The Use of c-mos Nuclear Gene as a Phylogenetic Marker in Tetraonidae Birds

A 480-bp fragment of nuclearc-mos gene was sequenced in nine bird species representing four genera of the family Tetraonidae. It was demonstrated that nuclear genome region examined was highly conservative. The data were used to construct phylogenetic relationships among the c-mos gene sequences in Tetraonidae. The data obtained point to a paraphyletic origin of hazel grouse (Bonasa bonasia) and ruffed grouse (Bonasa umbellus).     

Molecular phylogeny of grouse: individual and combined performance of W-linked,autosomal, and mitochondrial loci

The phylogeny of grouse (Aves: Tetraoninae) was reconstructed using four noncoding loci: two were W-linked, one was autosomal, and one was the mitochondrial control region (CR). The rapidly evolving CR provided resolution throughout the tree, whereas the slowly evolving nuclear loci failed to resolve deeper nodes. The tree based on all four loci combined was almost identical to the CR tree and did not improve resolution or bootstrap support. The stemminess and imbalance of the trees were good determinants of the quality of the phylogenetic signal. The skewness of the tree score distribution (g(1)) behaved contrary to prediction; loci that had a more symmetric tree score distribution produced trees that had greater stemminess and balance. The quality of the phylogenetic signal was related to the evolutionary rate. Four clades of grouse were discovered. Two of these clades corresponded to currently recognized genera Bonasa and Lagopus. Bonasa was the sister to other grouse and Lagopus was the sister to the other two non-Bonasa clades. The third clade included Falcipennis, Tetrao, and Lyrurus. The fourth clade included the genera Centrocercus, Dendragapus, and Tympanuchus. The data support recognition of Falcipennis canadensis franklinii and Dendragapus obscurus fuliginosus as species.     

Complete mitochondrial genome sequence of Bonasa sewerzowi(Galliformes: Phasianidae) and phylogenetic analysis

Bonasa sewerzowi, the smallest and most southerly distributed grouse species in the world, is a bird endemic to China. The population of B. sewerzowi had shown a declining trend, which made it to be the endangered species in the China Red Data Book and Category I of nationally protected animals. So far, however, most studies about this species were mainly focused on the morphological and ecological aspects. In order to further study the feature of B. sewerzowi, the complete mitochondrial genome(mitogenome) of B. sewerzowi was sequenced by Illumina Hiseq 2000 high-throughput sequencing. Then, we focused on comparative genomics of two Bonasa species to find their characteristics. Finally, phylogenetic position of Bonasa was made based on the mitogenome dataset. Our results revealed that:(1) the mitogenome of B. sewerzowi, consisting of 16 658 bp, displayed typical genome organization and gene order found in other previously determined Galliformes mitogenomes;(2) the structure and composition of mitogenomes were similar between B. sewerzowi and B. bonasia;(3) the monophyly of Bonasa was well supported, which had a closer phylogenetic relationship with Meleagris gallopavo.     

Morphometrics of the Chinese grouse Bonasa sewerzowi

We report on the body size and mass and egg size and mass of the Chinese grouse (Bonasa sewerzowi), one of the least known grouse in the world. Although reported to be the smallest grouse in the world, not enough data have been available to substantiate this claim. We document that the Chinese grouse is, in fact, the smallest grouse in the world. It shows significant changes in body mass between the reproductive and nonreproductive season (males average 310 g in the reproductive season and 341 g in the nonreproductive season; females averaged 338 and 315 g, respectively). Egg mass averages 20.6–21.6 g and shows significant annual variation. The Chinese grouse shows some of the highest relative reproductive investment among grouse, both for egg mass (6.1–6.9% of female body mass) and clutch mass (38–43%).     

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.     

Seasonal changes in territorial behaviour of hazel grouse ( Bonasa bonasia) in a temperate forest of South Korea

We studied the seasonal changes in territorial behaviour of hazel grouse (Bonasa bonasia) from December 1999 to July 2001 in a temperate forest of South Korea. The study was based on the use of radio telemetry. Territory size and degree of overlap increased after the breeding period, and index of advertisement and border disputes decreased. In spring, the territory was defended by both males and females. During autumn and winter, this behaviour was relaxed to such a degree that hazel grouse cannot be considered territorial, but as forming a home range.     

