveniles and adults were more diverse, but large volumes of microcrustacea, aquatic insect larvae and detritus were always present. In all three species diet overlap was especially low between larvae and adults. The proportion of detritus in the stomach was positively related to fish size, which probably reflected the inability of larger size classes to separate effectively benthic invertebrate prey from bottom substrata, and not a specialization towards detritivory. Adults showed only minor seasonality in diet composition, and no effect of season on diet breadth. Interspecific diet overlap among larvae, juveniles and adults did not differ significantly from 1 (complete overlap). Lack of food partitioning was probably related to a similar morphology of the three catfishes. Intraspecific between-size class diet overlap was considerably lower than interspecific within-size class overlap. It is suggested that large interspecific diet overlap occurred because food was not limiting the populations of these catfish. However, the nest habitats of the three species differed.     

Diet of age-0 tarpon (<Emphasis Type="Italic">Megalops atlanticus</Emphasis>) in anthropogenically-modified and natural nursery habitats along the Indian River Lagoon,Florida

As human development in coastal areas increases, the role of anthropogenically-created habitats in the life history of marine organisms is becoming increasingly important. We examined the diet of age-0 tarpon, Megalops atlanticus, in and around man-made mosquito control impoundments along the Indian River Lagoon in east-central Florida, with a particular focus on identifying dietary patterns associated with tarpon size and nursery habitat type (i.e., between perimeter pool habitats in established impoundments and newly-created restoration marsh habitats). Age-0 tarpon were found to consume a wide variety of prey organisms, and exhibited considerable dietary variation among study sites. Smaller juvenile tarpon consumed a limited number of small prey taxa, while larger individuals fed on a greater range of prey taxa and sizes. Overall, copepods and fishes were the dominant prey items; however, the consumption of these organisms varied considerably among size classes and sites. There was no clear difference in tarpon diet between the two types of habitat we examined. The ability of juvenile tarpon to utilize such a diverse range of prey organisms may allow populations to inhabit a variety of habitats, including man-made marshes. When natural systems have been degraded or destroyed, human-altered habitats can assume a nursery role for the species.     

山地麻蜥个体发育过程中头部两性异形和食性的变化

研究了山地麻蜥(Eremias brenchleyi)个体发育过程中头部两性异形和食性的变化．成体个体大小(SVL)无显著的两性差异,但雄体具有较大的头部(头长和头宽)．头部两性异形在孵出幼体就已存在,成体头部两性异形比幼体(包括孵出幼体)更为显著,雄性较大的头部与其头部随SVL的增长速率大于雌性有关．两性头部总体上随SVL呈异速增长,表现为个体发育过程中头长和头宽与SVL的线性回归方程斜率有显著的变化．孵出幼体有相对较大的头部,这种形态特征是胚胎优先保证生态学意义更为显著的头部生长的结果,有利于孵出幼体的早期生存和生长．相对头部大小在个体发育过程中有显著的变化．不同性别和大小的山地麻蜥摄入食物的种类及各种食物在摄入食物中所占的比例有一定程度的差别,食物生态位宽度和重叠度因此有一定的差别．然而,没有直接的证据表明头部两性异形能导致两性食物生态位的明显分离,并有利于减缓两性个体对食物资源的竞争。     

Ontogenetic and seasonal feeding habits of the annular seabream (Diplodus annularis L.) in Zostera sp. beds, eastern Adriatic Sea

The ontogenetic and seasonal feeding habits of 489 individuals of Diplodus annularis L. in Zostera sp. beds were studied using frequency of occurrence, numeric percentage, percentage by weight and an index of relative importance (IRI). The Schoener overlap index was used to compare diets by fish size and seasons. Diet composition including 18 different prey taxa was evidently heterogenous among annular seabream size groups, with no significant dietary overlap. In relation to body size, juveniles were omnivorous and preferred algae and plants, while adults were omnivorous and preferred animals. Principal prey items by decreasing importance were Chlorophyta (dominant taxa Cladophora sp. and Enteromorpha flexuosa), Decapoda (dominant species Upogebia pusilla), Gastropoda (important species Rissoa ventricosa and Alvania sp.), Bivalvia and Spermatophyta, as well as other food items (mostly unidentified Crustacea and fish eggs). Differences in diets among seasons were not significant.     

