Tiger Shark Galeocerdo Cuvier Research Articles

Age and growth of the tiger shark Galeocerdo cuvier off the east coast of Australia.

Total lengths (L(T)) at age and growth rates for south-west Pacific Galeocerdo cuvier were estimated from vertebral growth-band counts of 202 sagitally sectioned centra from 112 females (71-430 cm L(T)), 79 males (72-351 cm L(T)) and 11 of unknown sex. Captive growth data were also examined to complement vertebral age estimations. The sexes combined modelled growth coefficient (k = 0.08) was smaller than previously reported for G. cuvier populations elsewhere. Split-band and narrow banding patterns were identified as potential sources of age underestimation in this species.

Repeated, long-distance migrations by a philopatric predator targeting highly contrasting ecosystems.

Long-distance movements of animals are an important driver of population spatial dynamics and determine the extent of overlap with area-focused human activities, such as fishing. Despite global concerns of declining shark populations, a major limitation in assessments of population trends or spatial management options is the lack of information on their long-term migratory behaviour. For a large marine predator, the tiger shark Galeocerdo cuvier, we show from individuals satellite-tracked for multiple years (up to 1101 days) that adult males undertake annually repeated, round-trip migrations of over 7,500 km in the northwest Atlantic. Notably, these migrations occurred between the highly disparate ecosystems of Caribbean coral reef regions in winter and high latitude oceanic areas in summer, with strong, repeated philopatry to specific overwintering insular habitat. Partial migration also occurred, with smaller, immature individuals displaying reduced migration propensity. Foraging may be a putative motivation for these oceanic migrations, with summer behaviour showing higher path tortuosity at the oceanic range extremes. The predictable migratory patterns and use of highly divergent ecosystems shown by male tiger sharks appear broadly similar to migrations seen in birds, reptiles and mammals, and highlight opportunities for dynamic spatial management and conservation measures of highly mobile sharks.

Seasonal variability of bull and tiger shark presence on the west coast of Reunion Island, western Indian Ocean

A fisheries‑independent survey using longlines and drumlines, and an acoustic telemetry study, revealed that bull sharks Carcharhinus leucas and tiger sharks Galeocerdo cuvier occur throughout the year off the west coast of Reunion Island. The research, which commenced in 2011, was conducted in response to an increase in the rate of shark attacks. Juvenile, subadult and young adult tiger sharks of 111–390 cm total length (TL) were caught in all months except July; the sex ratio was 1:1 (M:F; n = 61). All bull sharks taken, except one (183 cm TL), were mature (205–329 cm TL), with a sex ratio of 0.7:1 (M:F; n = 54), and catches occurred throughout the year except in May and August, with the highest CPUE in September. Presence/absence of a total of 46 tagged tiger sharks and 36 tagged bull sharks was monitored by means of 42 acoustic receivers distributed along the west coast of the island. Tagged tiger sharks were detected in all months, with seasonal variation between sexes. Detections of males remained low throughout the year but with a peak in winter, whereas detections of females were low in May and August only and peaked in summer. Tagged bull sharks of both sexes were more abundant in winter than in summer, with females present year round. The reasons for the apparent differences in seasonality found in longline and drumline catches compared to acoustic monitoring are discussed.

Isolation and characterization of 20 microsatellite markers from Carcharhinus leucas (bull shark) and cross-amplification in Galeocerdo cuvier (tiger shark), Carcharhinus obscurus (dusky shark) and Carcharhinus plumbeus (sandbar shark)

With the development of genetics methods, it becomes possible to study the population structure and some aspects of the reproductive behaviour of endangered sharks. Here we describe the isolation of 20 polymorphic microsatellite markers in the bull shark Carcharhinus leucas (Carcharhinidae) and their characteristics. Two to 10 alleles per locus were detected. Observed and expected heterozygosities ranged from 0.00 to 0.78 and from 0.05 to 0.80, respectively. Four markers showed deviations from Hardy–Weinberg equilibrium; among them, three showed presence of null alleles. No linkage disequilibrium was detected among any of the loci. Moreover, four, 11 and 19 of these 20 markers successfully cross-amplified in the tiger shark Galeocerdo cuvier, the sandbar shark Carcharhinus plumbeus and the dusky shark Carcharhinus obscurus, respectively.

