Coevolutionary stability of host-symbiont systems with mixed-mode transmission.

The coevolution of hosts and symbionts based on virulence and mode of transmission is a complex and diverse biological phenomenon. We introduced a conceptual model to study the stable coexistence and coevolution of an obligate symbiont (mutualist or parasite) with mixed-mode transmission and its host. Using an age-structured Leslie model for the host, we demonstrated how the obligate symbiont could modify the host's life history traits (survival and fecundity) and the long-term growth rate of the infected lineage. When the symbiont is vertically transmitted, we found that the host and its symbiont could maximize the infected lineage's evolutionary success (multi-level selection). Our model showed that symbionts' effect on host longevity and reproduction might differ, even be opposing, and their net effect might often be counterintuitive. The evolutionary stability of the ecologically stable coexistence was analyzed in the framework of coevolutionary dynamics. Moreover, we found conditions for the ecological and evolutionary stability of the resident host-symbiont pair, which does not allow invasion by rare mutants (each mutant dies out by ecological selection). We concluded that, within the context of our simplified model conditions, a host-symbiont system with mixed-mode transmission is evolutionarily stable unconditionally only if the host can maximize the Malthusian parameters of the infected and non-infected lineages using the same strategy. Finally, we performed a game-theoretical analysis of our selection situation and compared two stability definitions.

Multivariate study of lice (Insecta: Psocodea: Phthiraptera) assemblages hosted by hummingbirds (Aves: Trochilidae).

Lice were collected from 579 hummingbirds, representing 49 species, in 19 locations in Brazil, Costa Rica, Honduras, Paraguay and Peru, at elevations 0­3000 m above sea level. The following variables were included in an ecological analysis (1) host species' mean body mass, sexual size dimorphism, sexual dichromatism, migratory behaviour and dominance behaviour; (2) mean elevation, mean and predictability of temperature, mean and predictability of precipitation of the host species' geographic area; (3) prevalence and mean abundance of species of lice as measures of infestation. Ordination methods were applied to evaluate data structure. Since the traits are expressed at different scales (nominal, interval and ratio), a principal component analysis based on d-correlations for the traits and a principal coordinates analysis based on the Gower index for species were applied. Lice or louse eggs were found on 80 (13.8%) birds of 22 species. A total of 267 lice of 4 genera, Trochiloecetes, Trochiliphagus, Myrsidea and Leremenopon, were collected, with a total mean intensity of 4.6. There were positive interactions between migration behaviour and infestation indices, with elevational migrants having a higher prevalence and abundance of lice than resident birds. Further, we found weak negative correlations between host body mass and infestation indices and positive correlations between mean elevation and prevalence and abundance of Trochiliphagus. Thus, formerly unknown differences in the ecological characteristics and infestation measures of Trochiliphagus and Trochiloecetes lice were revealed, which allows a better understanding of these associations and their potential impacts on hummingbirds.

Definitions of parasitism, considering its potentially opposing effects at different levels of hierarchical organization.

An annotated synthesis of textbook definitions of parasitism is presented. Most definitions declare parasitism is a long-lasting relationship between individuals of different species harming the hosts. The infection-induced costs are interpreted as diseases in the medical-veterinary literature. Alternatively, evolutionary ecologists interpret it as a reduction of host's fitness (longevity, fertility or both). Authors often assume that such effects decrease host population growth and select for antiparasitic defences, which is not necessarily true because infections may simultaneously express opposite effects at different levels of biological organization. (i) At the cellular level, infection-induced cell growth, longevity and multiplication may yield tumours maladaptive at higher levels. (ii) At the individual level, reduced host longevity, fertility or both are interpreted as disease symptoms or reduced fitness. (iii) Contrary to common sense, the growth rate of infected host lineages may increase in parallel with the individuals' reduced survival and fertility. This is because selection favours not only the production of more offspring but also their faster production. (iv) Finally, infections that reduce host individuals' or lineages' fitness may still increase infected host populations' growth rate in the context of ecological competition. Therefore, differences between parasitism and mutualism may depend on which level of organization one focuses on.

Relationship between body size and sexual size dimorphism in syringophilid quill mites.

