京都大学 / 0048 / 新制・課程博士 / 博士(地球環境学) / 甲第21236号 / 地環博第172号 / 新制||地環||35(附属図書館) / 京都大学大学院地球環境学舎地球環境学専攻 / (主査)教授 山下 洋, 教授 柴田 昌三, 教授 宮下 英明 / 学位規則第4条第1項該当 / Doctor of Global Environmental Studies / Kyoto University / DFAM

The stomachs of 183 bigeye tuna (Thunnus obesus) were collected from three Taiwanese longline vessels operating in the western Indian Ocean during the period from April to December 2006. Their fork lengths ranged from 89 cm to 198 cm. Distinguishable prey species, belonging to 28 families, consisted mostly of fishes, crustaceans, and cephalopods. Fourteen prey items were newly recorded, among them, five families (Anoplogastridae, Bothidae, Grammicolepididae, Pinguipedidae, and Sciaenidae) were firstly recorded as the diet of bigeye tuna. Two special dietary species were found for the first time. Oplophorus gracilirostris, the dominant numerical dietary species, had not been previously recorded; however, O. typus had been recorded in the past. The other special prey species found, Macrurocyttus acanthopodus (Grammicolepididae), was not only the first record in the stomach of bigeye tuna but also the first record in the Indian Ocean. The largest body length among prey species was 655 mm for an Alepisaurus ferox. The number of prey species appeared to decrease with the increasing size of the bigeye tuna, and larger Indian bigeye tuna tended to feed on larger prey species. In the stomachs of the bigeye tuna, Oplophorus gracilirostris was the most frequently found prey species (56.8%). The three numerical most dominant prey species were Oplophorus gracilirostris (38.2%), Charybdis smithii (8.8%), and Alepisaurus ferox (3.6%). In addition, the fish prey item, Alepisaurus ferox, had the highest percentage of prey weight. The total percentage of prey weight for prey fishes is similar to that reported in previous research, but the average weight of Alepisaurus ferox is heavier than the previous report. Additionally, the percentage of prey weight for Paralepididae is lower than the results of previous studies. Moreover, the occurrence and percentage of the number of Oplophorus gracilirostris are distinctly higher than the results of previous studies. These findings indicate that the percentage of mean preys number exist seasonal effect. Alepisaurus ferox was the most numerous prey in May to July; Oplophorus gracilirostris was the most numerous prey at 200 m depth in August to October 45¢XE to east; Charybdis smithii was the most numerous prey at 50 m depth in August to October 45¢XE to west. A possible reason to explain this phenomenon is that there are strong feeding migratory behavior in bigeye tuna.

Effective population management of green sea turtles (Chelonia mydas) necessitates understanding the temporal variation in foraging grounds used in ontogenetic stages, and the effect that the assimilated diet within those habitats has on nutritional gain, growth and eventual reproductive output. Texas coastal waters provide foraging grounds critical to meeting the nutritional needs of green turtles during early life history. To characterize temporal shifts in foraging strategy stomach contents combined with stable carbon (delta13C) and nitrogen (delta15N) isotopes of scute tissue were examined across size classes of stranded juvenile green turtles from the middle and lower Texas coast during 2007-2010. Findings from dietary analysis generally corroborated those from stable isotopes in scute samples. Results indicate green sea turtles exhibit multiple shifts in diet and habitat along the Texas coast. Although isotope values in the tissues of some <25 cm SCL turtles signified recent recruitment to jetty habitat, most in this size class exhibited depleted delta13C and enriched delta15N values indicative of oceanic life. Reinforcing oceanic occupancy from stable isotope results was forage material dominated by oceanic items such as Sargassum spp., Scyphozoa spp., and plastic debris. Diet analysis of 25-34.9 cm SCL turtles implied regional differences existed in macroalgae and seagrass consumption. Enriched delta13C and delta15N values in newest scute suggest most turtles inhabited the jetty environment, where macroalgae is the most available forage. A definitive shift by >35 cm SCL turtles to inshore seagrass habitat was revealed by a diet of seagrasses and tissue enriched in delta13C and depleted in delta15N. This is the first study to integrate stomach contents of several green turtle size classes with tissue analysis of stable isotopes. The combination of these techniques provided an assessment of the effectiveness of stable isotope analyses in documenting diet and habitat shifts. Stomach content examination determined the most recent diet consumed within the habitat occupied, whereas stable isotope analysis provided a time-integrated synopsis of diet and habitat shifts. Findings indicate integration of stomach content and stable isotope analysis is highly effective for characterizing habitat use and foraging strategy of ontogenetic-stage green sea turtles.