Occurrence of sub-adult tiger sharks (Galeocerdo cuvier) in a small and very remote atoll lagoon

Occurrence of sub-adult tiger sharks (Galeocerdo cuvier) in a small and very remote atoll lagoon

Long-Term Changes in Species Composition and Relative Abundances of Sharks at a Provisioning Site

Diving with sharks, often in combination with food baiting/provisioning, has become an important product of today’s recreational dive industry. Whereas the effects baiting/provisioning has on the behaviour and abundance of individual shark species are starting to become known, there is an almost complete lack of equivalent data from multi-species shark diving sites. In this study, changes in species composition and relative abundances were determined at the Shark Reef Marine Reserve, a multi-species shark feeding site in Fiji. Using direct observation sampling methods, eight species of sharks (bull shark Carcharhinus leucas, grey reef shark Carcharhinus amblyrhynchos, whitetip reef shark Triaenodon obesus, blacktip reef shark Carcharhinus melanopterus, tawny nurse shark Nebrius ferrugineus, silvertip shark Carcharhinus albimarginatus, sicklefin lemon shark Negaprion acutidens, and tiger shark Galeocerdo cuvier) displayed inter-annual site fidelity between 2003 and 2012. Encounter rates and/or relative abundances of some species changed over time, overall resulting in more individuals (mostly C. leucas) of fewer species being encountered on average on shark feeding dives at the end of the study period. Differences in shark community composition between the years 2004–2006 and 2007–2012 were evident, mostly because N. ferrugineus, C. albimarginatus and N. acutidens were much more abundant in 2004–2006 and very rare in the period of 2007–2012. Two explanations are offered for the observed changes in relative abundances over time, namely inter-specific interactions and operator-specific feeding protocols. Both, possibly in combination, are suggested to be important determinants of species composition and encounter rates, and relative abundances at this shark provisioning site in Fiji. This study, which includes the most species from a spatially confined shark provisioning site to date, suggests that long-term provisioning may result in competitive exclusion among shark species.

A revised estimate of daily ration in the tiger shark with implication for assessing ecosystem impacts of apex predators

NOAA, National Marine Fisheries Service, Southeast FisheriesScience Center, 3500 Delwood Beach Road, Panama City, Florida 32408, USAApex predators may impact ecosystem structure and func-tion through trophic cascades (Estes, Tinker & Williams1998). Studies of diets, feeding patterns and food webdynamics can contribute to an understanding of commu-nity structure and ecological interactions (Winemiller 1989;Krebs 1998). Thus, understanding trophodynamics isimportant for developing ecosystem models for predictingcommunity responses to anthropogenic changes (Walters,Christensen & Pauly 1997). Studies of this kind are espe-cially critical in the case of large sharks, given that severalspecies are experiencing varying levels of populationdeclines on a global scale (Dulvy et al. 2008; Camhi et al.2009). Accordingly, there is mounting concern – andincreasing effort – to predict the consequences of largeshark declines for food web dynamics (Estes et al. 2011).In a recent study, Hammerschlag et al. (2012) investi-gated the movement patterns of an apex marine predator,the tiger shark (Galeocerdo cuvier), in relation to an eco-tourism provisioning site. The authors rejected the nullhypothesis that sharks displayed restricted long-term habi-tat use at the site. However, they speculated that theobserved long-term and large-scale shark movements maybe related to cryptic, lesser-understood aspects of Atlantictiger shark life history (i.e. reproduction, mating, foragingforays).As part of their discussion, Hammerschlag et al. (2012)considered the daily ration of 210-kg adult female tigersharks. However, no published studies exist that havequantified this aspect of tiger shark food-consumptiveneeds. To get a broad approximation of daily ration, theauthors averaged values derived from two other species ofsharks found in the subtropical Atlantic: the lemon shark(Negaprion brevirostris), daily ration estimate of 2 1%body weight (Cortes & Gruber 1990); and shortfin makoshark (Isurus oxyrinchus), daily ration estimate of 4 6%body weight (Wood et al. 2009). The species used in thecalculation here were chosen in part because of theirbehavioural and geospatial overlap with the tiger sharkmovements found in Hammerschlag et al. (2012). How-ever, the resulting approximation of the daily ration valuefor an adult tiger shark (3 7%) is likely inflated for twoprimary reasons: (i) using values derived from juvenile spe-cies, although daily ration in sharks is known to decreasewith size (Wetherbee & Cortes 2004) and (ii) averaging val-ues from two species, one of which, the shortfin mako, isregionally endothermic, while tiger sharks are ectothermic(Carlson, Goldman & Lowe 2004).Here we present a different, more traditional approachto estimate the daily ration for tiger sharks followingWinberg (1956). Accordingly, daily ration (kcal day

