Where in Europe should we look for sources of the cutaneous trematode Collyriclum faba infections in migrating birds?

Cutaneous cysts with trematodes of Collyriclum faba have been found in birds during their spring and post-breeding migrations in the Czech Republic. During spring migrations, C. faba was found in one dunnock Prunella modularis, two European robins Erithacus rubecula, three common nightingales Luscinia megarhynchos, one song thrush Turdus philomelos and one great reed warbler Acrocephalus arundinaceus. During post-breeding migration, the same parasite was found in one garden warbler Sylvia borin, one whitehroat S. communis, three goldcrests Regulus regulus and one Eurasian treecreeper Certhia familiaris. The newly identified hosts of C. faba are dunnock, common nightingale, song thrush, great reed warbler and Eurasian treecreeper. Geographical areas of the birds' infection were identified from an analysis of reports on ringed birds of the same species, the time necessary for the development of cutaneous cysts with C. faba and the time of their survival, and hitherto known geographical areas of endemic occurrence of C. faba. It is presumed that birds trapped during spring migrations were infected in some montane and submontane regions in south-western Europe (the Alps, the Apennines). Birds infected during autumn migration or post-breeding vagrancy could have been infected in the Central European Carpathians, the region of C. faba endemic occurrence. For migrating birds, the impact of C. faba infections has not been hitherto assessed.     

Vertebrate host specificity of two avian malaria parasites of the subgenus Novyella: Plasmodium nucleophilum and Plasmodium vaughani

The susceptibility of wild-caught European passeriform birds to naturally isolated malaria parasites, Plasmodium (Novyella) nucleophilum and Plasmodium (Novyella) vaughani, was studied by means of intramuscular subinoculation of infected citrated blood. Plasmodium nucleophilum of the great tit, Parus major, was transmitted to 3 great tits, but 3 blackcaps (Sylvia atricapilla) were not susceptible. Plasmodium vaughani of the robin, Erithacus rubecula, was transmitted to 1 robin and 1 blackcap, but 1 dunnock, Prunella modularis, was not susceptible. The prepatent period was between 8 and 10 days in all experimental infections. Maximum experimental parasitemia (3.4% of red cells) was detected in great tits infected with P. nucleophilum 23 days postexposure. A light (<0.01%) transient parasitemia of P. vaughani developed in the robin and blackcap. This study is in accord with former experimental observations on host specificity of P. nucleophilum and P. vaughani, which are characterized by a wide, but selective, range of avian hosts. Two new host-parasite associations were recorded.     

Low frequency of extrapair paternity in the common redstart ( Phoenicurus phoenicurus )

Social monogamy is common among passerines whereas genetic monogamy is regarded as rare. Here we report the first study of paternity in the genus Phoenicurus (Aves, Muscicapidae). In the socially monogamous common redstart (Phoenicurus phoenicurus) 2.0% (5/253) of offspring in 10.5% (4/38) of the broods examined were sired by extrapair males. The observed pattern of extrapair paternity was consistent among two geographically separate nest-box populations sampled over two years. Our findings reveal low rates of extrapair paternity compared with other passerine birds, suggesting only a minor role of sperm competition in this sexually dichromatic species.     

Postexercise ketosis in night-migrating passerine birds

This study investigated the postexercise metabolism of six species of free-living, night-migrating passerine birds (European robin, pied flycatcher, wheatear, redstart, blackcap, and garden warbler). The birds were caught during autumn migration out of their nocturnal flight, and their metabolism changed from a fasting, highly active state to a fasting, resting state. Concentrations of six plasma metabolites of the fat, carbohydrate, and protein metabolism were measured during up to 10 h of recovery time. The metabolic changes indicated a biphasic pattern: (a) a quick first response to the reduced energy demands during the first 20 min of recovery, suggested by an increase and subsequent decrease of free fatty acid levels, and (b) subsequently, a postexercise ketosis and a reduction of lipolysis and proteolysis, suggested by high beta-hydroxy-butyrate and low free fatty acid, glycerol, triglyceride, and uric acid levels. This metabolic pattern differs from that of humans and rats, in which ketosis starts immediately postexercise or is absent in trained subjects. Since migrating birds are naturally adapted to endurance exercise, it is hypothesized that the high and long-lasting postexercise ketosis does not evoke physiological problems (such as hypoglycemia) but, by contrast, increases the ability of birds to rely on lipids, to a very high extent, during and after flight and decreases the dependence on glucose and glucogenic amino acids. Differences between species in fat stores and metabolic pattern support this hypothesis.     