Methods used to estimate the prey consumption by large carnivores include direct continuous observation, stomach content analysis, carcass observations and scat analysis. Continual observations are widely considered the best approach to estimate large carnivore diets, with lions (Panthera leo) being no exception. Continual observation allows the recording of all prey encounters and biases inherent in the other approaches are minimised. However, continuous observations are not always feasible, and in situations where animals cannot be observed at all times, diets are often estimated from observed carcasses. This often leads to an over-estimation of large kills in the estimated diet. Alternative methods that are free of the constraints placed on continuous observations are needed to provide data of a similar quality to that obtained using these continuous observation bouts. I employed a cluster follow up technique to locate lion kills from remotely accessed Global Positioning System (GPS) data from lions in the Kruger National Park (KNP). I develop Generalized Linear Models (GLMs) that increase the probability of locating kills at GPS cluster events. By increasing the predictive ability of detecting kills I show that this technique can be used to locate kills in a more efficient manner than random searching of GPS clusters, with further advantages in that multiple groups of lions can be monitored simultaneously. By incorporating this technique into an adaptive research framework, the diet of lions (and that of other large carnivores) can be estimated. In addition, I show that the spatial association between lions at kill sites, while feeding on carcasses, provides a further increase in the predictive ability of kill site models. Lionesses were found to be considerably closer together at the start of clusters associated with kills in comparison to clusters where no kill was found. This pattern remained consistent for both small and large kills. This proximity approach could therefore be incorporated into the GLMs that are developed to predict kill sites of large social carnivores. To further reduce the bias (where small kills are often missed) inherent in carcass observations, I combined scats and carcasses collected from known times, locations and lion groups to construct a temporal kill record for each group of lions. By combining scats and carcasses I estimate that at least 50% of the small prey items, namely impala (Aepyceros melampus) and warthog (Phacochoerus africanus) were missed when GPS clusters were investigated for carcasses. Ultimately, I show that a combination of GPS cluster investigations based on models developed using GPS movement data in combination with lion proximity data, augmented with scats collected at GPS clusters, could provide estimates of large carnivore diets that begin to approach estimated diets obtained through continuous monitoring. The resulting diet, estimated from the GPS cluster approach in combination with scat collection, indicated that the dominant prey item in the region was zebra (Equus quagga) followed by wildebeest (Connochaetes taurinus), impala and buffalo (Syncerus caffer). Selection indices for the eight dominant prey items were calculated using prey availability measures obtained from the aerial census data and ground counts of groups. It has been suggested that group level selection is a better approach to calculating predator-prey interactions, and that stability in predator-prey systems is improved if group metrics of prey are used as apposed to individual measures of availability. I show that there is a considerable shift in selection indices, as well as in the order that prey is selected, when using different measures of prey availability. In selection studies, more effort needs to be paid to the assessment and definition of prey availability to ensure results accurately reflect selection patterns in the field, especially when data are used for the development of management practices. Combining buffalo predation data collected from GPS cluster investigations with buffalo mortality data collected over five years prior to the commencement of the GPS cluster investigations, allowed an investigation into patterns of lion predation on buffalo between 2000 and 2007. Buffalo of both sexes were more vulnerable to predation in habitats that gave lions an ambush advantage (i.e. increased grass height and tree density). Despite this similarity in landscape risk, different processes lead to similar fates in dangerous habitats for buffalo of both sexes. Predation pressure by lions on buffalo increased following periods of reduced rainfall; with more buffalo predated on following drier six month periods. Predation on males constituted a significant proportion of all predation and was focused predominantly into the late dry season. The resulting method of locating kills by using GPS clusters and correcting carcass data with scats collected along the movement path represents a robust technique to estimate large carnivore diets. In the concluding chapter I present avenues where future research can build on the current thesis and present a framework that can be employed when attempting to estimate large carnivore diets. / Thesis (PhD)--University of Pretoria, 2010. / Zoology and Entomology / unrestricted

This study addressed a fundamental question in applied ecology and conservation; what is the predatory impact of introduced piscivorous sport fish on imperiled native fish populations? More specifically, which of many introduced species and size-classes represent the greatest threats and should be targeted for control? In order to explore this important question, an integrated analysis of stable isotopes, quantified observed diet analysis, and stable isotope mass-balance models were used to quantify trophic interactions. These tools were used to construct food web models that were then compared to draw inferences regarding the relative contribution of prey fish, including rare native fish, to the diet of introduced sport fish. The stable isotope-derived food web illustrated a slight decoupling in energy flow between a pelagic and a benthic-littoral sub-web. The quantified diet analysis suggested piscivory was low overall, and that the introduced sport fish assemblage relied heavily on zooplankton and aquatic insect prey. The integrated stable isotope and quantified diet analysis demonstrated that the consumption of prey fish, particularly pelagic prey fish, was typically underestimated using stomach content analyses. From the evaluation, comparison, and integration of food web models, I suggest that substantial predation was occurring on the early life stages of Utah Lake fishes, including native fishes, and it was not being observed using stomach content analysis. My comparative modeling demonstrated that introduced sport fish are an impediment to native fish conservation and identified the small size-class of white bass as the most immediate threat.