Complete mitochondrial genome of the tiger shark Galeocerdo cuvier (Carcharhiniformes: Carcharhinidae)

The tiger shark Galeocerdo cuvier is the only member of the genus Galeocerdo. The complete mitochondrial genome of G. cuvier is presented for the first time in this study. The gene composition and arrangement in the mitogenome of G. cuvier is identical to most animal mitogenome. There are 22 bp short noncoding sequences and 44 bp overlaps in the mitogenome. The overall base composition is 31.8% A, 23.9% C, 13.0% G and 31.3% T. The dihydrouridine arm of tRNA-Ser2 was replaced by a simple loop and the other tRNAs could be folded into the typical cloverleaf structure.

Apparent resource partitioning and trophic structure of large-bodied marine predators in a relatively pristine seagrass ecosystem

Large predators often play important roles in structuring marine communities. To understand the role that these predators play in ecosystems, it is crucial to have knowledge of their interactions and the degree to which their trophic roles are complementary or redundant among species. We used stable isotope analysis to examine the isotopic niche overlap of dolphins Tursiops cf. aduncus, large sharks (>1.5 m total length), and smaller elasmobranchs (sharks and batoids) in the relatively pristine seagrass community of Shark Bay, Australia. Dolphins and large sharks differed in their mean isotopic values for ! 13 C and ! 15 N, and each group occupied a rela- tively unique area in isotopic niche space. The standard ellipse areas (SEAc; based on bivariate standard deviations) of dolphins, large sharks, small sharks, and rays did not overlap. Tiger sharks Galeocerdo cuvier had the highest ! 15 N values, although the mean ! 13 C and ! 15 N values of pigeye sharks Carcharhinus amboinensis were similar. Other large sharks (e.g. sicklefin lemon sharks Negaprion acutidens and sandbar sharks Carcharhinus plumbeus) and dolphins appeared to feed at slightly lower trophic levels than tiger sharks. In this seagrass-dominated ecosystem, seagrass- derived carbon appears to be more important for elasmobranchs than it is for dolphins. Habitat use patterns did not correlate well with the sources of productivity supporting diets, suggesting that habitat use patterns may not necessarily be reflective of the resource pools supporting a popula- tion and highlights the importance of detailed datasets on trophic interactions for elucidating the ecological roles of predators.

Gulf‐Wide Decreases in the Size of Large Coastal Sharks Documented by Generations of Fishermen

AbstractLarge sharks are top predators in most coastal and marine ecosystems throughout the world, and evidence of their reduced prominence in marine ecosystems has been a serious concern for fisheries and ecosystem management. Unfortunately, quantitative data to document the extent, timing, and consequences of changes in shark populations are scarce, thwarting examination of long‐term (decadal, century) trends, and reconstructions based on incomplete data sets have been the subject of debate. Absence of quantitative descriptors of past ecological conditions is a generic problem facing many fields of science but is particularly troublesome for fisheries scientists who must develop specific targets for restoration. We were able to use quantitative measurements of shark sizes collected annually and independently of any scientific survey by thousands of recreational fishermen over the last century to document decreases in the size of large sharks from the northern Gulf of Mexico. Based on records from fishing rodeos in three U.S. coastal states, the size (weight or length) of large sharks captured by fishermen decreased by 50–70% during the 20 years after the 1980s. The pattern is largely driven by reductions in the occurrence and sizes of Tiger Sharks Galeocerdo cuvier and Bull Sharks Carcharhinus leucas and to a lesser extent Hammerheads Sphyrna spp. This decrease occurred despite increasing fishing effort and advances in technology, but it is coincident with the capitalization of the U.S. commercial shark long‐line fishery in the GOM.Received April 10, 2012; accepted March 7, 2013

