Sexual dichromatism and parasitism in british and irish freshwater fish

Hamilton & Zuk (1982) predicted that there should be a relationship between a species' parasite load and its sexual showiness. The relationship between the number of parasite genera reported from a fish family and its sexual dichromatism was examined in British and Irish freshwater fish. Eleven other ecological and life history variables which may also cause sexual dichromatism were also examined. The changes in appearance that take place are always more marked in males and occur only during the breeding season. This strongly implicates sexual selection as an important selective determinant. There was a significant positive correlation between a fish family's sexual dichromatism and the number of parasite genera reported from it. This remained significant when the influences of near-significantly correlated ecological and life history variables were removed. Using more detailed published parasite data on six species, there was also a significant correlation between the mean number of parasite species per host individual and host sexual dichromatism. These results support Hamilton & Zuk's bright, parasiteresistant male and choosy female hypothesis.     

MHC,parasites and antler development in red deer: no support for the Hamilton & Zuk hypothesis

The Hamilton‐Zuk hypothesis proposes that the genetic benefits of preferences for elaborated secondary sexual traits have their origins in the arms race between hosts and parasites, which maintains genetic variance in parasite resistance. Infection, in turn, can be reflected in the expression of costly sexual ornaments. However, the link between immune genes, infection and the expression of secondary sexual traits has rarely been investigated. Here, we explored whether the presence and identity of functional variants (supertypes) of the highly polymorphic major histocompatibility complex (MHC), which is responsible for the recognition of parasites, predict the load of lung and gut parasites and antler development in the red deer (Cervus elaphus). While we found MHC supertypes to be associated with infection by a number of parasite species, including debilitating lung nematodes, we did not find support for the Hamilton‐Zuk hypothesis. On the contrary, we found that lung nematode load was positively associated with antler development. We also found that the supertypes that were associated with resistance to certain parasites at the same time cause susceptibility to others. Such trade‐offs may undermine the potential genetic benefits of mate choice for resistant partners.     

Genetic variability in the European minnow,Phoxinus phoxinus (L.)

An electrophoretic study of genetic variation at 11 loci was performedfor a population of European minnows, Phoxinus phoxinus (L.). Ten loci, EST-1 ^*, EST-2 ^* EST-3 ^*,GPD-1 ^*,GPD-2 ^*,GPI-1 ^*,GPI-2 ^*,MPI ^*,6PGD ^* and PGM ^* were polymorphic. IDH ^*wasmonomorphic. The mean number of heterozygotic loci over all 176 fish was 3.05 ± 0.104(SE). Observed mean heterozygosity was 0.28±0.058(SE) and expected mean heterozygosity was 0.27±0.054(SE). EST-2 ^*, EST-3 ^* andPGM ^* were not in Hardy-Weinberg equilibrium. Length,condition, parasite numbers or male breeding characters, i.e. red colorationand tubercles, were not influenced by single enzyme loci.     

EFFECTS OF A HAEMATOPHAGOUS MITE ON THE BARN SWALLOW (HIRUNDO RUSTICA): A TEST OF THE HAMILTON AND ZUK HYPOTHESIS

I tested three assumptions of the Hamilton and Zuk hypothesis (1982), which suggests that the extravagant male plumage of many bird species allows females to choose mates that are resistant to the parasites exploiting the host population at a given time. By choosing such males as mates, females will rear offspring carrying the genes for resistance. I tested three necessary conditions for the Hamilton and Zuk model: (1) whether parasites affect the fitness of their hosts; (2) whether there is heritable variation in parasite resistance, and (3) whether the expression of the sexual ornament varies with parasite burden. The haematophagous mite Ornithonyssus bursa (Macronyssidae, Gamasida) sucks blood from their Barn Swallow (Hirundo rustica) hosts. Experimental manipulation of mite loads and partial cross-fostering experiments on Barn Swallows, where half of the nestlings in the brood were exchanged with nestlings from another nest, shows that parasite burdens and origin, but not rearing conditions, of Bam Swallow nestlings, affected their adult tarsus length and maximum body weight shortly before fledging. Mite loads of adult Barn Swallows at spring arrival were more similar to mite loads of their own offspring, whether reared in their own or in foster nests inoculated with mites, than to loads of foster offspring. Parent Barn Swallows with long tail ornaments had offspring with smaller mite loads in the partial cross-fostering experiments. The amount of increase in male tail ornaments from one year to another was negatively related to experimentally manipulated mite loads of Barn Swallow nests during the preceding breeding season. In conclusion, the three assumptions of the hypothesis were supported by the experimental tests.     

