Cuticular compounds inhibit cannibalism of early-instar spiderlings by pulli-carrying Pardosa pseudoannulata females.

Cannibalism is common in spiders. Wolf spider (Lycosidae) females, which exhibit extensive maternal care, have been reported to cannibalize less when they are carrying egg sacs and juveniles. In a laboratory experiment, we demonstrated that cannibalism of early-instar spiderlings (EIS) by a wolf spider (Pardosa pseudoannulata) mother was almost completely inhibited when she was carrying spiderlings. Compared with virgin and mated-females, mother spiders tolerated more and predated fewer spiderlings, including gregarious pulli and newly dispersed spiderlings (NDS). Cannibalism of EIS by females during their reproductive period exhibited a V-shaped pattern, with a gradual decrease from the egg sac-carrying to pulli-carrying (PC) stage, and a recovery from the PC stage to post-reproductive (PR) stage. Notably, there was 0 cannibalism at the PC stage. PC females exhibited no interest in pulli, while PR females were attracted to and predated pulli and NDS as they did their natural prey, Nilaparvata lugens. Interestingly, PC females captured and released NDS in a foraging assay, although attraction was observed from olfactometer measurements. PC mothers possessed a cuticular volatile profile that was closer to that of pulli and NDS than to that of PR females. Moreover, NDS cuticular extract provoked an electrophysiological response in legs of PC females. Therefore, cuticular compound-mediated chemical communication may be involved in inhibiting cannibalism of EIS by spider mothers, and especially in eliminating cannibalism by PC mothers. Future studies will aim to characterize the specific cuticular compounds and chemoreception mechanism in females, which will facilitate our understanding of intraspecific recognition and cannibalism in spiders.

Diet type influences the gut microbiome and nutrient assimilation of Genetically Improved Farmed Tilapia (Oreochromis niloticus).

Nile tilapia, Oreochromis niloticus is the third most commonly farmed finfish species in the world, accounting for nearly 5% of global aquaculture production. In the past few decades much of the success of this species has been attributed to the development and distribution of Genetically Improved Farmed Tilapia (GIFT). Despite the increasing availability of GIFT, the productivity of small-scale farming remains highly variable, particularly in developing nations. Commercial fish-feed pellets can increase fish farm productivity; however, many small-scale farmers rely on other means of feeding fish due to the high cost and limited availability of commercial fish feed pellets. Therefore, understanding how locally-sourced feeds affect the production of GIFT is an important step towards improving feeding practices, particularly for farmers with low financial capital. This study used stable isotope analysis (SIA) and 16S rRNA gene sequencing to compare the effects of a locally-sourced vegetable-based diet and commercial pellet-based diets on the relative condition, nutrient assimilation patterns and gastrointestinal microbiota of GIFT. GIFT fed a locally-sourced diet were smaller, and in a significantly poorer condition than those fed with commercial fish feeds. SIA showed no differences in dietary carbon between the two diets; however, δ13C, poor fish condition and the abundance of specific bacterial taxa (of such as Fusobacteria) were correlated. SIA revealed that GIFT fed locally-sourced diets that predominantly consisted of vegetables were significantly enriched in δ15N despite a perceived lack of dietary protein. This enrichment suggests that GIFT fed a locally-sourced diet may be supplementing their diet via cannibalism, a behaviour representative of poor farming practice. Overall this study highlights the need to increase the availability of suitable GIFT feeds in developing nations. The development a low-cost feed alternative could improve the success of small-scale GIFT farmers in PNG, increasing both food and income security within the region.

Black widows as plastic wallflowers: female choosiness increases with indicators of high mate availability in a natural population.

Female choice is an important driver of sexual selection, but can be costly, particularly when choosy females risk remaining unmated or experience delays to reproduction. Thus, females should reduce choosiness when mate encounter rates are low. We asked whether choosiness is affected by social context, which may provide reliable information about the local availability of mates. This has been demonstrated in the lab, but rarely under natural conditions. We studied western black widow spiders (Latrodectus hesperus) in the field, placing experimental final-instar immature females so they were either 'isolated' or 'clustered' near naturally occurring conspecifics (≥10 m or ≤1 m, respectively, from a microhabitat occupied by at least one other female). Upon maturity, females in both treatments were visited by similar numbers of males, but clustered females were visited by males earlier and in more rapid succession than isolated females, confirming that proximity to conspecifics reduces the risk of remaining unmated. As predicted, isolated females were less choosy in staged mating trials, neither rejecting males nor engaging in pre-copulatory cannibalism, in contrast to clustered females. These results demonstrate that exposure of females to natural variation in demography in the field can alter choosiness of adults. Thus, female behaviour in response to cues of local population density can affect the intensity of sexual selection on males in the wild.