A Comparison of the Seasonal Movements of Tiger Sharks and Green Turtles Provides Insight into Their Predator-Prey Relationship

During the reproductive season, sea turtles use a restricted area in the vicinity of their nesting beaches, making them vulnerable to predation. At Raine Island (Australia), the highest density green turtle Chelonia mydas rookery in the world, tiger sharks Galeocerdo cuvier have been observed to feed on green turtles, and it has been suggested that they may specialise on such air-breathing prey. However there is little information with which to examine this hypothesis. We compared the spatial and temporal components of movement behaviour of these two potentially interacting species in order to provide insight into the predator-prey relationship. Specifically, we tested the hypothesis that tiger shark movements are more concentrated at Raine Island during the green turtle nesting season than outside the turtle nesting season when turtles are not concentrated at Raine Island. Turtles showed area-restricted search behaviour around Raine Island for ∼3–4 months during the nesting period (November–February). This was followed by direct movement (transit) to putative foraging grounds mostly in the Torres Straight where they switched to area-restricted search mode again, and remained resident for the remainder of the deployment (53–304 days). In contrast, tiger sharks displayed high spatial and temporal variation in movement behaviour which was not closely linked to the movement behaviour of green turtles or recognised turtle foraging grounds. On average, tiger sharks were concentrated around Raine Island throughout the year. While information on diet is required to determine whether tiger sharks are turtle specialists our results support the hypothesis that they target this predictable and plentiful prey during turtle nesting season, but they might not focus on this less predictable food source outside the nesting season.

Extraordinary growth in tiger sharks Galeocerdo cuvier from the South Atlantic Ocean

Two tagged-and-recaptured tiger sharks Galeocerdo cuvier, measuring 172 and 304 cm total length (L(T) ) and at age 0.75 and 3.50 years, exhibited unmatched growth rates of 118.4 and 55.5 cm year(-1), respectively. The larger fish was nearly mature, indicating that G. cuvier off Brazil could mature considerably earlier than conspecifics from other regions.

Relative abundance and size of coastal sharks derived from commercial shark longline catch and effort data

In the north-west Atlantic Ocean, stock assessments conducted for some commercially harvested coastal sharks indicate declines from 64 to 80% with respect to virgin population levels. While the status of commercially important species is available, abundance trend information for other coastal shark species in the north-west Atlantic Ocean are unavailable. Using a generalized linear modelling (GLM) approach, a relative abundance index was derived from 1994 to 2009 using observer data collected in a commercial bottom longline fishery. Trends in abundance and average size were estimated for bull shark Carcharhinus leucas, spinner shark Carcharhinus brevipinna, tiger shark Galeocerdo cuvier and lemon shark Negaprion brevirostris. Increases in relative abundance for all shark species ranged from 14% for C. brevipinna, 12% for C. leucas, 6% for N. brevirostris and 3% for G. cuvier. There was no significant change in the size at capture over the time period considered for all species. While the status of shark populations should not be based exclusively on abundance trend information, but ultimately on stock assessment models, results from this study provide some cause for optimism on the status of these coastal shark species.

Shark scavenging and predation on sea turtles in northeastern Brazil

Large sharks have the potential to help structure ecosystem dynamics through top-down impacts on their prey, including sea turtles. Studies of interactions between large sharks and sea turtles, however, are practically nonexistent along the Brazilian coast. Between September 2002 and May 2011 we examined 655 sea turtles – including green turtlesChelonia mydas(n = 607), olive ridleysLepidochelys olivacea(n = 10), hawksbillsEretmochelys imbricata(n = 33), and loggerheadsCaretta caretta(n = 5) – that stranded on Paraíba coast, northeastern Brazil. A total of 63 green turtles (10.4%), two olive ridleys (20.0%) and one hawksbill (3.0%) had shark-inflicted bites. Most bites could not be definitively attributed to scavenging or attacks on living turtles, but the presence of healed shark bites and freshly bleeding bites suggests that some attacks occurred pre-mortem. Bite characteristics suggest that tiger sharksGaleocerdo cuvierwere responsible for most bites that could be identified to a particular species. Within green turtles, the only species with sufficient sample size, the probability of carcasses having been bitten increased with carapace length but did not vary across seasons and years. However, there was spatial variation in the probability of a carcass having been bitten by sharks. Our estimates of the minimum proportion of turtles attacked while alive (∼4%) and bitten overall are similar to other areas where shark-turtle interactions have been studied. Turtles likely are an important food for tiger sharks in northeastern Brazil, but further studies are needed to determine the relative frequencies of scavenging and predation.

