Ultra-violet reflectance of male and female red grouse, Lagopus lagopus scoticus, sexual ornaments reflect nematode parasite intensity

In many birds, females prefer males with the biggest or brightest sexual ornaments, which might reflect a higher phenotypic quality, such as fewer parasites. Unlike humans, most birds detect near-ultraviolet (UV) light, and UV signals can play an important role in sexual signalling and mate choice. Using a spectrophotometer, we analysed the colour of red grouse Lagopus lagopus scoticus sexual ornaments (their combs). We first show that combs reflect both in the red (600–700 nm) and UV (300–400 nm) part of the spectrum. Second, we investigated whether comb size and colour, and UV reflectance in particular, reflected an aspect of individual quality: the intensity of infection by a main nematode parasite, the caecal threadworm Trichostrongylus tenuis . We first analysed comb size and colour variation, and parasite intensity variation, in relation to sex and age. Males had bigger and redder combs than females, but UV brightness was greater for female than for male combs. Comb colour also differed between age groups, with young birds of both sexes showing brighter UV than old birds. Young grouse also had fewer T. tenuis worms than old grouse. We further tested whether intensity of infection could be predicted from comb characteristics (size and colour) in male and female red grouse. We found that parasite intensity was not significantly related to comb size or red brightness, but fewer worms were predicted from brighter UV in combs, in both males and females. The results indicate that UV reflectance of combs have a quality revealing function and might play an important role in grouse mate choice: UV brightness of combs could enable both male and female red grouse to assess the parasite loads of a potential mate.     

Cestodiasis in the red grouse in Scotland

Little is known about the cestode parasites of red grouse Lagopus lagopus scoticus. The carcasses of 71 free-living and grouse collected from northeastern Scotland (UK) between June 1991 and September 1992 were examined for the cestode Paroniella urogalli. Over 70% of young and old birds were infected in summer and winter. Cestode abundance ranged from 0 to 29 (mean +/- SE = 5.54 +/- 0.81) worms per host and did not differ significantly between young and old grouse, or between seasons. Cestode intensities did vary significantly between the sexes, but in opposite directions at two sites. A significantly greater total biomass (dry weight) of cestodes was found in grouse in the summer. Cestode biomass also was greater in young than in old grouse, although there was no significant difference after controlling for the effects of site, year and season. However, in the summer total cestode biomass in young grouse was significantly greater per unit bodyweight than in old grouse. No evidence was found for an effect of cestodiasis on grouse condition but a significant positive correlation existed between grouse comb condition and cestode abundance. Estimates of mean per capita worm mass suggested an intensity dependent reduction in the weight of individual cestodes in birds in their first summer. Per capita cestode mass was significantly lower in winter than summer, which may suggest that the production of mature proglottids varies with season. There was also a significant negative correlation between numbers of the nematode Trichostrongylus tenuis and the total biomass of P. urogalli in grouse.     

Temporal changes in age and sex ratios of forest grouse harvested in northeastern oregon

Ratio of immature (young of the year) grouse to adult birds (I:A) in the harvest of upland game birds is commonly used as an index to annual reproduction; however, I:A ratios can vary as the season progresses producing biased estimates. We analyzed I:A ratios in the daily harvest of dusky grouse (Dendragopus obscurus) and ruffed grouse (Bonasa umbellus) in northeastern Oregon over 28 years (1981–2008) and found that I:A ratios in the harvest declined for both species as the hunting season progressed. We also analyzed ratios of adult female to adult male (AF:AM) grouse to determine if female and male grouse were harvested in equal numbers throughout the harvest season. We found that more males than females of both species were harvested, but that AF:AM ratio of both dusky and ruffed grouse did not change during most of the hunting season. Approximately 50% of the annual harvest occurred during the first 14 days of the hunting season. Therefore, we recommend using the ratios of I:A birds in the first 14 days of the harvest season as the best index to annual reproduction of forest grouse in northeast Oregon. © 2011 The Wildlife Society.     