Effect of Removal from the Family Group on Feeding Behavior by Captive Callithrix jacchus

Common marmosets are omnivorous primates with a highly diversified diet. There is no study describing if and how the diet is learned. Infants get their first bits of solid food from other monkeys in the group, which suggests that they may need an introduction to food items by older individuals before including them in their diet. We assessed the acceptance of novel and familiar food items by common marmosets, both isolated and in their family groups. We tested adult, subadults and juveniles from 5 captive families while isolated and in their family groups. The test consisted of presenting for 10 min novel and familiar food items to isolated individuals or to the whole family. We recorded the latency to start eating and the number of food items ingested. When isolated, adults ate more novel and familiar food items than juveniles did. They also started eating sooner than juveniles did. When tested alone, all juveniles, except one, never tasted novel food, and juveniles ingested fewer familiar food items than adults did. When tested in their family groups, juveniles ingested more familiar and novel food than when they were isolated. Our results suggest that: 1. juvenile common marmosets show more food neophobia than adults do, especially when alone; 2. the family group may facilitate the acceptance of novel food items by juveniles; 3. the family group, besides promoting the acceptance of novel food, may also increase its ingestion; and 4. dietary acquisition in Callithrix jacchus involves social facilitation.     

Diet,intestinal morphology,and nitrogen assimilation efficiency in the damselfish,Stegastes lividus,in Guam

Synopsis Analysis of stomach contents of Stegastes lividus revealed a shift from omnivory in juveniles to herbivory in adults. Red algae, primarily Polysiphonia spp., Gelidiopsis intricata and Ceramium spp., formed the bulk of the diet in all size classes. Foraminifera and small crustaceans were of particular importance in the diet of juveniles. Net nitrogen assimilation efficiency of field-fed S. lividus was approximately 61%, and did not vary with fish size. The net assimilation efficiencies of fish fed on Enteromorpha in the laboratory ranged from size-class means of 36–79% for nitrogen and from 29–72% for total organic material, and increased from juveniles to adults. Apparently, juveniles compensate for a lower efficiency in assimilating plant food by including a higher percentage of animal material in their natural diets. The ratios of intestine length to standard length (IL/SL) and to intestine diameter (IL/ID) increased rapidly in juveniles and leveled off in adults. The retention time for ingested food items may only be about one-half as long in juveniles as in adults. The combination of observed IL/ID ratios and estimated retention times proved most valuable in the interpretation of ontogenetic changes in feeding habits and assimilation efficiency.     

Comparisons of the food niches of three native and two introduced fish species in an Australian river

Synopsis The breadth, correspondence and overlap of the diets of the small and large size classes of the three native species (Galaxias occidentalis, Bostockia porosa and Edelia vittata) and two introduced species (Gambusia holbrooki and Perca fluviatilis) of fishes found in the shallows of the main channel and in the tributary creeks of a south-western Australian river have been compared in each season. Classification and ordination were used to examine the overall interrelationships of the diets across species, size groups, seasons and the location where the fish were caught (channel or creek). The smaller fish had a narrower dietary breadth than larger fish in the spring and summer, presumably reflecting the size limit imposed on prey size by their possession of a relatively small mouth in these seasons. Intraspecific dietary overlap between large and small size classes was usually high in G. occidentalis, but generally low in G. holbrooki and P. fluviatilis, and also in B. porosa and E. vittata when the difference between the lengths of the two size groups was greatest. Dietary overlap was least in autumn when the main prey taxa were most abundant. During winter, the diets of the three native species in the tributaries converged, probably reflecting a relatively low faunal diversity in these highly seasonal water bodies. The only relatively consistent interspecific overlap in diet was between B. porosa and E. vittata. Classification and ordination of the dietary samples separated G. occidentalis (which fed extensively on terrestrial organisms from the water surface) from the smaller P. fluviatilis (that concentrated on copepods in the plankton) and from B. porosa and E. vittata (which ingested primarily benthic organisms). Furthermore, B. porosa tended to ingest larger prey taxa than E. vittata. The diet of Gambusia holbrooki is sometimes dominated by terrestrial insects and at other times by benthic organisms, demonstrating that this species is an opportunistic carnivore. It is concluded that food partitioning by the three native and two introduced fish species in the Collie River is likely to be one of the principal factors facilitating the coexistence of substantial populations of these species in this system.     