Arrival and onset of breeding of three passerine birds in eastern Finland tracks climatic variation and phenology of insects

Anthropogenic climate change poses a challenge to the annual cycles of migratory birds. It has become urgent to understand whether migratory birds are able to advance their spring phenology when the climate is warming and whether they are able to adjust these phenological phases to the spring phenology in their breeding areas. In this work, we studied long‐term trends in first arrival and onset of breeding for three passerine birds in eastern Finland; the pied flycatcher Ficedula hypoleuca, the common redstart Phoenicurus phoenicurus and the great tit Parus major. The pied flycatcher and the common redstart are long‐distance migrants while the great tit is a partial migrant in Finland. We asked what environmental variables best explain the first arrival or onset of breeding, if there is evidence of ‘thermal delay’ (long‐term increase in the accumulated temperatures) at arrival or onset of breeding and if the interannual variation in the onset of breeding correlates with variation in spring phenology of local insects. We found that the pied flycatcher and the common redstart had advanced their first arrival (explained by increased temperatures at the migration route), but we found no long‐term change in the onset of breeding (explained by local temperatures). Also, the onset of breeding of the great tit is tracking local temperatures. We found no or only weak evidence of thermal delay at arrival or onset of breeding for any of the species. The onsets of breeding for the pied flycatcher and the great tit are also closely tracking the spring phenology of the local insects. The stable or increasing population sizes of all three species in Finland could be a result from their ability to effectively track climatic and environmental variation.     

Leucocytozoonosis and trypanosomiasis in redstarts in Finland.

Leucocytozoon spp. and Trypanosoma spp. blood parasites in the redstart (Phoenicurus phoenicurus) were studied during spring migration 1994 in southern Finland (53 individuals) and the breeding season 1992-1994 in northern Finland (69). Parasite prevalence was higher during the breeding season (48%) than during the migration period (13%), with no age or sex differences in the breeding site birds. In both periods, redstarts were infected by the same blood parasites Leucocytozoon shaartusicum (46% prevalence at the breeding site and 71% during the migration period) and Trypanosoma avium, complex (58% and 43%, respectively). One individual at the breeding site had contracted L. dubreuili and one at the stop-over site had T. everetti. Our results may support the assumption that tissue-hidden parasites relapse during the breeding season when birds may have diminished immune response related to egg production and brood rearing. Another explanation could be that the high abundance of ornithophilic vectors enhance parasite transmission during breeding season in northern Finland.     

Nest biology of urban populations of cavity-nesting birds

Nesting features of four species of cavity-nesting birds—common starling (Sturnus vulgaris L.), great tit (Parus major L.), common redstart (Phoenicurus phoenicurus L.), and pied flycatcher (Ficedula hypoleuca Pall.)—have been studied in city. Under urban conditions, number of eggs in a nest of a redstart is significantly larger, while for great tit and pied flycatcher it is significantly smaller; differences between the populations of starling are insignificant. Success of reproduction for the cavity-nesters is significantly higher in a city and exceeds the reference value for all the species by 10–20%. The number of chicks per a reproduction attempt is also higher in city for all the species. Starling population in city exceeds the reference population in the success of reproduction mainly due to a decrease in the percentage of abandoned eggs. The main reasons for an increase in the reproduction success of other species at the urbanized locality are weakened pressure of predators and a decrease in the percentage of abandoned broods.     

Response of introduced European birds in New Zealand to experimental brood parasitism

The loss of anti-parasite adaptations against the European cuckoo Cuculus canorus was studied in three European passerine species, song thrush Turdus philomelos , blackbird T. merula , and chaffinch Fringilla coelebs , introduced to New Zealand in the 19th century. Chaffinches in New Zealand ejected non-mimetic eggs at a rate similar to their source population in the United Kingdom, but both song thrushes and blackbirds in New Zealand rejected non-mimetic eggs at a higher rate than the United Kingdom. It is not clear if this difference reflects variation among hosts in their response to brood parasitism or if it is an artefact of subtle differences in the types of non-mimetic eggs tested. In contrast, all three introduced species showed little aggression to a taxidermic model of a European cuckoo presented at their nests. This differs from European populations of these species, where model cuckoos are typically attacked. Our results suggest that in the∼130 years since their release in New Zealand, introduced birds have lost recognition of the European cuckoo but not their ability to discriminate non-mimetic eggs. The differential loss of anti-parasite adaptations by introduced birds in New Zealand suggests that cyclical models of host/parasite co-evolution may need to take into account the differing rates at which different host adaptations are lost and gained.     