Discovery of Maritrema obstipum (Digenea: Microphallidae) from migratory birds in Korea

Adults of Maritrema obstipum (Digenea: Microphallidae) were found in the intestines of 4 species of migratory birds, including the sanderling (Crocethia alba), Kentish plover (Charadrius alexandrines), Mongolian plover (Charadrius mongolus), and red-necked stint (Calidris ruficollis), collected from Yubu Island, Chungcheongnam-do, Korea. The worms of were 451 × 265 μm in size, and were easily identifiable as Maritrema species by the presence of the cirrus sac, and the ring-like distribution of the vitellaria. More specifically, the ejaculatory duct curved posteromedially, and the 2 parts of vitelline follicles were found to be distinct at the posterior end. The eggs were brown-colored, and 19.8 × 12.3 μm in size. All these findings implicated M. obstipum as the pertinent species of the worms. Beside these, adult worms of Gynaecotyla squatarolae, Parvatrema duboisi, and Acanthoparyphium sp. were also discovered. This is the first report establishing migratory birds as the natural definitive hosts for M. obstipum.     

Seasonality and causes of black grouse (Lyrurus tetrix,Galliformes, Tetraonidae) death in Western Russia according to count of remains

Black grouse (Lyrurus tetrix) death in north and middle zones of the Russian plain were studied in 1981–2013 according to counts of the remains of perished individuals. The average annual death of birds older than 1.5–2 months is estimated as 2.0–2.5 birds/km². Its maximum falls into spring or winter-spring season and minimum into summer. Predation is main cause of death; all other reasons (unusual weather, hunting, no established circumstances) determine only one-quarter of total loss. Raptors cause maximal death of black grouse in northern regions. No significant differences between death of males and females were detected, but seasonal patterns of their death is not the same, and factors that determine their act differently. The female losses as a result of activity of all predators are distributed relatively uniformly by seasons; male death is higher in spring than in other times of year. The dynamics of male and female removal by only raptors is similar: maximum in spring, decrease in summer, and drop to minimum in autumn and winter. Seasonal distribution of losses due to activity of carnivorous beasts is uneven; male death is higher in transitional seasons (as compared with winter and summer), and female losses are maximal in winter and minimal in spring-summer period.     

Territorial status and survival in red grouse Lagopus lagopus scoticus: hope for the doomed surplus?

A previous study of survival in territorial and non-territorial red grouse Lagopus lagopus scoticus conducted between 1957 and 1967 found that territorial status in the autumn pre-determined over-winter survival. A very high proportion of territorial birds survived and virtually all non-territorial birds died or emigrated. We tested the hypothesis that over-winter survival was dependent on territorial status within four grouse populations in Scotland between 1986 and 1993. In contrast to the previous study, 66% of non-territorial birds survived over winter compared to approximately 70% of territorial birds. There was no significant effect of territorial status on the survival estimates. Moreover, some of the birds considered to be non-territorial during autumn went on to successfully raise a brood. We suggest that on our study sites, territory ownership in autumn did not greatly influence over-winter survival, and territorial behaviour did not determine breeding density as previously supposed. We postulate differences with other studies may reflect variations in scale and predation pressure.     

Susceptibility of greater sage-grouse to experimental infection with West Nile virus

Populations of greater sage-grouse (Centrocercus urophasianus) have declined 45-80% in North America since 1950. Although much of this decline has been attributed to habitat loss, recent field studies have indicated that West Nile virus (WNV) has had a significant negative impact on local populations of grouse. We confirm the susceptibility of greater sage-grouse to WNV infection in laboratory experimental studies. Grouse were challenged by subcutaneous injection of WNV (10(3.2) plaque-forming units [PFUs]). All grouse died within 6 days of infection. The Kaplan-Meier estimate for 50% survival was 4.5 days. Mean peak viremia for nonvaccinated birds was 10(6.4) PFUs/ml (+/-10(0.2) PFUs/ml, standard error of the mean [SEM]). Virus was shed cloacally and orally. Four of the five vaccinated grouse died, but survival time was increased (50% survival=9.5 days), with 1 grouse surviving to the end-point of the experiment (14 days) with no signs of illness. Mean peak viremia for the vaccinated birds was 10(2.3) PFUs/ml (+/-10(0.6) PFUs/ml, SEM). Two birds cleared the virus from their blood before death or euthanasia. These data emphasize the high susceptibility of greater sage-grouse to infection with WNV.     