Behavioural transition probabilities in dugongs change with habitat and predator presence: implications for sirenian conservation

Although not widely appreciated, prey can manage predation risk by modifying the sequence of their behavioural states. We explored this phenomenon in dugongs (Dugong dugon) subject to spatially and temporally variable risk of tiger shark (Galeocerdo cuvier) predation in Shark Bay, Australia. Dugong behaviour was assayed using focal follows and organised into sequences of foraging, resting, and travelling bouts. We used log-linear analysis to test for sequence differences in relation to habitat (deep, shallow) and predation danger (sharks present, largely absent). Dugongs modified their behavioural sequences between periods of high and low shark abundance: those at risk alternated more frequently between foraging, which constrains anti-predator vigilance, and travelling, which facilitates predator detection. Dugongs also avoided continuous series of resting bouts, during which awareness is reduced, when sharks were present. These changes were only observed in relatively dangerous shallow habitat, which is hunted disproportionately by tiger sharks; behavioural responses to sharks in deep habitat were modest. We conclude that dugongs in risky habitat resort to safer behavioural sequences in response to sharks. Given that human disturbance and predators are perceived similarly by many species, some forms of vessel interaction could compromise the fitness of sirenians by eliciting similar behavioural adjustment.

Yo-yo vertical movements suggest a foraging strategy for tiger sharks Galeocerdo cuvier

Many pelagic fishes exhibit ‘yo-yo’ diving behavior, which may serve several possible functions, including energy conservation, prey searching and navigation. We deployed accelerometers and digital still cameras on 4 free-ranging tiger sharks Galeocerdo cuvier, to test whether their vertical movements are most consistent with energy conservation or prey searching. All sharks swam continuously, with frequent vertical movements through the water column at mean swimming speeds of 0.5 to 0.9 m s–1. Tail-beating was continuous except for sporadic, powerless gliding during descents (from 0 to 18% of total descent time). At a given tailbeat frequency, swimming speeds were lower during ascent than descent (consistent with negative buoyancy). Burst swimming events, which might represent prey pursuits, were observed during all phases of vertical movements. Camera images showed a variety of potential prey and the possible capture of a unicornfish. Collectively, results suggest that yo-yo diving by tiger sharks is not primarily for energy conservation, but probably represents an effective search strategy for locating prey throughout the water column.

The frequency and effect of shark-inflicted bite injuries to the reef manta ray Manta alfredi

Shark bite injuries on reef manta rays Manta alfredi off the coast of Inhambane, Mozambique, were examined over a three-year period (2003–2006). The frequency and seasonality of attacks, the rate of wound healing, and the possible identities of attackers were explored. This study presents the first examination of bite wounds on manta rays in the wild and the role sharks may play in the natural mortality of this species. The reported incidence of shark-inflicted injuries is high with over three-quarters of the sampled population affected. In total, 571 bite injuries were observed on 283 identified individuals. The number of bite injuries varied from one to seven, with a mean of 1.54 bite wounds. There was no significant difference in the frequency of bite injuries in male and female rays. The majority (96%) of the observed bite wounds were healed. Fresh wounds occurred throughout the year, with no obvious seasonality. The bull shark Carcharhinus leucas and tiger shark Galeocerdo cuvier are suggested as the primary mediators of attacks, although up to 11 other shark species are listed as potential attackers. The majority of the bite marks (96%) occurred to the most posterior region of the body, specifically the posterior edges of the pectoral and pelvic fins, with many injuries likely having a negative impact on the reproductive abilities and fitness of the rays.