Parasite load and brightness in lizards: an interspecific test of the Hamilton and Zuk hypothesis

Hamilton & Zuk (1982) hypothesized a positive correlation between a species' sexual showiness and its level of parasitic infection. We tested the hypothesis in 26 species of lizards, members of a class of vertebrates never before used to test the model. The prevalence of parasites was determined using published lists of parasites found in wild lizard populations. An index of showiness (brightness) was derived by scoring photographs of lizards in natural settings. Contrary to expectations of Hamilton & Zuk (1982), we found an inverse correlation between a lizard species' brightness and parasite prevalence. No correlation was found between a species' brightness and the number of parasite genera, species, or percentage of individual infecting parasite taxa. These results are discussed in relation to other interspecific tests of the hypothesis.     

Sexual selection and the role of parasites

Six years ago, William Hamilton and Marlene Zuk(1) proposed a novel and - on the face of it - bizarre explanation for the evolution of secondary sexual characters. Contrary to most theories of sexual selection through female choice(2), Hamilton and Zuk suggested that the bright colours and elaborate ornaments of many animal species have a utilitarian function: they allow females to assess a potential mate's ability to resist parasites. Recently, several studies have begun to test the hypothesis directly, and the idea has been discussed at two meetings(3,4). Here I review the work to date, and discuss the difficulties involved in gathering and interpreting the evidence still required.     

Towards a functional understanding of "good genes"

The Hamilton & Zuk hypothesis (1982) underpins our understanding of the relationship between secondary sexual characters, parasites, and immunological function. However, despite the wealth of empirical studies that attempt to address issues raised by the Hamilton & Zuk hypothesis, there have been no overt attempts to identify the "good genes" that females select or how those good genes influence the host's immune system. Behavioural ecologists have generally viewed this aspect of immunity as a black box. In this review we propose candidate good genes in vertebrates, discuss their role in immune function and parasite resistance, and discuss several aspects of the assumptions that pervade studies of parasite mediated sexual selection in vertebrates. We also examine invertebrates (specifically insects) where our current knowledge of these systems suggests the patterns apparent in vertebrates are not underpinned by the same genetic mechanisms.     

Parasites and showy males: malarial infection and color variation in fence lizards

Summary Hamilton and Zuk (1982) proposed that the quality of male showy traits reflects genetically-based resistance to parasites and can be used by females to select mates that are less prone to parasitic attack. The hypothesis requires that a particular state of a variable showy trait should be associated with parasite infection. We tested this idea with a population of western fence lizards, Sceloporus occidentalis, infected with the malarial parasite, Plasmodium mexicanum. Ventral color pattern is strongly dimorphic in fence lizards and varies greatly among males in this population. Malaria-infected males exhibited significantly more black and less pale on their ventral surface than did noninfected males of similar body size. This difference was not a function of differing ages of infected and noninfected animals of the same body size. However, logistic regression demonstrated that females using male ventral color as a gauge of infection status would only marginally improve their chance of choosing a noninfected lizard over random selection of mates.     