Ontogenetic shift in dietary preference and low dietary overlap in rohu (<Emphasis Type="Italic">Labeo rohita</Emphasis>) and common carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis>) in semi-intensive polyculture ponds

In order to investigate ontogenetic changes in diet and diet overlap between rohu (Labeo rohita) and common carp (Cyprinus carpio) in polyculture ponds, food preferences of different size classes of these fishes were quantified. Rohu diet consisted of both phytoplankton and zooplankton, and there was a distinct ontogenetic shift in the relative importance of these food items. Zooplankton was the dominant food for rohu up to 20.6 cm total length (TL) and then gradually decreased in importance as fish grew. Phytoplankton was initially a minor component of rohu diet but gradually increased in importance and became the dominant food for rohu at 24.2 cm TL. Phytoplankton biovolume in rohu guts was positively correlated with fish size (TL). Chesson’s α indicated that rohu of all sizes preferentially selected Cladocera and avoided Cyanophyceae and Euglenophyceae. Young rohu initially preferred Rotifera and Copepoda but gradually switched to Bacillariophyceae and Chlorophyceae. Common carp diet consisted of phytoplankton, zooplankton, and benthic macroinvertebrates, but was dominated by benthic macroinvertebrates (63–92% of total diet). As common carp grew, the proportion of zooplankton ingested decreased and the proportion of benthic macroinvertebrates increased. Benthic macroinvertebrate biovolume in common carp guts was positively correlated with fish size. Common carp of up to 15.4 cm TL preferentially selected zooplankton, but common carp larger than 18.9 cm TL avoided this food item. Common carp of all sizes avoided phytoplankton. A low dietary overlap was found between rohu and common carp (Schoener overlap index: 0.08–0.35), probably due to ingestion of smaller quantities of zooplankton by the latter. Dietary overlap also decreased with increasing rohu and common carp size because of divergent ontogenetic shifts in dietary preferences of the two species.     

Effects of riparian canopy cover on salmonid diet and prey selectivity in low nutrient streams

This study focuses on stream sections within a relatively low nutrient catchment in south‐east Ireland in an attempt to characterize the probable effects of riparian canopy on salmonid diet and prey selectivity within two size classes of nursery stream. Sampling found that brown trout Salmo trutta diet changed significantly in response to riparian canopy regardless of stream size. The observation that S. trutta within unshaded stream sites did not feed on drifting terrestrial prey items to the same extent as those within shaded streams was not due to a lack of availability of this food source. There was no evidence to suggest that S. trutta selectively choose particular prey items.     

Prey use of the fishing spider Dolomedes triton (Pisauridae,Araneae): an important predator of the neuston community

Summary Prey of feeding juvenile and adult Dolomedes triton (Walckenaer 1837) were sampled over two seasons on three small ponds in central Alberta, Canada. Prey were mainly insects active at the water surface with truly aquatic species making up about 14% of the diet. Throughout the season aquatic and semi-aquatic Heteroptera represented about 30% of the prey. Diptera and adult Odonata were also important prey items but their abundance in the diet was more variable seasonally. Of the 625 prey items recorded nearly 50% were represented by taxa taken no more than once by spiders in one of the five size classes (adult females, adult males, large, intermediate and small juveniles). Large spiders did not take the smallest prey available, although small and intermediate-sized spiders fed on nearly the full size range taken by larger spiders. Cannibalism was common, accounting for 5% of the observations, with females and large juveniles as the most frequently observed cannibals. We hypothesize that intraguild predation (including cannibalism) could be an important coevolutionary force structuring phenology, population dynamics and microhabitat use of the predatory guild of the neuston community.     

Feeding of the Brazilian sharpnose shark Rhizoprionodon lalandii (Müller & Henle, 1839) from southern Brazil

Stomach contents of 175 Brazilian sharpnose sharks, Rhizoprionodon lalandii, were examined to assess their feeding habits. Caught by artisanal fishers between June 2006 and July of 2007 at Ipanema Beach, on the central coast of the state of Paraná, southern Brazil, R. lalandii appears to be piscivorous (89.7% IRI), feeding mainly on families of Clupeidae (5.1% IRI) and Sciaenidae (3.7% IRI) and secondarily on squid Lolliguncula brevis (8.6% IRI). Diets were similar between sexes and seasons. However, there were ontogenetic changes in their diets. The major items for neonates were Penaeidae crustaceans (%IRI = 56.4) and Engraulidae fishes (%IRI = 29.3); for the juveniles, Sciaenidae (%IRI = 62.5) and Clupeidae (%IRI = 19.7), and for adults Clupeidae (%IRI = 45.8) and Sciaenidae (%IRI = 15.9). Only neonate sharks consumed crustaceans, whereas all size classes consumed cephalopods and teleosts. Neonates had the lowest trophic level (TR = 3.8), with the diet of juveniles and adults the highest (TR = 4.2). Given its abundance and diet, R. lalandii may be an important predator of demersal and pelagic prey in coastal waters of Brazil.     