Phylogeography of willow grouse (Lagopus lagopus) in the Arctic: taxonomic discordance as inferred from molecular data

Using independently segregating nuclear single nucleotide polymorphisms (SNPs) and mitochondrial control region sequences, we found an east–west division among sampled willow grouse Lagopus lagopus subspecies. This division cut across the range of the subspecies with the largest distribution (lagopus) and thus contradicted existing taxonomic classifications. Russian Lagopus lagopus lagopus tended to cluster with North American willow grouse partly classified as other subspecies. Scandinavian willow grouse (L. l. lagopus) clustered with red grouse from Britain and Ireland (Lagopus lagopus scoticus and Lagopus lagopus hibernicus) but substructuring confirmed the monophyly of the latter. In North America, we could not detect any major genetic divisions apart from two birds described as alexandrae from the Heceta Island (Alaska) when using mitochondrial sequences. Other samples from North America were intermingled regardless of whether they were described as muriei, alexandrae or lagopus. A specimen described as alexandrae was to some extent distinct when analysing the SNP data. The genetic analyses indicated some concordance between genetics and taxonomy but not complete congruence. This is particularly evident for mitochondrial DNA network analyses. We suggest that the taxonomy of this species would benefit by a careful re‐examination of the available evidence for subspecies. It appears as if subspecies status is a poor proxy for assigning evolutionary significant units and management units in this species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 77–90.     

On fruit eating by Scottish Red grouse

Fruit-eating behaviour of Red grouse was studied in the Grampians of Scotland from analyses of bird droppings for seeds and from direct observations of feeding birds. The commonest moorland species with edible fruits are blaeberry, Vaccinium myrtillus L., cowberry, Vaccinium vitis-idaea L., and crowberry, Empetrum nigrum L.
The fruit-frugivore interaction is not species specific, as grouse ate berries of all the fruiting species, but almost all individuals showed high fruit specificity for one type of fruit. Fruit selection is density-dependent and probably depends on the formation by grouse of a search specific image of the commonest fruit available in the feeding area. This is well known in polyphagous predator-prey interactions in animals but has not so far been reported in animal-plant feeding interactions.
We related fruit selection to both relative fruit abundance and plant cover. Some birds were feeding highly selectively on crowberries which were low in fruit abundance but had high relative cover. This indicated that the fruit resource had been large but had become depleted. Such birds would eventually lose their specific searching image and, following a short period of random search, would form new searching images.
The fruit cues and effect of density dependent selection on the evolution of fruiting behaviour was discussed.     

Behavioural evidence for ultraviolet vision in a tetraonid species – foraging experiment with black grouse Tetrao tetrix

In addition to wavelengths visible to humans (400–700 nm), many birds are able to detect near ultraviolet light (320–400 nm). Most studies of ultraviolet (UV) vision in birds have concentrated on the importance of UV vision in intraspecific signalling, especially in passerine birds. However, birds may also use UV vision for other purposes, e.g. foraging. We performed a laboratory experiment to test whether a tetraonid species, black grouse Tetrao tetrix , could detect the difference between UV-reflecting and non-UV-reflecting food items (two colour morphs of bilberry Vaccinium myrtillus ). Black grouse preferred UV-reflecting berries when UV light was used for illumination, but showed no preference in the absence of UV light. This observation establishes a potential UV sensitivity in this species; such a sensitivity should be considered in behavioural experiments with this species.     

