Extrapolating predation mortalities back in time: an example from North-east Arctic cod cannibalism

Cannibalism is known to be a significant source of natural mortality of young North-east Arctic (NEA) cod. Cannibalism data, starting from 1984, have been used in NEA cod stock assessments since 1995, which has led to inconsistency in the cod abundance time series from 1946 to the present. To address this inconsistency, this study estimates the cannibalism-induced mortality (M2) of NEA cod at age 3–5 for the period 1946–1983. Combined qualitative and quantitative cod stomach content data for 1984–2010 were used to make the M2 estimations for age groups 3–5 (ICES 2014), then different factors including SSB were used to examine which covariates explained variability in M2 and thus make predictions for 1946–1983. The level of cannibalism was estimated to be high in the 1950s – early1960s. VPA-based assessment was run using these estimated M2 values. As a result, numbers of cod eaten by their conspecifics in the historical period and new increased recruitment estimates at age 3 were computed. The main factors affecting cannibalism appeared to be young cod abundance, total stock biomass (TSB) of large cod, and capelin total stock biomass (which represents an alternative prey). The problems involved in using the new recruitment time series in fishery management are discussed. The methodology presented here represents a generic approach to extending predation mortalities back in time to improve historical stock estimates.     

Experimental evidence of cannibalism and prey specialization in Arctic charr,Salvelinus alpinus

Synopsis Use of a radiographic technique enabled the study of prey selection and individual specialization in Arctic charr, Salvelinus alpinus, fed with small charr and dry pellets under laboratory conditions. Both naive and experienced fish (mean weight 475 g, mean length 34.9 cm), selected the smallest individuals when offered juvenile charr (6–16 cm) as prey. The selected prey were, on average, 22% of the predator length. Cannibalism appeared to involve individual specialization, since when groups of large charr were given the opportunity to feed on juveniles one day every two weeks, the same individuals were cannibalistic throughout the sampling period of two months. When large charr were presented with alternate cycles of different food types consisting of dry pellets and fish prey, the charr exhibited a strong and consistent feeding specialization with three distinct groups being recognised: cannibals, pellet eaters and non-feeding fish.     

Cannibalism as a life boat mechanism

Under certain conditions a cannibalistic population can survive when food for the adults is too scarce to support a non-cannibalistic population. Cannibalism can have this lifeboat effect if (i) the juveniles feed on a resource inaccessible to the adults; and (ii) the adults are cannibalistic and thus incorporate indirectly the inaccessible resource. Using a simple model we conclude that the mechanism works when, at low population densities, the average yield, in terms of new offspring, due to the energy provided by one cannibalized juvenile is larger than one.     

Computer simulation study on the cyclicities ofTribolium population dynamics

Under a set of assumptions, a mathematical model was constructed to investigate the effect of cannibalistic behavior and medium renewal schedule on Tribolium population dynamics, and the results of simulation analyses were presented to show how modelling can contribute to a better understanding of experimental study. The analyses of the present model, the main concern of which is the cyclicity in Tribolium population. showed that there are two distinct factors which create cyclicity, the cannibalistic behavior in Tribolium itself, and the medium renewal schedule. Cannibalism per se does not necessarily cause cyclicity, but the combinations of cannibalistic behavior among various life stages and their relative intensities among them can cause cyclicity and can also determine the period of cycle. External factors also can generate cyclicity, but their interaction with cannibalistic behavior has a more significant effect in modifying the period of cycle. Some discrepancies between the model and experimental results were discussed.     

Diel feeding periodicity, daily ration and vertical migration of juvenile Cape hake off the west coast of South Africa

Feeding periodicity, daily ration and vertical migration of juvenile Cape hake Merluccius capensis are investigated from midwater and bottom trawl collections taken during a 42-h period between 29 February and 2 March 1992 at a fixed position off the west coast of South Africa. Feeding of 10-20 cm hake intensified during the night when they ascended into subsurface layers to prey on recruits of anchovy Engraulis capensis. Larger hake remained close to the bottom, were partially cannibalistic and exhibited no diel feeding periodicity. M. capensis appear to migrate vertically and feed asynchronously in midwater, as individuals, and not as a population, returning to the bottom when satiated. Based upon the exponential rate of decline in stomach fullness throughout the day, the evacuation rate by hake <20 cm was estimated as 0.054 h⁻¹; 90% evacuation of anchovy prey required an estimated 43 h. Using the Elliott & Persson and Eggers methods, the daily ration was estimated as 5.51 and 4.15% of wet body weight respectively. The effect of the foraging behaviour of M. capensis on the appropriateness of acoustic sampling for estimates of their abundance is discussed.     