New Antarctic deep-sea weird leech (Hirudinida: Piscicolidae): morphological features and phylogenetic relationships

A new fish leech Ambulobdella shandikovi n. g., n. sp. (Hirudinida: Piscicolidae), a parasite of Whitson’s grenadier Macrourus whitsoni (Regan) (Macrouridae: Gadiformes) collected in the Ross Sea at depths from 1,221 to 1,433 m, is described and compared with related taxa based on morphological and molecular characters. Ambulobdella shandikovi n. sp. is characterised by prominent segmental tubercles on the venter and dorsal segmental tubercles, an uncommon appearance of its anterior sucker with ear-like edges and an inner membrane around the mouth-pore, well-developed musculature and a unique combination of features of the reproductive and digestive systems. The presence of uncommon tubercles can be attributed, in part, to temporary associations of A. shandikovi n. sp. with its fish hosts and a need for well-developed sensory and locomotory organs. A certain locomotory function of ventrolateral tubercles of A. shandikovi n. sp. is hypothesised and discussed. Further deep-sea surveys are obviously needed to shed light on the behaviour and mode of locomotion of this species.     

Do highly ornamented and less parasitized males have high quality sperm? – an experimental test for parasite-induced reproductive trade-offs in European minnow (Phoxinus phoxinus)

Parasites take their resources from hosts and thus directly reduce available resources for hosts’ own body functions, such as growth and reproduction. Furthermore, parasite infections cause significant indirect costs to their hosts in terms of increased investments on immune defense. In this study, we investigated the impact of parasite infection on the sperm quality and expression of secondary sexual ornamentation (saturation of the red abdominal colouration and number of breeding tubercles) in the Eurasian minnow (Phoxinus phoxinus). We exposed minnows to a high and low dose of common nonspecific fish ectoparasite, the glochidia larvae of duck mussel (Anodonta anatina) and tested whether parasite infection leads to trade‐off in sperm quality and/or ornamental expression. We found that glochidia infection reduces the curvature of the sperm swimming trajectory, number of breeding tubercles, and possibly male competitive ability, but does not affect expression of male color ornamentation. Furthermore, glochidia infection was found to reduce sperm motility, but only when all the noninfected individuals were excluded from the model. Supporting one of the predictions by phenotype‐linked fertility hypothesis both in high‐infection and low‐infection group male breeding colouration was positively associated with sperm quality. Our results suggest that although glochidia infection may have negative impact on male reproductive success, parasite‐induced costs may not create strong trade‐off between breeding colouration and sperm quality or that such trade‐off become detectable only in resource‐limited conditions.     

Parasites and Sexual Selection in Birds of Paradise

We studied blood parasite infections in relation to aspectsof sexual selection and mate choice in 10 species of birds ofparadise. Across species there was a significant, positive correlationbetween relative parasite intensity and showiness in males.Parasite infections also correlated across species with thedegree of sexual dimorphism and varied with mating systems.Promiscuous species were showier and had significantly higherparasite prevalences than monogamous species. Within one species,Lawes' Parotia (Parotia lawesii), parasite intensity was negativelycorrelated with all phenotypic traits examined, a pattern significantlydifferent than random. The mating success of males with lowparasite intensities varied, but males with high intensitiesdid not mate. Sampling of individual males on repeated occasionsrevealed large temporal differences in parasite counts whichspanned the range believed to affect behavior and mating success.Whereas the interspecific correlations support one predictionof the Hamilton and Zuk hypothesis on parasites and sexual selection,the intraspecific data are equivocal with respect to a secondprediction of this hypothesis. Parasites appear to influencethe behavior of Lawes' Parotia, but alternative explanationsto that of Hamilton and Zuk for this effect are equally plausibleand there is no evidence of a link between female choice andthe traits in males indicative of parasite loads. We suggestthat female Lawes' Parotia may be avoiding highly infected malesrather than actively choosing parasite-resistant males.     