A positive relationship of body size and sexual size dimorphism (males' size relative to females), called Rensch's rule, is often observed in comparisons within non-parasitic taxa. However, this allometric relationship has rarely been tested in comparisons across closely related parasite species. Since male sexual rivalry is often regarded as the main cause of this phenomenon, the present study tests this rule in a taxon where sexual selection is almost totally absent in males. Body size data of (non-physogastric) female and male quill mites (Acari: Syringophilidae) were gathered from the literature to investigate this relationship. The data set consisted of 113 species representing 8 genera. For the data set as a whole, increasing body size came together with decreasing relative body size of males (relative to females), a phenomenon known as converse Rensch's rule. Repeating the same analysis for the 8 genera separately, similar patterns were found in 4 significant and 3 non-significant cases. There was a significant tendency to comply with Rensch's rule only in one genus, the Neoaulonastus. Thus, converse Rensch's rule is the primary trend in syringophilid quill mites that appears repeatedly and independently in several genera. This phenomenon is probably caused by their extreme inbreeding, which strongly reduces sexual competition among males in this taxon.

Sex-dependent changes in the louse abundance of red-footed falcons (Falco vespertinus).

Permanent ectoparasites live in stable environments; thus, their population dynamics are mostly adapted to changes in the host life cycle. We aimed to investigate how static and dynamic traits of red-footed falcons interplay with the dynamics of their louse subpopulations during breeding and how they affect the colonisation of new hosts by lice. We sampled red-footed falcon (Falco vespertinus) nestlings (two breeding seasons) and adults (one breeding season) in southern Hungary. The mean abundance of Colpocephalum subzerafae and Degeeriella rufa lice on the nestlings was modelled with generalized linear mixed models using clutch size and host sex in interaction with wing length. For adults, we used wing length and the number of days after laying the first egg, both in interaction with sex. D. rufa abundances increased with the nestlings' wing length. In one year, this trend was steeper on females. In adult birds, both louse species exhibited higher abundances on females at the beginning, but it decreased subsequently through the breeding season. Contrarily, abundances were constantly low on adult males. Apparently, D. rufa postpones transmission until nestlings develop juvenile plumage and choose the more feathered individual among siblings. The sexual difference in the observed abundance could either be caused by the different plumage, or by the females' preference for less parasitized males. Moreover, females likely have more time to preen during the incubation period, lowering their louse burdens. Thus, sex-biased infestation levels likely arise due to parasite preferences in the nestlings and host behavioural processes in the adult falcons.

Sex interacts with age-dependent change in the abundance of lice-infesting Amur Falcons (Falco amurensis).

Sex-biassed and age-biassed parasite infections are common in nature, including ectoparasites-vertebrate host systems. We investigated the effect of Amur Falcons' sex, age and body size on the abundance of their lice at a migratory stopover site, where the falcons' habitat use and behaviour are more homogeneous across sex and age categories than during the breeding season. We sampled Amur Falcons in Nagaland, India at major roosting sites in 2016. We applied generalized linear models (with negative binomial distribution and log-link) to model the abundance of their two most numerous lice (Colpocephalum subzerafae and Degeeriella rufa) using the host age category (juvenile or adult) and wing length, both in interaction with sex, as explanatory variables. The abundance of C. subzerafae was only affected by host age, being nearly four times higher on juveniles than on adults. Juveniles were also more infested with D. rufa than the adults. Additionally, the abundance of the latter species was lower on adult male Falcons as compared to adult females. A juvenile bias in ectoparasite infestations is common in nature, probably due to juveniles being immunologically naïve, more resource-limited and may be inexperienced in body maintenance behaviours like preening and grooming. On the other hand, female-biassed infestations are much rarer than male-biassed infestations. We briefly discuss the possible causes of female-biassed infestations on Amur Falcons reported here, and in the closely related Red-footed Falcon and Lesser Kestrel as reported in the literature.

Do small samples underestimate mean abundance? It depends on what type of bias we consider.

Former authors claimed that, due to parasites' aggregated distribution, small samples underestimate the true population mean abundance. Here we show that this claim is false or true, depending on what is meant by 'underestimate' or, mathematically speaking, how we define 'bias'. The 'how often' and 'on average' views lead to different conclusions because sample mean abundance itself exhibits an aggregated distribution: most often it falls slightly below the true population mean, while sometimes greatly exceeds it. Since the several small negative deviations are compensated by a few greater positive ones, the average of sample means approximates the true population mean.