The cannibalistic behaviour of two Gammarus species (Crustacea: Amphipoda)

This study examines the cannibalistic behaviour of the freshwater amphipods Gammarus duebeni celticus Stock & Pinkster, 1970 and G. pulex (L., 1758). In the first experiment, interactions were staged among all combinations of single adult males, single adult females, adults in the precopulatory mate-guarding phase and juveniles. Cannibalism by inter-moult individuals on newly moulted conspecifics occurred in all interaction categories in both species. Gammarus d. celticus , however, were significantly more cannibalistic than G. pulex. Cannibalism between and within sex and size categories (males > females > juveniles) was facilitated by the vulnerability of individuals at moult. Individuals of smaller size categories, however, did not cannibalize newly moulted conspecifics of larger size categories. Males were less cannibalistic on newly moulted females than on newly moulted males and juveniles and, when in the precopulatory condition, appeared to defend females from cannibalistic attacks. In a second experiment, stream conditions were simulated in the laboratory and replicated populations monitored for nine weeks. High levels of cannibalism, and the species and sex differences in cannibalism identified in the first experiment, were confirmed under these heterogeneous conditions. Cannibalism by males on their newly moulted female mating partners, termed 'reversed' sexual cannibalism, was further investigated. When males were deprived of foraging opportunities, cannibalism of precopulatory partners was significantly more frequent. The occurrence of 'reversed' sexual cannibalism is thus interpreted as a conflict between motivation to feed and motivation to mate.     

Difference in age,growth, and sexual demography of black‐spot tuskfish,Choerodon schoenleinii (Valenciennes, 1839), in two adjacent populations,Ryukyu Archipelago,southern Japan

To better understand the plasticity of life history traits in the blackspot tuskfish, Choerodon schoenleinii (Valenciennes, 1839), the characteristics of the population around the Yaeyama Islands (24°N, 124E) were examined and compared to those around Okinawajima Island (26°N, 128E) that had been investigated in a previous study. Age and growth of the Yaeyama population were examined based on 103 specimens collected at fish markets between 2006 and 2016. Specimens included 83 females (25.2–69.0 cm total length [TL]), and 20 males (43.1–71.8 cm TL). Ages determined from sectioned otoliths ranged from 1–9 for females, and 4–15 for males. Values for von Bertalanffy growth functions were L_t = 74.2 {1?exp[?0.23 (t + 0.38)]}, and the growth of the Yaeyama population was significantly faster than that of the previously studied population. Sexual demography of the two populations was compared using body length data on landings measured at the fish markets. In the Yaeyama population, females and males ranged from 24–65 cm TL and 39–75 cm TL, respectively; length at 50% individual sex change size was estimated at 54.7 ± 0.56 cm (±95% C.I.). In contrast, in the Okinawajima population, females and males ranged from 16–65 cm TL and 30–75 cm TL, respectively; meanwhile, 50% sex change size was estimated to be 50.0 ± 0.25 cm. There were thus significant differences in the size at sex change between the two populations. This difference may be related to the difference in population density between the sites.     

影响漠斑牙鲆稚鱼的自残因子的研究

漠斑牙鲆(Paralichthys lethostigma)稚鱼阶段,自残现象比较严重。本文以漠斑牙鲆稚鱼为实验对象,选取全长10.1—40.9mm的漠斑牙鲆进行试验观察。研究了密度、浑浊度、饥饿、个体大小差异对该幼鱼自残的影响程度。(1)密度、投饲量、个体大小差异对漠斑牙鲆稚鱼的自残行为影响极显著;高密度组的比低密度组自残现象严重;饱食组的自残率较1/3饱食投饲量组别的轻;个体大小差异是引起自残最重要的因子,当投喂不充足时,(10.1±0.4)—(20.3±0.3)mm组的自残现象最为严重自残死亡率为53%。(2)饥饿对自残行为影响显著,饥饿24h后自残死亡率为20%((15.0±0.4)mm组)。(3)浑浊度、饵料种类对自残行为影响不显著,在密度为20尾/L时,无浑浊度组稚鱼自残死亡率为8%,浑浊度组5%。       

Cannibalism in whitefish (Coregonus lavaretus)