Male Ornamentation in the European Minnow (Phoxinus phoxinus) Signals Swimming Performance

The association between male secondary sexual ornamentation and male quality has been under intensive study for decades. It has been shown that male ornaments in fish indicate a variety of physical and genetic qualities. However, only few studies have investigated whether male ornaments signal locomotor performance, a trait that is expected to be vital for individual fitness in the nature. In this study, we sampled two populations of European minnow (Phoxinus phoxinus), a cyprinid fish species with spectacular and colourful male breeding ornamentation, and examined the association between male ornaments (tubercle number, lateral darkness and abdominal redness), swimming performance, and three fitness‐related traits, body length (L_T), condition factor (K) and gonadosomatic index (GI). We found that male tubercle number and abdominal redness were positively correlated with their swimming performance. In addition, these two ornaments were correlated with male fitness‐related traits, which in turn were also directly associated with the swimming performance. In general, these results suggest that in the European minnow male ornamentation may act as an honest signal of several fitness‐related traits and locomotor performance in the nature.     

Barbus comizo infestation by Lernaea cyprinacea (Crustacea: Copepoda) in the Guadiana River,southwestern Spain

The purposes of this study were to collect baseline data on parasitology of a native species and to determine if parasite infestations by Lernaea cyprinacea were related to size, month and sex. An infestation of L. cyprinacea was monitored on Barbus comizo (n = 307) from March to October 2004 in the River Guadiana, south‐western Iberian Peninsula; over 55.7% of the examined B. comizo were infested. The number of female parasites per fish ranged from 1 to 8. There were significant differences in prevalence, abundance, and intensity among months. Prevalence was highest in August, while abundance (5.16) and intensity (5.32) were highest in September. Infestation levels were significantly related to fish size (r_s = 0.947, n = 171, P < 0.05) and condition factor (r_s = ?0.199, n = 169, P < 0.05). Haematocrit values were significantly lower in infected fish (Mann–Whitney U = 21, P < 0.05). The level of infestation was independent of sex (χ² = 0.87, P < 0.05). The dorsal and ventral areas of the fish were the most heavily parasitized.     

Experimental test of the influence of the size of shoals and density of fish on parasite infections

There is an increasing interest in the relationship between adult coral reef fishes and their parasites; however, there is almost no information concerning post larvae (i.e. newly recruited) fish and their parasites. Newly recruited and juvenile fish are highly sensitive to their environment and they have a very low survival rate. Fish grouping behavior has been studied in several ways and many studies have shown that increasing group size might be a way to share a cost linked to parasitism, via a dilution effect. To my knowledge, this hypothesis has never been tested for juveniles of fish. The potential influence of school size and density of a juvenile coral reef fish, Haemulon flavolineatum (Haemulidae), on the success of a monogenean fish parasite, Haliotrema sp., was studied. First, the fish acquired their monogeneans when they were more than 40 mm long, size corresponding also to a change in their foraging behavior. Second, there was a positive effect of fish density on the success of the establishment of the parasite on the host population. Finally, although the parasite success was not directly related to fish group size, the parasite abundance was higher for intermediate shoal size. These results are discussed in terms of the consequence parasitism may have on a fish's shoaling and to a larger extent on the fish population.     

Genetic effective size,Ne, tracks density in a small freshwater cyprinid,Pecos bluntnose shiner (Notropis simus pecosensis)

Genetic monitoring tracks changes in measures of diversity including allelic richness, heterozygosity and genetic effective size over time, and has emerged as an important tool for understanding evolutionary consequences of population management. One proposed application of genetic monitoring has been to estimate abundance and its trajectory through time. Here, genetic monitoring was conducted across five consecutive year for the Pecos bluntnose shiner, a federally threatened minnow. Temporal changes in allele frequencies at seven microsatellite DNA loci were used to estimate variance effective size (N_eV) across adjacent years in the time series. Likewise, effective size was computed using the linkage disequilibrium method (N_eD) for each sample. Estimates of N_e were then compared to estimates of adult fish density obtained from traditional demographic monitoring. For Pecos bluntnose shiner, density (catch‐per‐unit‐effort), N_eV and N_eD were positively associated across this time series. Results for Pecos bluntnose shiner were compared to a related and ecologically similar species, the Rio Grande silvery minnow. In this species, density and N_eV were negatively associated, which suggested decoupling of abundance and effective size trajectories. Conversely, density and N_eD were positively associated. For Rio Grande silvery minnow, discrepancies among estimates of N_e and their relationships with adult fish density could be related to effects of high variance in reproductive success in the wild and/or effects of supplementation of the wild population with captive‐bred and reared fish. The efficacy of N_e as a predictor of density and abundance may depend on intrinsic population dynamics of the species and how these dynamics are influenced by the landscape features, management protocols and other factors.     