Predator-induced modifications to diving behavior vary with foraging mode

Breath-hold divers are strongly interacting species whose top – down infl uence on aquatic communities is shaped by factors governing their diving decisions. Although some of these factors (e.g. physiological constraints, energetic needs) have been scrutinized, the possibility that predation risk infl uences diving behavior has been largely overlooked, and no study to date has asked if anti-predator responses by divers depend on foraging mode. We contrasted dive cycle changes by herbivorous dugongs Dugong dugon using two foraging tactics – cropping, which always permits anti-predator vigilance, and excavation, which limits surveillance at depth – in response to temporal variation in tiger shark Galeocerdo cuvier abundance. Dugongs responded to increasing shark abundance (one component of predation risk) by diving more frequently without changing their surface times and thereby spending a greater proportion of time at the surface, but only while excavating. When threatened, in other words, excavating dugongs sacrifi ced foraging time at depth to facilitate shark detection. In contrast, cropping dugongs at risk from sharks were able to continue diving and foraging normally. By implication, future studies should consider the infl uence of predation risk on diving decisions, even by large-bodied species, and the possibility that behavioral responses by divers to predators may vary with foraging mode.

Socio-economic aspects of the tiger shark diving industry within the Aliwal Shoal Marine Protected Area, South Africa

Understanding socio-economic aspects of the tiger shark Galeocerdo cuvier diving industry, including information on participant expectations, experiences and expenditure, is necessary for the effective management of the Aliwal Shoal Marine Protected Area on the east coast of South Africa. Between January and December 2007, a total of 2 133 tiger shark dives was conducted by 1 065 divers (95% CI = 946-1 198). Data were collected by means of the administration of a semi-structured survey questionnaire to 197 (18.6%) dive participants. Respondents indicated that the direct value of tiger shark diving to the Aliwal Shoal region was R12 405 274 (95% CI = R10 777 324-14 228 541). A total of 1 000 Monte Carlo simulations was used to estimate confidence intervals. On a ranking from one (poor) to five (excellent), the average participant response to overall quality of dive and standard of dive operator was 4.6 and 4.7 respectively. The majority of divers (98.0%) observed a tiger shark, at an average of four per dive. Although tiger sharks approached to an average distance of 1.6 m from divers, the majority (95.9%) felt safe and enjoyed the experience. The majority of interviewees (88.5%) supported the use of chumming for a closer ‘tiger shark experience’.

Olive-headed sea snakes Disteria major shift seagrass microhabitats to avoid shark predation

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 387:287-293 (2009) - DOI: https://doi.org/10.3354/meps08127 Olive-headed sea snakes Disteria major shift seagrass microhabitats to avoid shark predation Aaron J. Wirsing1,2,*, Michael R. Heithaus1 1Department of Biological Sciences, Marine Biology Program, Florida International University, Biscayne Bay Campus MSB, 3000 NE 151 Street, North Miami, Florida 33181, USA 2Present address: School of Forest Resources, University of Washington, Box 352100, Seattle, Washington 98195, USA *Email: wirsinga@u.washington.edu ABSTRACT: Predators elicit prey spatial shifts that may influence prey resources. The nature of these indirect effects is difficult to predict, however, in part because the means by which prey differentiate safe from dangerous space are poorly understood. Prey often avoid their predators, but they may instead favor predator-rich areas that facilitate escape, or discourage attack, when a predator is encountered. We investigated how olive-headed sea snakes Disteria major index their risk of tiger shark Galeocerdo cuvier predation over seagrass bank microhabitats (edges, interiors) in Shark Bay, Australia. D. major is equally likely to escape sharks in both microhabitats, so we expected to observe avoidance of predator-rich space. Supporting our prediction, snakes used microhabitats roughly in proportion to food supply when sharks were scarce and avoided edges, which are preferred by sharks, when sharks were abundant. Thus, D. major appears to measure danger across seagrass banks using variability in predator density and to seek low-encounter microhabitats when anti-predator investment is needed. Our results suggest that the influence of predators on sea snakes is underappreciated and, in the context of previous work, that sympatric prey species sharing predators may show opposite spatial shifts when threatened, potentially leading to different predator indirect effects. KEY WORDS: Anti-predator behavior · Predation risk · Predator indirect effects · Prey escape tactics · Tiger shark Full text in pdf format PreviousNextCite this article as: Wirsing AJ, Heithaus MR (2009) Olive-headed sea snakes Disteria major shift seagrass microhabitats to avoid shark predation. Mar Ecol Prog Ser 387:287-293. https://doi.org/10.3354/meps08127Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 387. Online publication date: July 28, 2009 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2009 Inter-Research.