Summary Cannibalism in whitefish (Coregonus lavaretus) is studied on the basis of an evolutionary model. The model predicts that individuals may be cannibalistic if they gain sufficiently in terms of fitness relative to the reduction in survival caused by this behaviour. If cannibalistic forms mainly increase survival it is predicted that cannibalism is particularly likely to be an evolutionarily optimal strategy if only a few age classes are being cannibalized and if, simultaneously, many age classes are cannibalistic. The same pattern, although less apparent, emerges on the basis of a model assuming that cannibalistic forms mainly increase fecundity.The whitefish in River Sokna spawn during late October to late November. Males dominate the spawning run numerically. Growth rate in mature stages is low, 1.0 cm·y^-1 for males and 1.3 cm·y^-1 for females. Yearly survival rate in mature stages is 0.574, and survival from egg to mature is 0.000078%. Larger whitefish on the average lay more and larger eggs than do smaller. Females were more cannibalistic than males; in total cannibalism contributed to an egg mortality of at least 8.8%. This is consistent with the derived theoretical predictions.Our theoretical and empirical analyses on whitefish are supported by the observed lack of sexual differences in the cannibalistic behaviour of brown trout (Salmo trutta) and Arctic char (Salvelinus alpinus).     

On the (dis) advantages of cannibalism

Cannibalism is an interaction between individuals that can produce counter- intuitive effects at the population level. A striking effect is that a population may persist under food conditions such that the non-cannibalistic variant is doomed to go extinct. This so-called life boat mechanism has received considerable attention. Implicitly, such studies sometimes suggest, that the life boat mechanism procures an evolutionary advantage to the cannibalistic trait. Here we compare, in the context of a size structured population model, the conditions under which the life boat mechanism works, with those that guarantee, that a cannibalistic mutant can invade successfully under the steady environmental conditions as set by a non-cannibalistic resident. We find qualitative agreement and quantitative difference. In particular, we find that a prerequisite for the life boat mechanism is, that cannibalistic mutants are successful invaders. Roughly speaking, our results show that cannibalism brings advantages to both the individuals and the population when adult food is limiting.     

Faster or slower: has growth of eastern Baltic cod changed?

Recent environmental changes have influenced the ecology and biology of eastern Baltic cod. Declining somatic condition, maturation at smaller size and restricted size distribution of the population suggest that growth rates have decreased between the early 2000s and the 2010s. Extensive age estimation problems have until now precluded testing of this hypothesis. This study presents evidence for a decrease in somatic growth rate of Baltic cod. Temporal patterns of growth, condition and maturation were analysed based on two complementary analyses: length frequency mode progression derived from DATRAS bottom trawl survey data and known-age samples, where size at age was back-calculated from daily otolith growth patterns. In the known-age samples, growth was positively related to somatic condition at capture with maturity dependent differences. Immature individuals had experienced significantly lower growth and were in lower condition at capture than mature individuals. Growth rates in the known-age samples were estimated at 9.5, 7.8 and 5.7?cm per year for age classes 1, 2 and 3 respectively. Growth between age 2 and 3 decreased significantly from 8.8?cm in the 1997 year class to 7.6?cm in the 2010 year class. While the 2001 and 2004 known-age samples were representative for the population, the 2013 sample was biased towards individuals with a higher condition and growth. Complementary length frequency analysis following the length mode of fish from age 2 to age 3 confirmed growth estimates from the early 2000s, while suggesting a 37.5% lower growth in 2013 compared with 2005.     

Cannibalism in the Andean lizard Liolaemus orientalis

Cannibalism is a recurrent behavior across the animal kingdom, with important ecological and evolutionary consequences due to its potential trade-offs on the fitness of involved individuals and demography dynamics. Cannibalism has been reported in eight species of several phylogenetic lineages within Liolaemus, a highly diverse genus of Neotropical lizards. Within the Liolaemus montanus series, two species are reported to eat juveniles in captivity. We observed a female Liolaemus orientalis, another L. montanus series member, which had ingested a juvenile of its own species in the wild. As it generally happens with other lizards, cannibalism in Liolaemus has been suggested to be performed by the larger sex, with L. chiliensis being the only other case observed until now in which the cannibalistic individual was a female. The fact that females are slightly smaller than males in L. orientalis means that our observation is an exception to previously observed trends. We suggest that there is a relationship between this behavior and the presence of a masculine trait in the cannibalistic female: developed precloacal pores. We also discuss how refuge availability may influence demography and the probability of dispersing juveniles encountering adults, increasing the chances for cannibalism to play some role in population density regulation.     