Variation of diet and food consumption of the scaldfish Arnoglossus laterna(Walbaum, 1792)

Although Arnoglossus laterna (Walbaum, 1792) is a common benthic fish in Portugal, several aspects of its feeding ecology remain incomplete. In this study, diet was examined and the food consumption estimated on inshore waters in the central coast of Portugal. The diet of the scaldfish included mainly mysids, amphipods and polychaetes, while the decapod Philocheras bispinosus Hailstone was the most frequent prey. Variation in the diet according to fish size (two size‐classes), sex and spawning/non‐spawning seasons was examined. Significant differences in diet composition were found between seasons, whereby the most frequently consumed prey in the non‐spawning season were crustaceans, while polychaetes and nemerteneans comprised most of its diet in the spawning season. The average value of the vacuity index was very similar between seasons (24.7% non‐spawning season and 25.0% spawning season). Diet overlap between size‐classes and sexes was high, but low between the two seasons. Food consumption of A. laterna was estimated based on diet characterization and evacuation rate. The estimate of the daily food consumption was 1.15 mg dry weight for the non‐spawning season and 1.67 mg dry weight for the spawning season. Significant differences in consumption rates were found between size‐classes and sexes. Larger individuals showed a higher consumption rate than smaller individuals. Females consumed more food than males in the non‐spawning season, while in the spawning season males consumed more than females.     

Influence of breeding on foraging behaviour and diet of Crowned Cranes

Gichuki, N. 2000. Influence of breeding on foraging behaviour and diet of crowned cranes. Ostrich 71 (1 & 2): 74–79.

This study investigated the effects of breeding on the foraging behaviour and diet of Grey Crowned Cranes Balearica regulorum in the Kitale area, western Kenya. During the study period (July 1994 to June 1996), 86 pairs successfully bred in 290 ha of wetlands. Egg-laying extended from July to November, indicating that the timing of nesting was flexible. Breeding and non-breeding adult cranes as well as immature birds foraged in grasslands, crop stubble, sown fields and wetlands. These habitats were not used according to the available proportions. Breeding commitments restricted parental adults to habitats within their territories while non-parental adults and sub-adults exploited a wider range of habitats. Further, breeding adults and juveniles had smaller foraging range sizes than those of non- breeding adults and sub-adults. Reduced foraging ranges by breeding birds appear to be a trade-off between energy expenditure in self-maintenance and reproduction. The diet of breeding and non-breeding adults was generally similar consisting of graminoid seeds and arthropods. Breeding birds, however, increased consumption of protein rich animal food and non-seed plant material. Selectivity of food items was greatest among breeding adults. Overall, breeding commitments resulted in reduced foraging range and variety of food items available but the major components of the diet remained the same for all adults and immature birds.     

Partitioning seasonal time: interactions among size, foraging activity and diet in leaf-litter frogs

This study investigates hypotheses about partitioning of food resources among all species and several size classes in an assemblage of diurnal leaf-litter frogs in central Amazonia. All species in this assemblage change the type and size of prey as they grow. An ordination of diet composition was significantly associated with frog size and species-specific behaviour. However, a partial Mantel analysis indicated that species explained about 1.5 times more of the variation in diet overlap between individuals than frog size. Diet and foraging activity are correlated in juveniles, but not in adults, and this result holds whether species are considered as statistically independent observations or whether relationships are analysed using phylogenetically independent contrasts. This study showed that the partitioning of food resources between species changes with the population size structures. Thus, intraspecific and interspecific changes in diet, coupled with different patterns of juvenile recruitment, cause diet segregation among species due to temporal segregation of equivalent size classes. Received: 25 August 1997 / Accepted: 1 April 1998     

Diet and endogenous carbohydrases in the temperate marine herbivorous fish Kyphosus sydneyanus

Stomach content analysis showed that Kyphosus sydneyanus collected by spearing from coastal reefs near Leigh, north-east New Zealand, is herbivorous throughout its post-settlement life. All size classes collected consumed a wide range of macroalgae, predominantly the phaeophytes Carpophyllum maschalocarpum and Ecklonia radiata , and the rhodophytes Gigartina macrocarpa and Caulacanthus ustulatus . Multivariate analysis of variance and discriminant function analysis showed an ontogenetic diet shift, with gut contents dominated by rhodophytes and chlorophytes in juveniles and by phaeophytes in adults. No significant seasonal variation in diet was apparent for any size class. Amylase and amylopectinase activity was highest in the smaller size classes. Laminarinase activity was relatively low throughout all size classes, and did not correlate with the ontogenetic increase in phaeophyte consumption. The activity of endogenous enzymes hydrolysing starch broadly correlated with rhodophyte consumption in adult specimens of three other species of marine herbivorous fishes: Odax pullus, Girella tricuspidata and Parma alboscapularis; laminarin hydrolysis, however, was negatively correlated with phaeophyte consumption. Overall, the diet and enzyme results suggest that juvenile K. sydneyanus rely mainly on the endogenous digestion of rhodophytes and chlorophytes, while exogenous (microbial) digestion is used to digest the phaeophyte-dominated diet of adults.     