Molecular phylogenetic relationships based on mitochondrial and nuclear gene sequences for the Todies (Todus, Todidae) of the Caribbean

We used mitochondrial/nuclear gene sequence analyses to determine the historical relationships of the endemic species of Todus (Aves: Todidae) from the Caribbean. We collected 1920-bp of nucleotide sequence data from the mitochondrial genes cytochrome b, ATPase 6, ATPase 8, and 591-bp of the single-copy nuclear gene c-mos for all Todus species and representatives of their outgroup taxa (Hylomanes, Barypthengus, Chloroceryle, Ceryle, and Galbula) to reconstruct the evolutionary history (via parsimony and maximum likelihood) of the five Todus species. The substitution rates among the mitochondrial genes were found to be much higher than the substitution rate for the c-mos gene, consequently resulting in higher substitutional saturation for the mitochondrial genes. When we applied weighting schemes to account for the variance in substitutional heterogeneity among the genes then parsimony and likelihood analyses both demonstrate that the genus Todus is monophyletic and closer to the Hylomanes and Barypthengus genera than the Chloroceryle and Ceryle genera. The mitochondrial-gene trees and nuclear-gene trees both show similar results, thus providing support for the relationships among the taxa from loci within two independently evolving genomes. The nuclear gene c-mos was found, therefore, to be a viable nuclear gene candidate for resolving intermediate and deep divergences.     

Inference of hazel grouse population structure using multilocus data: a landscape genetic approach

In conservation and management of species it is important to make inferences about gene flow, dispersal and population structure. In this study, we used 613 georeferenced tissue samples from hazel grouse (Bonasa bonasia) where each individual was genotyped at 12 microsatellite loci to make inference on population genetic structure, gene flow and dispersal in northern Sweden. Observed levels of genetic diversity suggest that Swedish hazel grouse do not suffer loss of genetic diversity compared with other grouse species. We found significant F(IS) (deviation from Hardy-Weinberg expectations) over the entire sample using jack-knifed estimators over loci, which is most likely explained by a Wahlund effect. With the use of spatial autocorrelation methods, we detected significant isolation by distance among individuals. Neighbourhood size was estimated in the order of 62-158 individuals corresponding to a dispersal distance of 950-1500 m. Using a spatial statistical model for landscape genetics to infer the number of populations and the spatial location of genetic discontinuities between these populations we found indications that Swedish hazel grouse are divided into a northern and a southern population. We could not find a sharp border between these two populations and none of the observed borders appeared to coincide with any potential geographical barriers.These results imply that gene flow appears somewhat unrestricted in the boreal taiga forests of northern Sweden and that the two populations of hazel grouse in Sweden may be explained by the post-glacial reinvasion history of the Scandinavian Peninsula.     

Rising burden of immature sheep ticks (Ixodes ricinus) on red grouse (Lagopus lagopus scoticus) chicks in the Scottish uplands

The sheep tick Ixodes ricinus (L.) (Acari: Ixodidae) is an ectoparasite of major economic and pathogenic importance in Scotland. Its distribution in the Scottish uplands is assumed to be governed by the abundance and distribution of its definitive hosts (deer and sheep) and climatic variables such as temperature and rainfall. As the numbers of its major host in Scotland, red deer, have increased dramatically and climatic conditions have become more favourable, the level of parasitism could have been expected to rise. We use data gathered from tick counts on over 4000 red grouse chicks Lagopus lagopus scoticus Latham (Galliformes: Tetraonidae) in various experiments over the past 19 years to ascertain whether the intensity and prevalence of parasitism has been increasing. From 1985 to 2003 the average tick burden of a parasitized red grouse chick has grown from 2.60 +/- 1.12 ticks per chick to 12.71 +/- 1.44. Over this period the percentage of chicks of a given brood parasitized has also increased from 4 +/- 2% to 92 +/- 3%. The possible implications of this increase in parasitism for red grouse production are discussed.     

Evolution of the mitochondrial DNA cytochrome b gene in Tetraonidae birds

Mitochondrial fragments containing the cytochrome b gene (1020 bp in size) of four bird species belonging to four genera of the family Tetraonidae (Tetrao parvirostris, Bonasa umbellus, Lagopus lagopus scoticus, and Falcipennis falcipennis) were directly sequenced. Of the 1020 nucleotide positions, 186 were variable and uniformly distributed over the gene and only 46 were parsimony informative. Most substitutions were synonymous. Replacement substitutions were detected for 15 out of 340 amino acid sites; only four replacements were parsimony informative. The greatest codon bias was found for leucine and serine. The C-T transitions and the G-C transversions were, respectively, the most common (60.7%) and the most rare (5.9%). The mutation frequencies were high at the third codon position (85.2%) and relatively low at the first and the second position. At the third codon position of the species examined, the guanine content was the lowest (3.3%) and the cytosine content was the highest (44.5%). Based on the cytochrome b gene sequences, phylogenetic relationships in the order Galliformes are inferred.     