Hen harriers and red grouse: economic aspects of red grouse shooting and the implications for moorland conservation

1.  Thirgood & Redpath (2008) propose ways in which red grouse : hen harrier conflicts could be resolved. It has also been suggested that grouse management could accept lower bag sizes (number of birds shot) thus reducing the need for intensive management of predators and habitats. This would allow hen harriers to co-exist more easily on grouse moors.
2.  We compare the bags, costs and incomes from these less intensive forms of grouse shooting with the more intensive driven shooting.
3.  Allowing high density grouse moors to decline to low density ones will result in greater loss of income than the corresponding saving of costs. This can result in moor owners abandoning grouse management and thus gamekeepers losing their employment.
4.  Losing gamekeepers from the uplands would jeopardize the protection of heather moorland and Special Protection Areas for birds, large areas of which are keepered and which currently support high numbers of breeding waders.
5.   Synthesis and applications . We agree with the study by Thirgood & Redpath that consideration of social and economic factors will be needed to resolve conflict but a reduction in management effort from driven to walked-up shooting is not the answer. A more satisfactory approach to the harrier : grouse conflict could be to try to reduce harrier predation by means of diversionary feeding and to address the problem of the rapid build-up in harrier numbers by exploring the use of a ceiling on harrier densities.     

A population genetic comparison of large- and small-bodied sage grouse in colorado using microsatellite and mitochondrial DNA markers

Sage grouse (Centrocercus urophasianus) from southwestern Colorado and southeastern Utah (United States) are 33% smaller than all other sage grouse and have obvious plumage and behavioural differences. Because of these differences, they have been tentatively recog-nized as a separate 'small-bodied' species. We collected genetic evidence to further test this proposal, using mitochondrial sequence data and microsatellite markers to determine whether there was gene flow between the two proposed species. Significant differences in the distribution of alleles between the large- and small-bodied birds were found in both data sets. Analysis of molecular variance (AMOVA) revealed that 65% of the variation in mitochondrial DNA (mtDNA) haplotypes could be explained by the large- vs. small-bodied distinction. Genetic distances and neighbour-joining trees based on allelic frequency data showed a distinct separation between the proposed species, although cladistic analysis of the phylogenetic history of the mitochondrial sequence haplotypes has shown a lack of reciprocal monophyly. These results further support the recognition of the small-bodied sage grouse as a distinct species based on the biological species concept, providing additional genetic evidence to augment the morphological and behavioural data. Furthermore, small-bodied sage grouse had much less genetic variation than large-bodied sage grouse, which may have implications for conservation issues.     

Increased DNA typing success for feces and feathers of capercaillie (Tetrao urogallus) and black grouse (Tetrao tetrix)

Noninvasive sampling, for example, of droppings or feathers, is a promising approach for molecular genetic studies on endangered and elusive animal species. Yet, such specimens are known for containing only minute amounts of DNA, resulting in lower typing success rates relative to analyses on fresh tissues such as muscle or blood. Furthermore, artefactual signals as well as contamination are more likely to occur when DNA is limited. To increase the reliability of DNA typing from noninvasive samples, optimized DNA extraction and polymerase chain reaction protocols were developed, taking advantage of developments in the forensic field aiming at successful molecular genetic analysis of DNA templates being low in quality and quantity. In the framework of an extensive monitoring project on population dynamics of capercaillie and black grouse in the Tyrolean Alps, feces samples and molted feathers from both species were collected. On a subset comprising about 200 specimens of either species, eight polymorphic short tandem repeat (STR) markers were analyzed to test these improved protocols. Besides optimizing DNA yields, both lowered sample consumption and reduced hands‐on time were achieved, and the rates of informative profiles amounted to 90.7% for capercaillie and 92.4% for black grouse. Similarly, high success rates had not been achieved in earlier studies and demonstrate the benefit of the improved methodology, which should be easily adaptable for use on animal species other than those studied here. The STR genotypes were not only powerful enough to discriminate among unrelated birds but also appeared fit for telling apart closely related animals, as indicated by Pi and Pi_sib values. The software package allelematch aided analysis of genotypes featuring possible dropout and drop‐in effects. Finally, a comparison between molecular genetic and morphology‐based species‐of‐origin determination revealed a high degree of concordance.     