Synchrony of woodland bird populations: the effect of landscape structure

The influence of environmental stochasticity and dispersal in producing patterns in population synchrony was examined for 53 woods censused annually from 1990 to 1999 for nine resident bird species (wren Troglodytes troglodytes , dunnock Prunella modularis , robin Erithacus rubecula , blackbird Turdus merula , song thrush Turdus philomelos , long-tailed tit Aegithalos caudatus , blue tit Parus caeruleus , great tit Parus major , and chaffinch Fringilla coelebs ) and four migrant bird species (garden warbler Sylvia borin , blackcap Sylvia atricapilla , chiffchaff Phylloscopus collybita and willow warbler Phylloscopus trochilus ). Twelve species showed global synchrony of population counts due to regional population trends and widespread annual population fluctuations. There was a clear link between population fluctuations and winter weather for wren, and three other species showed their largest population declines after the coldest winters. Eight species showed a decline in synchrony with distance between woods, and there was evidence for dispersal causing this pattern in three species. Landscape structure affected patterns of synchrony in several species, with lower synchrony in landscapes with less woodland. For three species, this difference in synchrony across a landscape gradient of decreasing woodland cover accounted for the decline in synchrony over distance. Three species showed greater synchrony between woods with similar amounts of hedgerow in the surrounding landscape, suggesting that the surroundings of a wood influence the population dynamics of some species breeding in the wood. Habitat fragmentation can alter the processes contributing to population synchrony. Loss of woodland reduces the relative abundance of woodland bird species. The remaining patches of habitat are smaller, more isolated and are set in a more hostile landscape, all of which may disrupt dispersal between patches and alter the population dynamics within woods.     

Fruit tracking between sites and years by birds in Mediterranean wintering grounds

Knowledge of the ability of birds to track spatiotemporal variation in fruit distribution is essential for understanding plant-frugivore interactions. Arguably, although total fruit availability sets an upper limit to the number of birds that can exploit a habitat patch, not all species can equally distribute abundance according to variation in fruit resources. To explore this, we studied bird and fruit abundance in 1999–2005 in Mediterranean scrublands and woodlands of southern Spain. We analysed whether changes of fruit abundance in eight different sites during six winters could predict numerical changes of a set of frugivorous passerines of the area (blackcap Sylvia atricapilla , Sardinian warbler S. melanocephala , robin Erithacus rubecula , song thrush Turdus philomelos and blackbird T. merula ). We also investigated if all frugivores together tracked fruits better than individual species, thereby supporting a shared use of resources. Results showed strong inter-specific differences. Only the most abundant species (blackcaps and robins) tracked the spatial patterning of food despite strong differences in the use of space (vagrant and territorial, respectively). This suggests plastic behaviour of territorial robins, with individuals changing from strictly territorial to wandering, a flexibility that would favour between-site numerical arrangements according to food resources. Annual changes in bird numbers were independent of the availability of fruits, except for blackcaps, an abundant vagrant bird that tracked inter-winter changes in fruit abundance. The abundance of blackcaps fitted the spatiotemporal patterning of fruit resources better than the whole guild of frugivorous birds, inconsistent with the idea that these species track together the changing availability of fruit resources.     

Egg colour mimicry in the common cuckoo Cuculus canorus as revealed by modelling host retinal function

Some parasite cuckoo species lay eggs that, to the human eye, appear to mimic the appearance of the eggs of their favourite hosts, which hinders discrimination and removal of their eggs by host species. Hitherto, perception of cuckoo-host egg mimicry has been estimated based on human vision or spectrophotometry, which does not account for what the receivers' eye (i.e. hosts) actually discriminates. Using a discrimination model approach that reproduces host retinal functioning, and museum egg collections collected in the south of Finland, where at least six different races of the European cuckoo (Cuculus canorus) coexist, I first assess whether the colour design of cuckoo eggs of different races maximizes matching for two favourite avian hosts, viz. the redstart (Phoenicurus phoenicurus) and the pied wagtail (Motacilla alba). Second, I assess the role of nest luminosity on host perception of mimicry by the same two hosts. Phoenicurus-cuckoo eggs showed a better chromatic matching with the redstart-host eggs than other cuckoo races, and in most cases can not be discriminated. Sylvia-cuckoo eggs, however, showed better achromatic matching with redstart-host eggs than Phoenicurus-cuckoo eggs. Also, Motacilla-cuckoo eggs showed poorer chromatic and achromatic matching with pied wagtail-host eggs than Sylvia-cuckoo eggs. Nest luminosity affected chromatic and achromatic differences between cuckoo and host eggs, although only minimally affected the proportion of cuckoo eggs discriminated by chromatic signals. These results reveal that cuckoo races as assessed by humans do not entirely match with host perception of matching and that achromatic mechanisms could play a main role in the discrimination of cuckoo eggs at low-light levels.     