Co-infections in Visceral Pentastomiasis, Democratic Republic of the Congo.

Snakeborne Armillifer pentastomiasis is an emerging human parasitic infection in rural tropical areas where snake meat is eaten. After a series of severe ocular A. grandis larval infections and anecdotal abdominal infection in Sankuru District, Democratic Republic of the Congo, during 2014-2015, we systematically investigated possible pentastomid etiology in patients who underwent surgery in the region. Histologic and molecular analyses by established pentastomid 18S rDNA- and newly developed Armillifer-specific cytochrome oxidase PCRs revealed larval pentastomid lesions in 3.7% of patients. Some persons had A. armillatus and A. grandis co-infections. Another pentastomid larva, Raillietiella sp., was molecularly detected in 1 patient who had concomitant A. grandis and A. armillatus infection. The PCRs used were suitable for detecting pentastomid species even in highly necrotic tissues. Phylogenetic analyses of Armillifer cytochrome oxidase genes detected multiple local strains.

Distribution of the myrmecoparasitic fungus Rickia wasmannii (Ascomycota: Laboulbeniales) across colonies, individuals, and body parts of Myrmica scabrinodis.

The ant species Myrmica scabrinodis plays a markedly important ecological role through much of the humid grasslands of Eurasia. It hosts a species-rich community of pathogens and parasites, including Rickia wasmannii, an enigmatic member of entomoparasitic laboulbenialean fungi. This study provides a descriptive ecology of R. wasmannii by characterizing its prevalence and distribution across several hierarchical levels: colonies, individuals, and anatomic body parts. Infections were restricted to a single ant species, M. scabrinodis, and infected colonies occurred predominantly in wet habitats. Infections tended to be highly prevalent within infected colonies, often reaching 100% sample prevalence among workers. Individual infections exhibited an aggregated distribution typical to host-parasite systems. Workers from the aboveground part of nests (presumably older ones acting as foragers) were more infected than those from the belowground part. Fungal thalli could be found all over the body of the hosts, the head and the abdomen being the most infected parts of the body. The fungi's distribution among host body parts statistically differed between low versus high-intensity infections: the initial dominance of the head decreased with advancing infection. These findings may provide baseline data for future comparative or monitoring studies.

Molecular Diagnosis of Abdominal Armillifer grandis Pentastomiasis in the Democratic Republic of Congo.

Pentastomiasis is an emerging snake-borne parasitic zoonosis in the tropics. We describe a molecular and morphological study to diagnose a cluster of asymptomatic abdominal human infections caused by Armillifer grandis. The findings may indicate a silent epidemic in a rural area where severe symptomatic ocular cases with the same parasite species have recently surfaced. Molecular diagnostics are of increasing importance when patient material from remote areas cannot be thoroughly examined locally for logistic reasons.

Why infest the loved ones--inherent human behaviour indicates former mutualism with head lice.

Head lice transmit to new hosts when people lean their heads together. Humans frequently touch their heads to express friendship or love, while this behaviour is absent in apes. We hypothesize that this behaviour was adaptive because it enabled people to acquire head lice infestations as early as possible to provoke an immune response effective against both head lice and body lice throughout the subsequent periods of their life. This cross-immunity could provide some defence against the body-louse-borne lethal diseases like epidemic typhus, trench fever, relapsing fever and the classical plague. Thus the human 'touching heads' behaviour probably acts as an inherent and unconscious 'vaccination' against body lice to reduce the threat exposed by the pathogens they may transmit. Recently, the eradication of body-louse-borne diseases rendered the transmission of head lice a maladaptive, though still widespread, behaviour in developed societies.

Evolutionary co-variation of host and parasite diversity-the first test of Eichler's rule using parasitic lice (Insecta: Phthiraptera).