The occurrence of gapeworm (Syngamus trachea) in willow grouse.

21.4% of the willow grouse (Lagopus lagopus) examined from a coastal locality in northern Norway were infected with Syngamus trachea. None of the willow grouse examined from an inland locality with colder and longer winters were infected with the parasite. It is suggested that the parasite may influence survival of the grouse population.     

Populations of Leucocytozoon gametocytes in blue grouse (Dendragapus obscurus) from Hardwicke Island, British Columbia

Population dynamics of round and elongate gametocytes of Leucocytozoon in wild and captive blue grouse (Dendragapus obscurus (Say)) from Hardwicke Island, British Columbia, were studied from 1980 to 1982. Blue grouse chicks were sampled weekly throughout each transmission season. Three patterns in the type of gametocyte produced during primary infection were observed in naturally infected captive and wild blue grouse chicks. Such variation in the expression of the gametocyte stage within a single host population suggests a different interpretation than has been previously reported for species of Leucocytozoon. The data from the primary patterns and profiles coupled with reexposure data and the asynchronous appearance of round and elongate gametocytes can be best interpreted as infection with two concurrent species of Leucocytozoon in blue grouse. More detailed research on the life cycle is necessary to confirm if two species of Leucocytozoon exist in blue grouse.     

The cost of dispersal: predation as a function of movement and site familiarity in ruffed grouse

Ecologists often assume that dispersing individuals experienceincreased predation risk owing to increased exposure to predatorswhile moving. To test the hypothesis that predation risk isa function of movement distance or rate of movement, we usedradio-telemetry data collected from 193 ruffed grouse (Bonasaumbellus) during 1996–1999 in southeastern Ohio. Cox'sproportional hazards model was used to examine whether the riskof predation was affected by the rate of movement and site familiarity.We found evidence indicating that increased movement rates mayincrease the risk of predation for adult birds but not juveniles.We also found juvenile and adult birds inhabiting unfamiliarspace were consistently at a much higher risk of predation (threeto 7.5 times greater) than those in familiar space. Our resultsindicate that although movement itself may have some effecton the risk of being preyed upon, moving through unfamiliarspace has a much greater effect on risk for ruffed grouse. Thissupports the hypothesis that increased predation risk may bean important cost of dispersal for birds.     

Myoglobin levels and mATPase activity in pectoral muscles of spruce and ruffed grouse (Aves: Tetraoninae)

Myoglobin concentration and myosin ATPase activity were measured in the pectoral muscle of wild spruce grouse (Dendragapus canadensis) and ruffed grouse (Bonasa umbellus), together with the weight of the Mm. pectoralis, supracoracoideus and heart. mATPase activities were similar in both species, but spruce grouse contained 15 times more myoglobin in the pectoralis muscle and the heart was three times heavier than that of the ruffed grouse. The relative mass of the flight muscles and wing loading were similar between species. Characteristics of the pectoral muscle of both grouse species reflect adaptations to predation and advertising displays. The glycolytic nature of the ruffed grouse pectoral muscle and small heart size is an adaptation to a sedentary existence within a small home range. The more oxidative pectoral muscle of spruce grouse together with its larger heart are adaptations to seasonal dispersals requiring more sustained flight.     