High-quality alternative food reduces cannibalism in the predatory mite Amblyseius herbicolus (Acari: Phytoseiidae).

Predatory mites of the Phytoseiidae family are important biological control agents. Many species of this family are omnivores, i.e., besides on prey, they can feed on plant resources such as nectar and pollen. It has been shown that the addition of alternative food for predators to a crop enhances biological control. However, factors such as food availability and quality can also affect interactions such as cannibalism, and thus influence biological control. We investigated the role of quality of the alternative food in the tendency of Amblyseius herbicolus to engage in cannibalism, a common ecological interaction in many phytoseiid mite species. Cannibalism on eggs by A. herbicolus was significantly reduced in the presence of high-quality food (cattail pollen) compared to egg cannibalism without alternative food, whereas this was not the case in the presence of low-quality food (cotton pollen). This suggests that cattail pollen is a high-quality alternative food, not only because it results in increased development and reproduction of predators, but also because it can minimize cannibalism.

Aggressive or funerary cannibalism? Skull-cup and human bone manipulation in Cueva de El Toro (Early Neolithic, southern Iberia).

OBJECTIVE: We analyze the processing sequence involved in the manufacture of a skull-cup and the manipulation of human bones from the Early Neolithic of Cueva de El Toro (Málaga, Spain). MATERIALS AND METHODS: The Early Neolithic material studied includes human remains found in two separate assemblages. Assemblage A consists of one skull-cup, a non-manipulated adult human mandible, and four ceramic vessels. Assemblage B contains manipulated and non-manipulated human remains that appeared mingled with domestic waste. Using a taphonomic approach, we evaluate the skull-cup processing and the anthropogenic alteration of human bones. RESULTS: The skull-cup was processed by careful paring away of skin, fragmentation of the facial skeleton and base of the skull, and controlled percussion of the edges of the calotte to achieve a regular shape. It was later boiled for some time in a container that caused pot polish in a specific area. The other human bones appeared scattered throughout the living area, mixed with other remains of domestic activity. Some of these bones show cut marks, percussion damage for marrow extraction, and tooth/chewing marks. DISCUSSION: Evidence from Cueva de El Toro suggests that cannibalism was conducted in the domestic sphere, likely following ritualized practices where the skull-cup could have played a part. Interpretation of this evidence suggests two hypotheses: (a) aggressive cannibalism relates to extreme inter-group violence; and (b) funerary cannibalism is a facet of multi-stage burial practices. Similar evidence has been found in other Neolithic sites of this region and suggests that cannibalism and skull-cups were elements widespread in these communities. These practices may be linked to significant transformations associated with the end of the Early Neolithic in southern Iberia.

Asymmetric interactions and their consequences for vital rates and dynamics: the smaller tea tortrix as a model system.

Asymmetric interactions among conspecifics can have diverse effects on population dynamics including stabilization, generation of cycles, and induction of chaotic fluctuations. A difficult challenge, however, is establishing the link between the impact of asymmetric interactions on life history and the consequences for population dynamics. The smaller tea tortrix, Adoxophyes honmai, is a good example. Larval instars differ dramatically in size and have a tendency for cannibalism, which suggests the potential for strong asymmetric interactions among instars. Yet whether these asymmetries have any role in generating the distinct single-generation cycles observed in the field and laboratory is unclear. Here we report on the development of a new experimental approach to characterize the impact of asymmetric interactions on life history that can be directly embedded into stage-structured population models. The experiments use donor-replacement protocols in which focal individuals are challenged to complete their life cycles in competitive environments where the instar and density of the competitors are held constant. The experimentally derived interaction surface contains all the information about stage-specific interactions and provides a straightforward framework for evaluating alternative ways of abstracting the interactions into traditional models of asymmetric competition. Working with the smaller tea tortrix, we found strong evidence of asymmetric interactions and identified critical "tipping points" in the competitive environment that strongly affected survival but not development. We incorporated the experimentally derived interaction surface into a stage-structured population model and found that despite the strong impact that asymmetric interactions have on tea tortrix life history, they do not scale-up to impact the predicted asymptotic population dynamics. Comparing these dynamics with two abstracted models of stage-structured interactions revealed that while the quantitative details of the emergent dynamics depends on the shape of the interaction surface, the qualitative features, such as the emergence of single-generation cycles and rapid synchronization of development among individuals, are pleasingly robust.