Three new species of Tricorythopsis (Ephemeroptera: Leptohyphidae) from southeastern Brazil

Three new species of Tricorythopsis Traver (Ephemeroptera: Leptohyphidae) are described and illustrated based on nymphs from southeastern Brazil. These new species can be distinguished from other species of the genus by the following characters: Tricorythopsis araponga sp. n.: (1) femora with long setae; (2) abdominal segments 5–7 with dorsal tubercles; (3) tarsal claws with 4–6 marginal denticles and 7 + 4 submarginal denticles. Tricorythopsis baptistai sp. n.: (1) tarsal claws with 4–5 large marginal denticles and one submarginal denticle on each side; (2) abdominal colour pattern; (3) abdomen without tubercles; (4) coxae without projections. Tricorythopsis pseudogibbus sp. n.: (1) abdominal segments 6–8 with small dorsal tubercles; (2) tarsal claws with four large marginal denticles, and 3 + 1 or 2 submarginal denticles; (3) coxae dorsally projected; (4) femora broad and with short setae; (5) pronotum with anterolateral projection.     

The relations between 0+ fish density,zooplankton size and the vulnerability of pikeperch,Stizostedion lucioperca,to angling in the Frisian lakes

During a five year study, the size and abundance of zooplankton and 0+ fish in September were registered in a number of Frisian lakes. The mean size of Daphnia was related to the abundance of planktivorous 0+ fish in September. Immigration of fish larvae into the Frisian Lake District from the IJsselmeer in June was observed in 1979 and 1980. The abundance of 0+ fish was negatively related and the size of Daphnia was positively related to the vulnerability to angling of predatory pikeperch.     

MHC VARIATION IS RELATED TO A SEXUALLY SELECTED ORNAMENT,SURVIVAL, AND PARASITE RESISTANCE IN COMMON YELLOWTHROATS

Hamilton and Zuk proposed that females choose mates based on ornaments whose expression is dependent on their genetically based resistance to parasites. The major histocompatibility complex (MHC) plays an important role in pathogen recognition and is a good candidate for testing the relationships between immune genes and both ornament expression and parasite resistance. We tested the hypothesis that female common yellowthroats prefer to mate with more ornamented males, because it is a signal of their MHC‐based resistance to parasites and likelihood of survival. In this species, females prefer males that have larger black facial masks as extrapair mates. Using pyrosequencing, we found that mask size was positively related to the number of different MHC class II alleles, as predicted if greater variation at the MHC allows for the recognition of a greater variety of pathogens. Furthermore, males with more MHC class II alleles had greater apparent survival, and resistance to malaria infection was associated with the presence of a particular MHC class II allele. Thus, extrapair mating may provide female warblers with immunity genes that are related to parasite resistance, survival, and the expression of a male ornament, consistent with good genes models of sexual selection.     

A new species of the genus Mekongina Fowler, 1937 (Cypriniformes: Cyprinidae) from South China

A new species of cyprinid fish, Mekongina lancangensis, is described from the upper Mekong River drainage in Southern Yunnan, China. The new species is distinguished from the other species of Mekongina occurring in the lower Mekong River drainage by possessing the following combination of characters: one pair of rostral barbels; two rows of tubercles irregularly scattered on the snout and cheeks, with two enlarged tubercles present at each side of anterior of the snout; 19–27 rostral marginal lappets; lateral line with 38–41 scales; 5·5 or 6·5 scales in transverse series from dorsal‐fin origin to lateral line; 18–20 circumpeduncular scales; snout length 31·9–36·9% head length; tip of depressed anal‐fin rays extending to the caudal‐fin base.