Populations of Leucocytozoon Gametocytes in Blue Grouse (Dendragapus obscurus) from Hardwicke Island,British Columbia1

Population dynamics of round and elongate gametocytes of Leucocytozoon in wild and captive blue grouse (Dendragapus obscurus (Say)) from Hardwicke Island, British Columbia, were studied from 1980 to 1982. Blue grouse chicks were sampled weekly throughout each transmission season. Three patterns in the type of gametocyte produced during primary infection were observed in naturally-infected captive and wild blue grouse chicks. Such variation in the expression of the gametocyte stage within a single host population suggests a different interpretation than has been previously reported for species of Leucocytozoon. The data from the primary patterns and profiles coupled with reexposure data and the asynchronous appearance of round and elongate gametocytes can be best interpreted as infection with two concurrent species of Leucocytozoon in blue grouse. More detailed research on the life cycle is necessary to confirm if two species of Leucocytozoon exist in blue grouse.     

Patterns in grouse and Woodcock Scolopax rusticola hunting yields from central Norway 1901–24 do not support the alternative prey hypothesis for grouse cycles

According to the alternative prey hypothesis, autumn populations of ground-nesting game birds fluctuate in synchrony with vole numbers because generalist predators that mainly eat voles switch to alternative prey, such as eggs and chicks, when vole numbers decline. In hunting statistics from Nord-Trøndelag, central Norway, 1901–24, annual fluctuations in the number of Willow Grouse Lagopus lagopus and Western Capercaillie Tetrao urogallus , but not of Woodcock Scolopax rusticola , were positively related to vole numbers in the current year. Both Woodcock and grouse indices were related to hunting indices of Goshawk Accipiter gentilis and to weather variables assumed to influence the birds' survival or reproduction, suggesting that the indices actually reflected local population levels. Synchronous vole and grouse fluctuations are consistent with the alternative prey hypothesis (although predator densities were low in the early 1900s), but the asynchronous Woodcock fluctuations refute the hypothesis. Rather, because the Woodcock does not feed on plants utilized by voles and grouse, I suggest that food quality is the ultimate factor for the synchrony in vole and grouse numbers in Norway.     

Killing of raptors on grouse moors: evidence and effects

Owing to the intensity of game management in Britain, managers of grouse moors have illegally killed raptors to increase the numbers of Red Grouse Lagopus l. scotica available for shooting. This paper summarizes evidence for the recent scale of illegal raptor killing on grouse moors and its effects on populations. It provides insights into how raptors themselves respond demographically to different levels of killing. Over Britain as a whole, most raptors have increased and expanded considerably since the 1970s, in response to reduced killing and nest destruction, and the diminished impacts of organo‐chlorine pesticides; however, in recent decades the populations of some species have declined on and around grouse moors. This is widely evident in Hen Harrier Circus cyaneus, Peregrine Falcon Falco peregrinus and Golden Eagle Aquila chrysaetos populations and in more restricted areas also in Northern Goshawk Accipiter gentilis and Red Kite Milvus milvus populations, in all of which illegal killing has been sufficient to affect numbers over wider areas. The evidence consists mainly of: (1) greater disappearance of nesting pairs, lower breeding densities or reduced occupancy of apparently suitable traditional territories on grouse moors compared with other areas; (2) reduced nest success compared with other areas; (3) reduced adult survival compared with other areas; (4) reduced age of first breeding, reflecting the removal of adults from nesting territories and their replacement by birds in immature plumage; (5) greater levels of disappearance of satellite‐tracked birds on grouse moors than elsewhere; and (6) the finding of poisoned baits and traps, and shot or poisoned carcasses of raptors. Not all these types of evidence are available for every species, and other types of evidence are available for some. The Common Buzzard Buteo buteo is currently the most numerous raptor in Britain and also seems to be killed in the greatest numbers. Other raptor species, including Merlin Falco columbarius, Common Kestrel Falco tinnunculus and Eurasian Sparrowhawk Accipiter nisus which nest on or near grouse moors, have little or no significant impact on grouse and are killed less often or not at all. In the absence of illegal killing, some raptor species breed as well or better on grouse moors than in other habitats. Merlins, in particular, seem to thrive on grouse moors, benefiting from the management involved (including predator control). Other aspects of illegal raptor killing are discussed, including suggestions for ways in which it might be reduced.