Death and Survival of Patients with Hansen's Disease in Colonial Korea / 의사학

The purpose of this research is to describe how Hansen's disease patients experienced the modern system of control of Hansen's disease introduced by Japan, and the inimical attitude of society against them in colonial Korea. The study also seeks to reveal the development of the system to eliminate Hansen's disease patients from their home and community to larger society and leprosarium in this era. Sorokdo Charity hospital (SCH), a hospital for Hansen's disease patients, was built in 1916, and vagrant Hansen's disease patients began to be isolated in this hospital beginning in 1917 by the Japanese Government-General of Korea (JGGK). Once the police detained and sent vagrant Hansen's disease patients to SCH, stigma and discrimination against them strengthened in Korean society. Because of strong stigma and discrimination in Korean society, Hansen's disease patients suffered from daily threats of death. First, their family members were not only afraid of the contagiousness of Hansen's disease but also the stigma and discrimination against themselves by community members. If a family had a Hansen's disease patient, the rest of community members would discriminate against the entire family. Furthermore, because Hansen's disease patients were excluded from any economic livelihood such as getting a job, the existence of the patients was a big burden for their families. Therefore, many patients left their homes and began their vagrancy. The patients who could not leave their homes committed suicide or were killed by their family members. The victims of such deaths were usually women, who were at the lower position in the family hierarchy. In the strong Confucian society in Korea, more female patients were killed by themselves than male patients. Moreover, all of patients victims in the murder were women. This shows that the stigma and discrimination against Hansen's disease patients within their families were stronger against women than men. Strong stigma and discrimination made the patients rely on superstition such as cannibalism. Patients believed that there were not any effective medicine. There were a few reports of patients who were cured, and many were treated with chaulmoogra oil in the modern Hansen's disease hospitals. Eating human flesh was known as a folk remedy for Hansen's disease. As such, patients began to kill healthy people, usually children, to eat their flesh. Increased stigma led to increased victims. Hansen's disease patients who left their homes faced many threats during their vagrancy. For survival, they established their own organizations in the late 1920's. The patients who were rejected to be hospitalized in the Western Hansen's disease hospital at Busan, Daegu, and Yeosu organized self-help organizations. The purpose of these organizations was first to secure the medicine supply of chaulmoogra oil. However, as stigma and discrimination strengthened, these organizations formed by Hansen's disease patients demanded the Japanese Government-General of Korea to send and segregate them on Sorok island. They did not know the situation of the inside of this island because news media described it as a haven for patients, and very few patients were discharged from this island to tell the truth. On this island, several hundreds of patients were killed by compulsory heavy labor, starvation, and violence. They were not treated as patients, but as something to be eliminated. Under strong suppression on this island, the patients resisted first by escaping this island. However, in 1937, some patients tried to kill a Korean staff but failed. Attempted murderers were all put in the jail, also located on this island. In 1941, a patient murdered another patient who had harassed other patients, and in 1942, Chunsang Lee, a patient, killed the director of Sorok island. These instances show that there was a system to eliminate Hansen's disease patients in colonial Korea.

The Quality of Nonprey Food Affects Cannibalism, Intraguild Predation, and Hyperpredation in Two Species of Phytoseiid Mites.