Trophic behaviour of juvenile reef fishes inhabiting interlinked mangrove–seagrass habitats in offshore mangrove islets

Stable isotope (δ¹³C and δ¹⁵N) and gut content analyses were used to investigate size‐related feeding habits of four reef fishes (the beaugregory Stegastes leucostictus, the french grunt Haemulon flavolineatum, the schoolmaster snapper Lutjanus apodus and the yellowtail snapper Ocyurus chrysurus) inhabiting an offshore (non‐estuarine) mangrove islet off Belize, Central America. Comparisons of isotopic niche space and Schoener diet similarity index suggested a low to moderate degree of niche overlap between fish size groups. The δ¹³C gradient between mangrove and seagrass prey as well as results of Bayesian mixing models revealed that sampled fishes relied mostly on seagrass prey items. Only small and large juveniles of the carnivorous species L. apodus derived a part of their diet from mangroves by targeting mangrove‐associated Grapsidae crabs and fish prey, respectively. Isotopic niche shifts were particularly obvious for carnivorous fishes that ingested larger prey items (Xanthidae crabs and fishes) during their ontogeny. The utilization of mangrove food resources is less than expected and depends on the ecology and life history of the fish species considered. This research highlights that mangrove‐derived carbon contributed relatively little to the diets of four fish taxa from an offshore mangrove islet.     

Prey selection of subadult male northern fur seals (Callorhinus ursinus) and evidence of dietary niche overlap with adult females during the breeding season

During the breeding season northern fur seals (Callorhinus ursinus) congregate on the Pribilof Islands in large numbers creating the potential for intraspecific competition. Due to the declining trend in the Pribilof Islands population of fur seals, it is important to understand how prey resources are partitioned among the population. Fur seals exhibit a high degree of sexual dimorphism resulting in energetic differences among age and sex classes. Therefore, we hypothesized that subadult male and adult female fur seals would differ in the type and size of prey consumed. We examined the diets of subadult male (age 2–8; mean mass 28–176 kg) and adult female (age ≥ 3 yr; mean mass 13–50 kg) seals on St. Paul Island from 1992 to 2000. Prey remains found in fecal samples were compared using niche overlap indices. There was nearly complete dietary niche overlap between subadult male and adult female fur seals. Walleye pollock (Theragra chalcogramma), Pacific salmon (Oncorhynchus spp.), Pacific herring (Clupeia pallasi), and cephalopods were common prey items found in the diets of both groups. We found differences in the size of pollock consumed and that geographic location of sample collection may be important in determining diet differences. Our results indicate high levels of dietary overlap among subadult male and adult female fur seals.     

Ontogenetic,seasonal, and spatial variation in the diet of <Emphasis Type="Italic">Heterotis niloticus</Emphasis> (Osteoglossiformes: Osteoglossidae) in the Sô River and Lake Hlan,Benin, West Africa

The African bonytongue, Heterotis niloticus (Osteoglossidae), is an important fisheries and aquaculture species in West Africa. This species has frequently been characterized either as an omnivore, insectivore or detritivore, the latter, in part, because of its benthic feeding habitats and possession of a gizzard (thick-walled pyloric stomach). We examined diets of two populations of H. niloticus in the Sô River in southern Benin. A population from the river channel and seasonally flooded marginal plains was dominated by juvenile and subadult size classes. Adults size classes were common in a second population from Lake Hlan, a natural lake in the river floodplain located upstream from the channel study region. Heterotis of all sizes consumed a variety of food resources, ranging from aquatic invertebrates to small seeds. Aquatic invertebrates composed a large proportion of the diets of juveniles, and adults consumed a mixture of aquatic invertebrates, seeds, and detritus. Seasonal dietary variation was observed in both populations, and diet breadth was not significantly different between populations. Aquatic invertebrates remained significant in diets of larger size classes; diets of fish between 100 and 200 mm began to include seeds and detritus, with a marked increase in the volumetric proportion of detritus in diets of fish between 300 and 400 mm in Lake Hlan and between 500 600 mm in the river. Relative gut length was inversely related to body size, which supports the notion that Heterotis is an omnivore and not a specialized detritivore. The thick-walled gizzard of Heterotis, which generally contained sand, probably aids digestion of seed coats. Because Heterotis consume mostly invertebrates and grass seeds in shallow waters of seasonal aquatic habitats and lakes the river floodplain, foraging success and fishery production should be strongly dependent on the annual flood pulse.