Temporal dynamics of grouse populations at the southern edge of their distribution

Studies of grouse conducted at northern latitudes have shown that tetraonids frequently exhibit cyclic fluctuations in abundance but little is known about the dynamics of grouse species at the southerly edge of their range. Hunting statistics from four species of grouse based on 30 yr of data collected from 210 hunting areas were examined from the Dolomitic Alps in the province of Trentino. These data were summed to represent 18 time series from discrete mountain groups. Analyses identified cycles of ca 5 yr in the minority of rock ptarmigan Lagopus mutus and hazel grouse Bonasa bonasia populations. These cycles only showed significant negative autocorrelation at half the cycle period and were classified as phase-forgetting quasi-cycles. Cycles were not found in time series of black grouse Tetrao terix or capereaillie Tetrao urogallus. Correcting time series for hunting effort or hunting restrictions tended to increase the proportion of populations that exhibited cycles but no difference in the strength of second order density dependence, A linear first order density-dependent autoregressive model described the dynamics of most of the populations with the exception of a proportion of rock ptarmigan and black grouse populations where a non linear first order model provided the best tit. We compare the findings with studies conducted in Finland and suggest possible reasons for the reduced tendency to cycle in the populations of southern Europe.     

Vulnerability of black grouse hens to goshawk predation: result of food supply or predation facilitation?

The plant cycle hypothesis says that poor-quality food affects both herbivorous voles (Microtinae spp.) and grouse (Tetraonidae spp.) in vole decline years, leading to increased foraging effort in female grouse and thus a higher risk of predation by the goshawk Accipiter gentilis. Poor-quality food (mainly the bilberry Vaccinium myrtillus) for these herbivores is induced by seed masting failure in the previous year, when the bilberry is able to allocate resources for chemical defence (the mast depression hypothesis; MDH). The predation facilitation hypothesis (PFH) in turn states that increased searching activity of vole-eating predators during or after the decline year of voles disturbs incubating and brooding grouse females. The behaviours used by grouse to avoid these terrestrial predators make them more vulnerable to predation by goshawks. We tested the main predictions of the MDH and PFH by collecting long-term (21-year) data from black grouse Tetrao tetrix hens and cocks killed by breeding goshawks supplemented with indices of bilberry crop, vole abundance and small carnivores in the vicinity of Oulu, northern Finland. We did not find obvious support for the prediction of the MDH that there is a negative correlation of bilberry crop in year t with vole abundance and with predation index of black grouse hens in year t + 1. We did find obvious support for the prediction of the PFH that there is a positive correlation between predator abundance and predation index of grouse hens, because the stoat Mustela erminea abundance index was positively related to the predation index of black grouse hens. We suggest that changes in vulnerability of grouse hens may mainly be caused by the guild of vole-eating predators, who shift to alternative prey in the decline phase of the vole cycle, and thus chase grouse hens and chicks to the talons of goshawks and other avian predators.     

Sex and age influence intestinal parasite burden in three boreal grouse species

Parasite infections are often more common in male vertebrates than in females. Sexual selection leading to dimorphism can cause sexual differences in immune defence, behaviour or body size. Possible proximate explanations for male bias in parasitism are the immunosuppressive effect of male sex hormones and the large body size of males which increases the likelihood of being parasitized. To evaluate these hypotheses, we studied the prevalence and abundance of intestinal helminth parasites in three boreal grouse species, the capercaillie Tetrao urogallus , the black grouse Tetrao tetrix , and the hazel grouse Bonasa bonasia . The first two are sexually dimorphic polygynous species while the latter species is sexually monomorphic and monogamous. We found a male-bias in the prevalence and abundance of the nematode Ascaridia compar . The bias was most pronounced in the polygynous black grouse and capercaillie. In the monogamous hazel grouse, there was a slight male-bias in occurrence of ascarids, but no bias in abundance. In juvenile grouse, the male-bias was larger than in adult grouse. No sexual bias was found in regard to the cestodes ( Skrjabinia cesticillus , Paroniella urogalli and Hymenolepis spp.). However, age was a factor in cestode prevalence: juvenile grouse were more commonly infected than adults. Differences in growth rates and body size are potential factors that may lead to male-biased parasitism in these grouse species, and their impact requires further studies.     