Generalist arthropod predators not only prey on herbivores but also may engage in competitive interactions by attacking and consuming conspecifics (cannibalism) or other predators (intraguild predation [IGP] and hyperpredation). These types of interactions are quite common among predators used in biological control. Although there is evidence that nonprey food relaxes cannibalism and IGP, there is little information regarding the impact of the quality of the nonprey food. Herein, we examined how pollen of different nutritional quality (pine, narrow-leaf cattail, or apple) impacted 1) the cannibalism by females of Euseius stipulatus (Athias-Henriot) (Acari: Phytoseiidae) on conspecific larvae, 2) the reciprocal predation between gravid females of E. stipulatus or Iphiseius degenerans (Berlese) (Acari: Phytoseiidae) and heterospecific larvae, and 3) the predation of E. stipulatus on the eggs of the aphid predator Aphidoletes aphidimyza (Rondani) (Diptera: Cecidomyiidae). E. stipulatus cannibalism was significantly reduced in the cattail pollen treatment, whereas in the pine pollen it did not significantly differ from control (no food). Predation between I. degenerans and E. stipulatus was significantly reduced in the cattail pollen treatment as compared to the control treatment. Finally, predation of E. stipulatus on A. aphidimyza eggs was significantly reduced when cattail or apple pollen was provided compared to the pine pollen or control treatments. These results suggest that cattail or apple pollen is suitable for mitigating negative interactions among generalist predatory mites used in biological control.

The diet of the first Europeans from Atapuerca.

Hominin dietary specialization is crucial to understanding the evolutionary changes of craniofacial biomechanics and the interaction of food processing methods' effects on teeth. However, the diet-related dental wear processes of the earliest European hominins remain unknown because most of the academic attention has focused on Neandertals. Non-occlusal dental microwear provides direct evidence of the effect of chewed food particles on tooth enamel surfaces and reflects dietary signals over time. Here, we report for the first time the direct effect of dietary abrasiveness as evidenced by the buccal microwear patterns on the teeth of the Sima del Elefante-TE9 and Gran Dolina-TD6 Atapuerca hominins (1.2-0.8 million years ago - Myr) as compared with other Lower and Middle Pleistocene populations. A unique buccal microwear pattern that is found in Homo antecessor (0.96-0.8 Myr), a well-known cannibal species, indicates dietary practices that are consistent with the consumption of hard and brittle foods. Our findings confirm that the oldest European inhabitants ingested more mechanically-demanding diets than later populations because they were confronted with harsh, fluctuating environmental conditions. Furthermore, the influence of grit-laden food suggests that a high-quality meat diet from butchering processes could have fueled evolutionary changes in brain size.

Cannibalism versus funerary defleshing and disarticulation after a period of decay: comparisons of bone modifications from four prehistoric sites.

OBJECTIVES: Humanly induced modifications on human and non-human bones from four archaeological sites of known funerary rituals (one interpreted as cannibalism and three interpreted as funerary defleshing and disarticulation after a period of decay) were analyzed to ascertain whether macromorphological and micromorphological characteristics of cut marks can be used to distinguish cannibalistic from secondary burial practices. MATERIAL AND METHODS: Four collections were analyzed: the Magdalenian assemblage from Gough's Cave (UK) and the Mesolithic-Neolithic bone samples from Lepenski Vir, Padina and Vlasac (Serbia). A total of 647 cut marks (345 on human and 302 on non-human remains) were imaged and measured using an optical surface measurement system, the Alicona InfiniteFocus, housed at the Natural History Museum (London, UK). RESULTS: The frequency of cut marks at Gough's Cave exceeds 65%, while it is below 1% in the Serbian sites, and no human tooth marks and only one case of percussion damage have been observed on the three Serbian collections. The distribution of cut marks on human bones is comparable in the four assemblages. Cannibalized human remains, however, present a uniform cut mark distribution, which can be associated with disarticulation of persistent and labile articulations, and the scalping and filleting of muscles. For secondary burials where modification occurred after a period of decay, disarticulation marks are less common and the disarticulation of labile joints is rare. The micromorphometric analyses of cut marks on human and non-human remains suggest that cut marks produced when cleaning partially decayed bodies are significantly different from cut marks produced during butchery of fresh bodies. CONCLUSIONS: A distinction between cannibalism and secondary treatment of human bodies can be made based on frequency, distribution and micromorphometric characteristics of cut marks.

Effects of dietary peppermint (Mentha piperita) essential oil on survival, growth, cannibalism and hepatopancreatic histopathology of Portunus pelagicus juveniles.