The taxonomic richness of lice (Phthiraptera) varies considerably among their avian and mammalian hosts. Previous studies explored some factors shaping louse diversity; however, the so-called Eichler's rule-according to which taxonomic richness of parasites co-varies with that of their hosts-has never been tested. Our study incorporates all families of birds and mammals and the whole order of lice to test this co-variation, thus we present the widest taxonomic range to test any correlates of louse richness. Louse richness data were controlled for uneven sampling effort. We used the method of independent contrasts to control for phylogenetic effects. We found a strong correlation between the species richness of avian and mammalian families and generic richness of their lice. We discuss some alternative macroevolutionary and macroecological hypotheses that may explain this phenomenon that may well be a general feature of parasitism and it seems possible that this effect contribute considerably to global biodiversity.

Ectoparasites, uropygial glands and hatching success in birds.

The uropygial gland of birds secretes wax that is applied to the plumage, where the secretions are hypothesized to eliminate fungi and bacteria, thereby potentially providing important benefits in terms of plumage maintenance. We analyzed variation in size of the uropygial gland in 212 species of birds to determine the function and the ecological correlates of variation in gland size. Bird species with larger uropygial glands had more genera of chewing lice of the sub-order Amblycera, but not of the sub-order Ischnocera, and more feather mites. There was a fitness advantage associated with relatively large uropygial glands because such species had higher hatching success. These findings are consistent with the hypothesis that the uropygial gland functions to manage the community of microorganisms, and that certain taxa of chewing lice have diverged as a consequence of these defenses.

A negative covariation between toxoplasmosis and CoVID-19 with alternative interpretations.

Coronaviruses may exert severely negative effects on the mortality and morbidity of birds and mammals including humans and domestic animals. Most recently CoVID-19 has killed about half million people (27th of June, 2020). Susceptibility to this disease appears to differ markedly across different societies but the factors underlying this variability are not known. Given that prevalence of toxoplasmosis in human societies may serve as a proxy for hygiene, and it also exerts both direct and immune-mediated antiviral effects, we hypothesize a negative covariation between toxoplasmosis and measures of the CoVID-19 pandemic across countries. We obtained aged-adjusted toxoplasmosis prevalence of pregnant women from the literature. Since the differences in the CoVID-19 morbidity and mortality may depend on the different timing of the epidemics in each country, we applied the date of first documented CoVID-19 in each country as a proxy of susceptibility, with a statistical control for population size effects. Using these two indices, we show a highly significant negative co-variation between the two pandemics across 86 countries. Then, considering that the wealth of nations often co-varies with the prevalence of diseases, we introduced GDP per capita into our model. The prevalence of toxoplasmosis co-varies negatively, while the date of first CoVID-19 co-varies positively with GDP per capita across countries. Further, to control for the strong spatial autocorrelation among countries, we carried out a Spatial Structure Analyses of the relationships between the date of first CoVID-19, prevalence of toxoplasmosis, and GDP per capita. Results of this analysis did not confirm a direct causal relationship between toxoplasmosis and susceptibility to the CoVID-19 pandemics. As far as an analysis of observational data let us to suggest, it appears that the interaction between CoVID-19 and toxoplasmosis is mediated by GDP per capita and spatial effects. This prompts the question whether the formerly known covariations of CoVID-19 and BCG vaccination or air pollution might have also emerged as spurious indirect effects.

The motivation for biological aggression is an inherent and common aspect of the human behavioural repertoire.

According to a widespread opinion shared by the vast majority of historians, instances of aggression using pathogen weapons constitute extremely rare events in human history. Similarly, students of human behaviour tend to believe that their science plays no role in explaining this phenomenon, which is held to be exceptional and abnormal. Contrary to this dominant view, I argue that Hamiltonian spite - like Hamiltonian altruism - is an inherent part of the human behavioural repertoire and it includes the use of pathogens for spiteful purposes. This paradigm is supported by the following observations. The use of pathogens as weapons emerged far before the scientific understanding of the nature of infections and epidemics, though it has been underrepresented in written history ever since. It is also present in our expectations concerning the likely behaviour of an enemy and it is also a frequent component of threats. Several languages appear to bear linguistic references to our motivation for biological aggression in profanity. Finally, given that wartime epidemics kill people at a rate comparable to (or exceeding) that of mechanical weapons, all wars fought in recorded history incorporated an element of aggression through biological means. On the basis of these arguments, I claim that the motivation for biological aggression is an inherent and common aspect of past and present human behaviour.