Evidence of Cannibalism in Macrolophus pygmaeus, a Natural Enemy of Whiteflies

Cannibalism and the effects of host plant, sex, time and food resources on its expression were studied for the zoophagous mirid Macrolophus pygmaeus Wagner (Hemiptera: Miridae). Cannibalistic behaviour was studied by offering 5 conspecific larvae (first instar) to newly emerging adults. Four treatments were studied: without water, with water only, with a host plant (tobacco) and with both a host plant and prey (eggs of Ephestia kuehniella). Cannibalism was observed in all treatments. In the “host plant + eggs of E. kuehniella” treatment, very few individuals displayed cannibalistic behaviour. The proportion of cannibalism was only reduced when eggs of E. kuehniella were offered. Water (free or via a host plant) was very important for both survival and feeding. The cannibalistic behaviour of M. pygmaeus should be taken into account when planning a release strategy in the context of biological control.     

Maternal cannibalistic preferences affect offspring preferences and development in Menochilus sexmaculatus Fabricius (Coleoptera: Coccinellidae): A transgenerational investigation

Cannibalism has been widely reported across taxa. However, the heritability and expression of cannibalistic traits have been least explored. The variation in the expression of cannibalism is likely to exist amongst the population affecting the propensity of cannibalism. Thus, to know whether the mother has any role in the transgenerational transmission of this trait in a ladybird beetle, Menochilus sexmaculatus, we studied the interaction between maternal and offspring prey preferences and its effect on, development duration and body weight of offspring over generations. An insignificant effect of maternal dietary history on offspring prey preference was observed across generations except for the non-cannibalistic adults who significantly preferred aphids over eggs. The long-term detrimental effect of cannibalism was found in cannibals with increased developmental duration and decreased body weight of offspring over generations. In conclusion, the results show that maternal diet did not affect the offspring preferences in M. sexmaculatus but cannibalism had a profound generational effect on the cannibalistic propensity, development and body weight of offspring across generations shows that larval dietary history and nutritional composition of prey contribute to the expression of cannibalistic behaviour across generations.     

Analysis of sibling cannibalism among pike,Esox lucius,juveniles reared under semi-natural conditions

Synopsis Sibling cannibalism in pike, Esox lucius, larvae and juveniles living in outdoor rearing ponds was studied using stomach contents analysis. For the two initial densities tested (6 and 18 larvae m^–2, equivalent to 12 and 36 larvae m^–3), cannibalism was non-existent during the larval period (13 to 35 mm total length) and was observed only during the juvenile stages. Initial density of larvae influenced both the date of first detection of cannibalistic individuals and the rate of development of cannibalism in the population. At initial stocking densities of 18 larvae m^–2 (36 larvae m^–3), cannibalism was observed from 21 days after the start of exogenous feeding (mean total length: 60 mm) onwards. At a mean total length of 100 mm and for initial stocking densities of 6 and 18 larvae m^–2, (12 and 36 larvae m^–3), the average proportions of cannibals in the populations of juveniles were 7.8% and 41.3% and the cannibals accounted for 15.5% and 65.9% of the total pike biomass, respectively. In stomachs of cannibals, young pike were the dominant prey in terms of weight. Dry weights of invertebrate-prey were lower in cannibals than in non-cannibals of similar size. Cannibalism among pike juveniles was characterized by the prey being swallowed whole and head first in the vast majority of cases. There was a strong positive correlation between predator and prey size and the mouth size of a cannibal was found to be an important constraint determining maximum victim size. The overall mean ratio of pike prey length to pike cannibal length was 66.2% and the average ratio of prey head depth to predator mouth width amounted to 87.6%. Prey size selection could be demonstrated for several length-groups of cannibals. These results are compared with the characteristics of early cannibalism in other fish species.     