A two-part experiment was performed to determine whether dietary peppermint oil could improve the growth and/or decrease aggression among blue swimmer crab, Portunus pelagicus early juveniles. A total of five isonitrogenous diets were made that contained increasing peppermint oil levels of 0.00, 0.05, 0.10, 0.50 or 1.00%.? These diets were fed to 45 replicate crabs in each treatment (total of 225 crabs) for 12 days, the final sizes and weights were measured, and then placed in 3 replicate containers (30 in total/treatment) to allow the opportunity for cannibalism over 10 days.? After 10 days, the remaining crabs were examined for any histopathological changes in gills or hepatopancreas.? Results showed dietary peppermint oil, at the tested levels, had no effect on the growth or cannibalism, in either experiments (p > 0.05).? However, there were substantial changes in the hepatopancreatic histopathology that included thinner tubules and significantly less B- and R-cells from 0.10% dietary peppermint oil and above.? The unaffected growth or cannibalism indicate that the levels of dietary peppermint oil used were insufficient and further investigations are required, particularly on the implications to the hepatopancreatic changes. ?

Upper Palaeolithic ritualistic cannibalism at Gough's Cave (Somerset, UK): The human remains from head to toe.

A recurring theme of late Upper Palaeolithic Magdalenian human bone assemblages is the remarkable rarity of primary burials and the common occurrence of highly-fragmentary human remains mixed with occupation waste at many sites. One of the most extensive Magdalenian human bone assemblages comes from Gough's Cave, a sizeable limestone cave set in Cheddar Gorge (Somerset), UK. After its discovery in the 1880s, the site was developed as a show cave and largely emptied of sediment, at times with minimal archaeological supervision. Some of the last surviving remnants of sediment within the cave were excavated between 1986 and 1992. The excavations uncovered intensively-processed human bones intermingled with abundant butchered large mammal remains and a diverse range of flint, bone, antler, and ivory artefacts. New ultrafiltrated radiocarbon determinations demonstrate that the Upper Palaeolithic human remains were deposited over a very short period of time, possibly during a series of seasonal occupations, about 14,700 years BP (before present). The human remains have been the subject of several taphonomic studies, culminating in a detailed reanalysis of the cranial remains that showed they had been carefully modified to make skull-cups. Our present analysis of the postcrania has identified a far greater degree of human modification than recorded in earlier studies. We identify extensive evidence for defleshing, disarticulation, chewing, crushing of spongy bone, and the cracking of bones to extract marrow. The presence of human tooth marks on many of the postcranial bones provides incontrovertible evidence for cannibalism. In a wider context, the treatment of the human corpses and the manufacture and use of skull-cups at Gough Cave have parallels with other Magdalenian sites in central and western Europe. This suggests that cannibalism during the Magdalenian was part of a customary mortuary practice that combined intensive processing and consumption of the bodies with ritual use of skull-cups.

Effect of water flow rate and feed training on "pacamã" (Siluriforme: Pseudopimelodidae) juvenile production / Efeito do fluxo de água e do condicionamento alimentar na produção de juvenis de pacamã (Siluriforme: Pseudopimelodidae)

The effects of different water flow rates and feed training on the production of "pacamã" Lophiosilurus alexandri juveniles were evaluated. In the first experiment, nine day post-hatch larvae (n= 2,400) were stocked at a density of 5 larvae/L. Different water flow (F) rates were tested: F1 = 180; F2 = 600; F3 = 1,300; and F4 = 2,600mL/min. Artemia nauplii were offered as food during the first 15 days of active feeding. In the second experiment for feed training, 720 juveniles (total length of 22.2mm) were stocked at a density of 1.5 juveniles/L. A water flow rate similar to F1 was used. The use of extruded dry diet was tested, and feed training was done with and without other enhanced flavors (Artemia nauplii or Scott emulsion). The water flow rates did not influence the survival or growth of L. alexandri. Cannibalism occurred during feed training. The worst survival, specific growth rate and high mortality were found with the use of extruded dry diet, while similar values were registered with the different feed training diets used. Reduced water flow rate can be used to lower water consumption during larviculture and feed training of L. alexandri.