The tripartite immune conflict in placentals and a hypothesis on fetal-->maternal microchimerism.

There is a two-way traffic of immune cells through the placenta; and fetal immune cells are often present in the maternal body even long after giving birth. We present an adaptationist theory to interpret fetal-->maternal microchimerism and the diverse set of concomitant medical phenomena. We handle fetal, maternal, and paternal adaptive interests separately and in interaction with one another. Fetuses may benefit from immunological information gathered by migrant cells in the maternal body, and also from improved maternal defence. However, they may be jeopardized by a selfish maternal usage of fetal-->maternal microchimerism - i.e., some mothers get pregnant only to improve their immune system and then to abort. The use of microchimeric cells by the maternal immune system may contribute to the adaptive benefits of female choosiness and polyandry. While fathers may enjoy an indirect benefit from enhanced fetal and maternal health, they also face the risk of wasting sexual efforts due to selfish pregnancies of cheating females. Paternal alleles acting via clones of microchimeric cells in the maternal body could launch an immunological attack against the non-kin sperm in the female genitalia, or against the non-kin fetus in the womb. Furthermore, an intraspecific version of Zahavi's Mafia Hypothesis could explain a potential interaction between the abortion of fetuses and a subsequent rise of an autoimmune disease. We suggest that males may be capable to provoke microchimerism-induced autoimmune-like diseases in the mother in revenge of selfish pregnancies. This hypothetic paternal threat could increase the maternal costs associated to selfish pregnancies. From a medical point of view, we propose new interpretations for autoimmune-like diseases, infertility, miscarriage, and also for the prevailing connections among them. Specifically, we argue that miscarriages may cause autoimmune diseases, a reversed causality as compared to the currently accepted one.

A psychochemical weapon considered by the Warsaw Pact: a research note.

Contrary to widespread rumours during the Cold War era, little, if any, evidence existed in the scientific literature to support the view that the Soviet Union or its Warsaw Pact allies considered the use of psychochemical weapons militarily. The Hungarian State Archives have recently opened up declassified records of Hungary's State Defence Council meetings held between 1962 and 1978. Materials submitted to the Council include reports about the coordinative meetings of the Warsaw Pact military medical services. Research into possible countermeasures against psychotropic drugs is listed as a research priority assigned to Hungary in 1962. Hungary rejected this task in 1963, but joined the ongoing project again in 1965. Methylamphetamine was produced in Budapest for use as an experimental model of such weapons. Within the context of contemporary western research, this drug was considered to be an effective interrogation tool. Similarly to the CIA, Hungary also failed to develop an antidote against it and the project was terminated, fruitlessly, in 1972. These documents serve as evidence that a Warsaw Pact forum had, in fact, been considering a psychochemical weapon as a "warfare agent."

Rensch's rule in avian lice: contradictory allometric trends for sexual size dimorphism.

Rensch's rule (RR) postulates that in comparisons across closely related species, male body size relative to female size increases with the average size of the species. This holds true in several vertebrate and also in certain free-living invertebrate taxa. Here, we document the validity of RR in avian lice using three families (Philopteridae, Menoponidae, and Ricinidae). Using published data on the body length of 989 louse species, subspecies, or distinct intraspecific lineages, we applied phylogenetic reduced major axis regression to analyse the body size of females vs. males while accounting for phylogenetic non-independence. Our results indicate that philopterid and menoponid lice follow RR, while ricinids exhibit the opposite pattern. In the case of philopterids and menoponids, we argue that larger-bodied bird species tend to host lice that are both larger in size and more abundant. Thus, sexual selection acting on males makes them relatively larger, and this is stronger than fecundity selection acting on females. Ricinids exhibit converse RR, likely because fecundity selection is stronger in their case.

Biostatistics for Parasitologists - A Primer to Quantitative Parasitology.

The aggregated distributions of host-parasite systems require several different infection parameters to characterize them. We advise readers how to choose infection indices with clear and distinct biological interpretations, and recommend statistical tests to compare them across samples. A user-friendly and free software is available online to overcome technical difficulties.