Effect of the presence of predatory cues and hunger on cannibalistic behaviour of ladybird beetle,Menochilus sexmaculatus Fabricius (Coleoptera: Coccinellidae)

1. Cannibalism and its role in population dynamics, along with various factors influencing the rate of cannibalism, have been widely studied across taxa.
2. The effects of predatory cue concentrations and starvation on cannibalistic tendencies and other life-history traits in cannibalistic and non-cannibalistic adults of Menochilus sexmaculatus were investigated.
3. Egg cannibalism increased with increasing predatory cue concentration and was maximum in high predatory cue treatment; increased with starvation irrespective of their diet history.
4. Latency to cannibalise eggs was shorter for cannibalistic adults as compared to non-cannibalistic adults irrespective of their hunger status and concentration of conspecific cues.
5. Non-cannibalistic females have higher fecundity than cannibalistic females.
6. In conclusion, both cannibalistic and non-cannibalistic individuals respond differently to intrinsic (hunger) and extrinsic cues (predatory cue concentrations). Cannibalistic individuals cannibalise more than non-cannibalistic ones at low levels of starvation and predatory cues.
7. Our finding suggests that cannibalistic individuals are better at assessing the risk and removing future competitors, which might favour these individuals over non-cannibalistic ones in adverse conditions and increased competition.

     

Population dynamic theory of size-dependent cannibalism

Cannibalism is characterized by four aspects: killing victims, gaining energy from victims, size-dependent interactions and intraspecific competition. In this review of mathematical models of cannibalistic populations, we relate the predicted population dynamic consequences of cannibalism to its four defining aspects. We distinguish five classes of effects of cannibalism: (i) regulation of population size; (ii) destabilization resulting in population cycles or chaos; (iii) stabilization by damping population cycles caused by other interactions; (iv) bistability such that, depending on the initial conditions, the population converges to one of two possible stable states; and (v) modification of the population size structure. The same effects of cannibalism may be caused by different combinations of aspects of cannibalism. By contrast, the same combination of aspects may lead to different effects. For particular cannibalistic species, the consequences of cannibalism will depend on the presence and details of the four defining aspects. Empirical evidence for the emerged theory of cannibalism is discussed briefly. The implications of the described dynamic effects of cannibalism are discussed in the context of community structure, making a comparison with the community effects of intraguild predation.     

Length–weight relationships of poor cod (Trisopterus minutus Linnaeus, 1758) in the Central Aegean Sea

A total of 1527 specimens of poor cod, Trisopterus minutus, was collected by trawl fishing between April 2001 and March 2002 from the ?zmir Bay (Middle Aegean Sea). Fish size ranged from 10.6 cm total length (TL, minimum) in September to 24.8 cm TL in April. Of the sampled population 53% were males and 41% females (sex ratio female : male = 1 : 1.3). Length–weight relationships were determined for males, females and combined sexes as W = 0.007L^3.16, W = 0.009L^3.04 and W = 0.007L^3.14 respectively.     

Length‐weight relationships of four fish species from the continental shelf of Yucatán (SW Gulf of México)

This paper presents length‐weight relationships (LWR) for four fish species from the continental shelf of Yucatán, at the Sonda de Campeche (southwestern Gulf of México). Samples were taken using shrimp trawl nets (5.71 cm squared mesh size in the cod‐end) in the mid‐summer of 2016. All LWRs were significant with r² values ranged from .946 to .994 and the values of exponent b, estimated using simple linear least squares of log‐transformed weight and length data, ranged from 2.54 to 3.10 (p < .001).     

Life cycle and growth ofPotamogeton crispus L. in a shallow pond,ojaga-ike

The life cycle and growth ofPotamogeton crispus L. were studied in a shallow pond, Ojaga-ike. With respect to the shoot elongation and seed and turion formations, the life cycle of this plant in the pond could be divided into following five stages: germination, inactive growth, active growth, reproductive and dormant stages. It was suggested that the plant showed these successive stages depending mainly upon water temperature. The turions germinated on the bottom in autumn when the water temperature fell below ca. 20 C. The plant showed hardly any growth during winter (December—early March) when the temperature was below 10 C. In the spring when the bottom water temperature rose to above 10 C (mid-March), the plant started to grow again and the shoot elongated rapidly at the rate of 4.2 cm day⁻¹ until the shoot apex reached the pond surface in late April. Both the increment of node number and the internodal elongation were associated with this rapid shoot growth. On 10 May (last sampling date), the mean values of shoot length, internodal length and the number of nodes estimated for 10 predominant plants were 238.2±5.6 cm, 7.1±0.8 cm and 34.9±4.0 cm, respectively. The turion formation and flowering occurred during the period from mid-April to mid-May when the surface water temperature ranged 19 and 22 C. The dry weight of a plant reached the maximum mean value of 1180 mg on 10 May. At its peak biomass, an individual plant produced 1–10 turions (5.5 on average) of which the mean individual turion dry weight was 53.2 mg. The turion dry weight accounted for ca. 42% of the total plant biomass m⁻² at that time.