Coupling in goshawk and grouse population dynamics in Finland

Different prey species can vary in their significance to a particular predator. In the simplest case, the total available density or biomass of a guild of several prey species might be most relevant to the predator, but behavioural and ecological traits of different prey species can alter the picture. We studied the population dynamics of a predator–prey setting in Finland by fitting first-order log-linear vector autoregressive models to long-term count data from active breeding sites of the northern goshawk (Accipiter gentilis; 1986–2009), and to three of its main prey species (1983–2010): hazel grouse (Bonasa bonasia), black grouse (Tetrao tetrix) and capercaillie (T. urogallus), which belong to the same forest grouse guild and show synchronous fluctuations. Our focus was on modelling the relative significance of prey species and estimating the tightness of predator–prey coupling in order to explain the observed population dynamics, simultaneously accounting for effects of density dependence, winter severity and spatial correlation. We established nine competing candidate models, where different combinations of grouse species affect goshawk dynamics with lags of 1–3 years. Effects of goshawk on grouse were investigated using one model for each grouse species. The most parsimonious model for goshawk indicated separate density effects of hazel grouse and black grouse, and different effects with lags of 1 and 3 years. Capercaillie showed no effects on goshawk populations, while the effect of goshawk on grouse was clearly negative only in capercaillie. Winter severity had significant adverse effects on goshawk and hazel grouse populations. In combination, large-scale goshawk–grouse population dynamics are coupled, but there are no clear mutual effects for any of the individual guild members. In a broader context, our study suggests that pooling data on closely related, synchronously fluctuating prey species can result in the loss of relevant information, rather than increased model parsimony.     

The effect of matrix on the occurrence of hazel grouse (Bonasa bonasia) in isolated habitat fragments

The aim of this study was to determine the effect of matrix on the occurrence of hazel grouse (Bonasa bonasia) in habitat fragments. The study was conducted in two kinds of landscape: (1) an agricultural landscape, where the censused forest habitat fragments were surrounded by farmland, and (2) in an intensively managed forested landscape, where the censused habitat fragments were surrounded by nonhabitat coniferous forest. Occupied and unoccupied habitat fragments in the agricultural landscape differed significantly in distance to the nearest suitable continuous habitat, with hazel grouse occurring only in habitat fragments closer than 100 m from continuous forest. In the intensively managed forest landscape, the effect of isolation was less evident, but there might be a threshold around 2 km. Effects of isolation occurred over much shorter distances when the surrounding habitats consisted of farmland than when it was forested habitats. The size of the habitat fragments was important in both landscapes, with larger habitat fragments more often containing hazel grouse.     

Phylogeny of Tetraoninae and other galliform birds using mitochondrial 12S and ND2 genes

The avian subfamily Tetraoninae (grouse and ptarmigan) is a Holarctic group in the order Galliformes distinguished by morphological adaptations to cold environments and behavioral traits associated with elaborate courtship. Here we investigate the relationships of 17 tetraonines and 12 other galliform species using mitochondrial 12S and ND2 sequence data. We found support for the recent phylogenetic classification that separates the genus Dendragapus into two genera, Falcipennis and Dendragapus. In addition, we found support for a tetraonine clade in which the first divergence is between Bonasa umbellus and all others, followed by divergence between a Bonasa bonasia/Bonasa sewerzowi clade and the remaining tetraonines. Falcipennis canadensis is sister to a clade with four Tetrao species, and the genus Centrocercus is sister to a Dendragapus obscurus/Tympanuchus clade. Our data indicate a basal position for Cracidae and Megapodiidae among the five recognized galliform families. We also found strong support for the monophyly of Phasianidae, although the relative positions of Numididae and Odontiphoridae remains unresolved. We use a maximum likelihood approach to infer ages of 37mya for divergence of Numididae and Phasianidae and 28mya for the divergence of Tetraoninae and Meleagris gallopavo. These estimates must be viewed as tentative as they depend on tests of rates of molecular evolution and accurate fossil dates.     