Herbivorous fishes include a diverse assemblage of species that target primarily benthic autotrophs. This is perhaps one of the most well-studied groups of coral reef fishes, often reputed to be key components of coral reef communities, contributing to coral reef health in numerous ways. Through their feeding ecology and benthic interactions, they help mediate algae-coral interactions which can allow for improved coral survival and health. Despite the wealth of literature documenting the prominent roles of these fishes in coral reef ecosystems, studies from the Red Sea are surprisingly lacking. The Red Sea is a marginal reef environment, with a host of unique environmental and biological characteristics making it a unique environment where dynamics of herbivory may differ. This dissertation aims to fill key gaps in our knowledge of herbivorous fishes through the study of their distribution and trophic ecology. Herein, I describe habitat-specific partitioning of Red Sea herbivorous fish assemblages, discovering higher diversity and abundance found in reefs closer to shower, dissimilar to findings from other regions. Cross-shelf variation in assemblage structure seems to be quite robust through time, indicating short-term stability in herbivore assemblages. Through the use of stomach contents and stable isotope analyses, I then investigate the trophic ecology of browsing herbivores across the same shelf-gradient. I found higher trophic redundancy on nearshore reefs through time, with increased variation in diet and high levels of complementarity on offshore reefs where macroalgae are scarce. Stable isotope analyses of both liver and muscle revealed the stability of this resource partitioning through time, demonstrating for the first-time temporal stability of resource partitioning within this group. This dissertation broadens our knowledge of herbivorous fishes, filling important gaps. It offers new insight into the role of habitat in structuring trophic ecology and how flexible the diets of browsing species can be. Together, this information creates a foundation where improved knowledge of herbivorous fish ecology could be incorporated into future management plans of ongoing giga projects within the Kingdom. Incorporating herbivores into these plans could allow for increased resiliency for Red Sea coral reefs in the face of future development and shifting climatology.

Understanding how diversity is partitioned along natural and anthropogenic gradients within ecosystems is important to predict the persistence of species and the ecological functions they provide. Dottybacks (Pseudochromidae) are a diverse group of mesopredators that feed on cryptic macroinvertebrates and newly recruited fishes. This diet behavior may modify the composition and abundance of cryptobenthic fauna within coral reef ecosystems. Understanding how mesopredators partition their diet and the functional role provided by available prey within reefs can assist in understanding the ecological role these predators contribute to coral reef trophodynamics and the effect of their population changes on the reef ecosystem. To assess the diet of three common Pseudochromis species and two distinct color morphs of P. flavivertex in the Red Sea, I used a combination of i) visual stomach content analysis, ii) stomach DNA metabarcoding (18S, COI), and iii) stable isotope analysis (δ15N, δ13C). I evaluated i) dietary niche breadth, ii) variation in diet composition, iii) degree of dietary specialization, and iv) trophic level. These techniques revealed partitioning in the dietary composition and resource use between P. flavivertex, P. fridmani, and P. olivaceus. Although the two technics used for stomach content analysis did not show differences in the dietary composition within color morphs of P. flavivertex, the isotopic signature showed marked differences in the isotopic niche and resource use between morphs. Resource partitioning appears to be driven by variation in resource availability in the fish habitat and by subtle differences in the ecology of these species. These findings provide evidence of species-specific differences in the trophic ecology of pseudochromids in the Red Sea and demonstrate their important role as predators of cryptic invertebrates and small fish, being key components in energy transfer in coral reef ecosystems by acting as a link between cryptofauna and higher trophic levels. This study highlights the importance of combining several approaches (short-term: visual analysis and DNA metabarcoding; and long-term: isotope analysis) when assessing the feeding habits of coral reef fish, as they provide different and complementary information necessary to delimit their niches and understand the role that small mesopredators play in coral reef ecosystems.