Avaliaram-se os efeitos de diferentes fluxos de água e do condicionamento alimentar sobre a produção de juvenis de pacamã Lophiosilurus alexandri. Larvas com nove dias pós-eclosão (n=2.400) foram estocadas na densidade de 5 larvas/L. Diferentes fluxos (F) de água foram testados: F1 = 180; F2 = 600; F3 = 1.300; e F4 = 2.600mL/min. Náuplios de Artemia foram usados como alimento nos primeiros 15 dias de alimentação ativa. No segundo experimento, para o condicionamento alimentar, 720 juvenis (comprimento total de 22,2mm) foram estocados na densidade de 1,5 juvenis/L. Fluxo de água similar a F1 foi utilizado. Foram testados o uso de dieta extrusada e o condicionamento alimentar adicionado ou não de náuplios de Artemia ou emulsão Scott. Os diferentes fluxos de água não afetaram a sobrevivência e o crescimento de L. alexandri. Durante o condicionamento alimentar foi registrada a ocorrência de canibalismo. Pior sobrevivência, taxa de crescimento específica e maior mortalidade foram registradas para o uso da dieta comercial, enquanto valores similares foram verificados para as diferentes dietas usadas no condicionamento alimentar. Fluxo de água reduzido pode ser empregado durante a larvicultura e condicionamento alimentar de L. alexandri.

[Was cannibalism practiced in ancient Gaul? Critognatus, the leader of the Celts, in the focus of Caesar's propaganda]. / Kannibalismus im alten Gallien? Der Keltenfürst Critognatus im Brennpunkt der Propaganda Caesars.

Critognatus, the leader of the Celts, is mentioned only once in the extant ancient literature, namely in Caesar's description of the siege of Alesia in BG VII 77.2-78.2. Here he is portrayed as a determined patriot who wants to encounter the Roman invader bravely and at the risk of all available means. Nevertheless, crafty Caesar succeeds in stamping him by propagandistic pinches to an evil monster and cannibal. On the one hand Caesar falls back on current Roman prejudices towards the Gauls. On the other hand, the endocannibalism practised among Celts to a certain extent as a cult action seems to have played a rôle. Caesar's propagandistic methods are transparent and at the same time so effective that the label of an ogre sticks to Critognatus until the present day. Caesar's portrayal aims above all at the justification of his Gallic War which he wages against uncivilized and inhuman opponents who are a menace to Rome and even to the culture itself.

New radiocarbon dates and isotope analysis of Neolithic human and animal bone from the Fontbrégoua Cave (Salernes, Var, France).

This article presents the results of stable carbon and nitrogen isotope analysis carried out on 12 human and 13 animal bones from the Neolithic cave of Fonbrégoua located in southern France. The stable isotope data shows that the humans had diets in which the protein sources were mainly from terrestrial animals. Six new radiocarbon dates on human bone indicate that the deposition of human remains at the site occurred during the Cardial phase (ca. 5450-5100 BCE cal.) as well as through to the transition to the middle Neolithic (ca. 5100-4800 BCE cal.). The isotopic human dietary patterns at Fontbrégoua are similar to other Neolithic sites in the South-east of France and in Liguria (Italy), indicating similar dietary adaptations in this area during this period. Radiocarbon dates on humans from the site indicate that human burial practices at this site occurred over a long time period.

Repeated evolution of male sacrifice behavior in spiders correlated with genital mutilation.

According to sexual cannibalism theory, male complicity in terminal mating can be adaptive when the male's future reproductive value is low relative to the benefits of self sacrifice. Spiders and insects that exhibit male sacrifice behavior (either complicity in cannibalism or spontaneous death associated with copulation) often also have male genitalia that stereotypically become broken or disfigured the first time they are used for copulation, potentially lowering his future reproductive value. Theoretical work on monogamy has identified male bias in the effective sex ratio as a precursor to the evolution of monogamy (including male sacrifice) as an adaptive form of paternity protection. Using phylogeny-based statistics and drawing on several phylogenetic studies of araneoid spiders, I investigate relationships between male sacrifice behavior, genital mutilation, extreme sexual size dimorphism, and the accumulation of multiple males in the female web (as an indicator of a male-based effective sex ratio). This investigation focuses on araneoid spiders because several independent origins of sacrifice behavior are known for this group and the phylogenetic structure of the lineage is relatively well studied. I report that male genital mutilation is significantly correlated with sacrifice behavior and argue that this finding is consistent with sexual cannibalism theory. Male sacrifice behavior is also correlated with male accumulation, a result that is consistent with theoretical work on the evolution of monogamy. Male accumulation and extreme sexual size dimorphism are correlated suggesting that sex-based differences in maturation time can lead to a male biased effective sex ratio. Similar patterns of correlated characters may hold for some insect taxa. Studying traits that have appeared independently in multiple lineages is a powerful method for developing general theories about the evolution of biological phenomena.