The importance of alder to hazel grouse in Fennoscandian boreal forest: evidence from four levels of scale

Buds and staminate catkins of alder (Alnus spp) form an important winter food for hazel grouse Bonasa bonasia in the Fennoscandian boreal forest Alder was found to be highly preferred over other deciduous trees, particularly alders ≥ 10 m tall and ≤15 m from spruce forest Winter territories were probably feeding territories, as size was correlated negatively with alder density and almost significantly correlated negatively with competitor density All winter territories were found to contain ample winter food resources for hazel grouse However, the distribution of territories was associated significantly with the distribution of alders at two levels of scale, the territory level and the landscape level Moreover, relationships between the abundance of alders and hazel grouse were found at two additional levels of scale the local patch level and the biogeographic region level This agreement of the results from four levels of scale strongly suggested that the abundance and distribution of alder was a major factor limiting hazel grouse winter territories within dense Norway spruce Picea abies forests in the boreal zone of Fennoscandia Alder was relatively uncommon and exhibited a clumped dispersion pattern at the local and landscape scales, being associated with wet and rich soils The close relationship to alder implies that hazel grouse winter habitats, even in natural forests, also should be distributed patchily Hazel grouse may select the catkins and buds of alder because it is a very nutntous food source, and small species, such as the hazel grouse, require more nutritious food than larger species     

Occupancy modeling of ruffed grouse in the Black Hills National Forest

Ruffed grouse (Bonasa umbellus) are a popular game bird and the management indicator species for quaking aspen (Populus tremuloides) in the Black Hills National Forest (BHNF), which requires development of a robust monitoring protocol to evaluate trends in ruffed grouse populations. We used roadside drumming surveys in spring 2007 and 2008 to estimate ruffed grouse occupancy and detection probabilities in the BHNF while simultaneously assessing the influence of sampling and site covariates on these processes. Ruffed grouse occupancy estimates were constant between spring 2007 and 2008 (Ψ = 0.12, SE = 0.03) and were positively influenced by the amount of aspen surrounding the site. Detection probability estimates were constant between spring 2007 and 2008 (p = 0.27, SE = 0.06) and were influenced by survey date in a quadratic form and negatively influenced by wind speed and time of the survey. Collectively, our results demonstrated that ruffed grouse occupancy and detection probabilities in the BHNF were low. Occupancy could be increased by increasing the extent of aspen. To improve monitoring efficiency and maximize probability of detecting ruffed grouse, ruffed grouse monitoring should be conducted during the peak of drumming (mid-May), during favorable weather conditions such as low wind speeds and little precipitation, and during early morning, near sunrise. © 2010 The Wildlife Society.     

Evaluating spatially explicit viability of a declining ruffed grouse population

Species associated with early successional habitats have experienced dramatic declines in the eastern United States as a result of land use changes and human disruption of natural disturbance regimes. Consequently, active management is required to create early successional habitat and promote plant and animal communities that depend on periodic forest disturbance. Ruffed grouse (Bonasa umbellus) depend on recently disturbed forest habitat, and have experienced dramatic declines over the last half-century. Although ruffed grouse are extensively studied, little effort has been made to link population dynamics with habitat management at landscape scales. We used stochastic, spatially explicit population models that combined landscape conditions derived from a Geographic Information System with demographic data, and applied the model to a declining ruffed grouse population in Rhode Island, USA. We identified vital rates that influence ruffed grouse population dynamics using baseline models constructed with current demographic rates and landscape conditions, and assessed the effect of landscape-scale forest management alternatives on population persistence by running multiple management simulations. Baseline models typically predicted population decline, and we concluded that vital rates (survival and recruitment) had a greater influence on population persistence than did dispersal capability, carrying capacity, or initial population size. Management simulations predicted greater population persistence under a scenario where high-quality habitat was provided in fewer large blocks as opposed to many small blocks, and the rate at which we allowed ruffed grouse to colonize newly created habitat had a substantial impact on management success. Populations of ruffed grouse in the eastern United States are likely to continue to decline given current disturbance regimes, and our work provides a link between ruffed grouse demography and landscape-scale habitat conditions to support management decisions. © 2011 The Wildlife Society.     

长白山花尾榛鸡繁殖成功率的研究

１９８６～１９９５年,对长白山花尾榛鸡的繁殖率进行了研究。无线电遥测的结果表明,花尾榛鸡的孵化率为６５５％（ｎ＝５）,繁殖成功率为２０９％（ｎ＝４）,幼鸟成活率３７５％（ｎ＝３）。花尾榛鸡家族群的幼鸟数随日龄增加而减少,幼鸟日龄为０～２５日时死亡率较高,２５日龄后的死亡率显著减少。１０年花尾榛鸡的繁殖成功率指数的变化与６月的降水量及＞１ｍｍ的降水天数呈显著的负相关（Ｐ＜００５,Ｐ＜００２５）,说明育雏初期降水对幼鸟成活率的影响较大。