The combined methods of stomach content analysis and stable 15N and 13C isotope biochemistry analysis were used to investigate the trophic dynamics and feeding ecology of coastal pelagic fishes in the waters off southeastern Florida, USA. The coastal pelagic fish complex includes blackfin tuna Thunnus atlanticus, dolphinfish Coryphaena hippurus, king mackerel Scomberomorus cavalla, little tunny Euthynnus alletteratus, skipjack tuna Katsuwonus pelamis, and wahoo Acanthocybium solandri. These coastal teleosts, particularly the tunas and wahoo, are primarily targeted by recreational anglers. However, there is a shortage of available trophic and diet composition data concerning these fishes of the coastal pelagic ecosystem. Stomach and muscle tissue samples were collected from the species of various lengths over a three-year period from March 2010 and March 2013. Across all six species, teleost fishes dominated the prey with an average 64.5% by occurrence, 63.7% by number, and 89.9% by weight. There were two dominant prey families: Clupeidae and Carangidae. Dolphinfish showed the lowest diet overlap among the six species, due to the highly diverse diet. The highest diet overlap occurred between king mackerel and little tunny. The mean δ15N ranged from 8.21 ‰ (wahoo) to 13.18 ‰ (king mackerel), and the mean δ13C ranged from -18.41 ‰ (king mackerel) to -16.70 ‰ (dolphinfish). Blackfin tuna exhibited the largest δ15N range (7.22 to 13.21 ‰), as well as the largest δ13C range (-19.13 to -12.99 ‰). The δ15N and δ13C signatures in the muscle tissue showed evidence of shifts to higher trophic levels with an increase in fish size and the formation of distinct trophic groups among the coastal pelagic predators. The δ13C also suggested an inshore-offshore spatial relationship among the coastal pelagic fish. The trophic dynamics and feeding ecology data generated by this study will provide valuable baseline data for the coastal pelagic complex and future ecosystem studies.

Det finns många frågetecken kvar kring konsumtionsbeteende inom förhistoriska kulturer. Målet med denna studie var därför att studera och förbättra förståelsen av det arkeobotaniska materialet i ugnar gjorda för att torka sädeskorn och hur ugnarnas innehåll korrelerar med den växtbaserade matkonsumtionen, samt att försöka se potentiella samband mellan ugnarna, deponerade växtrester i gravar och de två mossliken Grauballemannen och Tollundmannens maginnehåll. De 14 ugnarna, belägna i Sydskandinavien och daterade till järnåldern, analyserades och jämfördes för att se hur deras innehåll förhåller sig både i tid och rum men även jämfört med det övriga åkerbruket från bronsåldern till medeltiden. Det arkeobotaniska fynden visade att i yngre bronsåldern odlades det Hordeum vulgare var. vulgare, vilket följdes av en period då Hordeum vulgare var. nudum odlades. Detta pågick till slutet av romersk järnålder då Hordeum vulgare var. vulgare igen blev den vanligaste grödan att odla. I slutet av järnåldern odlades både Avena sativa och Hordeum vulgare var. vulgare. Övriga sädeskorn som Triticum sp. och Secale cereale har även odlats, dock endast till en mindre grad enligt fynden från ugnarna. De arkeobotaniska fynden jämfördes sedan med den rådande bilden av matkulturen under järnåldern, vilket visar indikationer på att Chenopodium album, Fallopia convolvulus, Persicaria maculosa/lapathifolia, Plantago lanceolata, Poacea, Polygonum aviculare och Spergula arvensis troligen har konsumerats tillsammans med de funna sädeskornen. Flera av dessa påträffades i de samtliga analyserade fornlämningarna.  Jämförelseanalyser i dendrogram mellan gravarnas och ugnars makrofossila rester samt de båda mosslikens maginnehåll visade inga definitiva samband. Dock fanns det gravar som korrelerade relativt starkt med ugnarnas innehåll, möjligen på grund av dess stora fyndmaterial av makrofossil. Dessa kunde visa indikationer på mat och måltid har deponerats i gravar.

Large, mesopelagic teleosts have a potentially keystone position in the ecology of the pelagic water column, yet remain relatively unstudied when compared to large, commercially important fishes of the epipelagic zone. In this study, the ecological roles of four large, vertically migrating teleosts, oilfish, Ruvettus pretiosus, escolar, Lepidocybium flavobrunneum, snake mackerel, Gempylus serpens, longnose lancetfish, Alepisaurus ferox, and shortnose lancetfish, Alepisaurus brevirostris, were examined. The stomachs of 61 oilfish, 35 escolar, 33 snake mackerel, and 34 lancetfish were collected from nighttime, pelagic, longline fishing operations in the epipelagic zone of the western North Atlantic Ocean and Gulf of Mexico from 2007 to 2010. Stomach content analysis was used to determine predator-prey interactions. Stable carbon and nitrogen isotope analyses were performed on white dorsal muscle tissue of 38 oilfish, 23 escolar, 28 snake mackerel, and 19 lancetfish also sampled for stomach content analysis. Crustaceans were abundant prey of small oilfish, lancetfish, and to a lesser degree, snake mackerel, and they were absent from large escolar diets. Lancetfish and snake mackerel showed diverse diets that included crustaceans, polychaetes, gymnosomes, salps, cephalopods, and fishes. Large escolar stomachs contained only squid and fish. Stable isotope analyses placed small oilfish in the lowest trophic position of the group, followed by lancetfish and snake mackerel, and large escolar occupied the highest trophic position of the four species.