Breeding bird communities on Scottish farms

The Common Birds Census, begun by the British Trust for Ornithology has now been monitoring bird population levels on a nationwide basis since 1962. Separate indices were constructed for Scotland and the rest of Great Britain for 12 of the commonest species. Of the species examined blue tit (Parus caeruleus), wren (Troglodytes troglodytes), song thrush (Turdus philomelos), lapwing (Vanellus vanellus), chaffinch (Fringilla coelebs) and skylark (Alauda arvensis) have increased since 1963 and show very similar population trends to the rest of Great Britain. Whitethroat (Sylvia communis), willow warbler (Phylloscopus trochilus) and swallow (Hirundo rustica) have all declined nationally, the decline in the swallow appears to have begun earlier in Scotland. The blackbird (Turdus merula) and dunnock (Prunella modularis) showed similar upward trends until 1972 when the Scottish population started to decline; a much more noticeable decline in the Scottish robin (Erithacus rubecula) population began in 1966. The total density of territory-holding males has not increased but the biomass has and this is due to more, bigger birds but fewer passerines. The species diversity measured by the Shannon Wiener index (H.) has improved a little since 1971 and it is concluded that during the study period the farmland plots censused in Scotland have on average not shown evidence of deterioration as habitats for territory-holding birds.     

Does Gray Squirrel (Sciurus carolinensis) Response to Heterospecific Alarm Calls Depend on Familiarity or Acoustic Similarity?

In habitats in which multiple species are prey to the same predators, individuals can greatly benefit from recognizing information regarding predators that is provided by other species. Past studies have demonstrated that various mammals respond to familiar heterospecific alarm calls, but whether acoustic similarity to a familiar call can prompt a mammal's recognition of an unfamiliar call has yet to be shown. We presented alarm calls to free‐ranging eastern gray squirrels (Sciurus carolinensis) and recorded behavioral changes in vigilance and antipredatory response. Playbacks included alarm calls of a sympatric bird (American robin, Turdus migratorius), an allopatric bird with a call structure similar to that of the robin (common blackbird, Turdus merula), and an allopatric bird with a distinct call structure (New Holland honeyeater, Phylidonyris novaehollandiae). Squirrels responded significantly more frequently to squirrel alarm calls (positive control) than to robin song (negative control) or honeyeater calls. Squirrel response to robin and blackbird alarm calls was statistically similar to their response to squirrel alarm calls, indicating that squirrels responded to those alarm calls as if they provided information about the presence of predators. However, squirrel response to robin song was not statistically different from response to any of the other avian calls, including the robin and blackbird alarms, suggesting that squirrels neither respond to blackbird alarms as if they clearly signify danger, nor as if they clearly do not signify danger, perhaps reflecting some ambiguity in interpretation of the calls. These results suggest that squirrel responses to alarm calls are generally based on call familiarity, but that acoustic similarity of an unfamiliar allopatric call to a familiar call also can elicit antipredator behavior. The lack of response to honeyeater alarm calls also supports the hypothesis that call recognition by gray squirrels is dependent on familiarity, rather than simply detection of an acoustic feature common to alarm calls across a variety of avian species.     

东北地区北红尾鸲巢址选择及繁殖成效

北红尾鸲(Phoenicurus auroreus)是一种分布广泛的小型雀形目鸟类,主要分布于南亚东北部,东南亚北部、东亚及俄罗斯等地区,我国东北、华北、华中至西南等地也均有分布,是重要的食虫益鸟。为探究北红尾鸲巢址选择的影响因素,找到影响北红尾鸲繁殖成功率的主要巢址因子,于2017年4—7月,在辽宁仙人洞国家级自然保护区开展系统研究。共发现北红尾鸲自然巢44个,其中29巢繁殖成功,15巢繁殖失败。北红尾鸲主要筑巢于石墙缝、空心砖墙缝及废旧电表箱中。巢址参数的主成分分析结果表明:巢口因子(27.738%)、巢位因子(14.195%)、光照因子(12.145%)、人为干扰因子(10.440%)、安全因子(9.266%)和隐蔽因子(7.187%)是影响北红尾鸲巢址选择的重要因子。采用二元逻辑斯蒂回归分析繁殖成功巢与失败巢参数发现,成功巢的巢口最大高度显著小于失败巢(P=0.047),且其距顶的距离更近(P=0.043)。多元线性回归分析表明,巢上方盖度对繁殖成功率有极显著影响(t=2.883,P=0.009)。总的来说,北红尾鸲虽偏爱筑巢于人为干扰较大的村庄房屋附近,但较小的巢口能有效避免巢捕食者的捕食,更近的距顶距离和更大的巢上方盖度能有效降低巢上方的可视程度和降水等不利因素的影响,从而提高繁殖成功率。     

Deep sympatric mitochondrial divergence without reproductive isolation in the common redstart Phoenicurus phoenicurus

Mitochondrial DNA usually shows low sequence variation within and high sequence divergence among species, which makes it a useful marker for phylogenetic inference and DNA barcoding. A previous study on the common redstart (Phoenicurus phoenicurus) revealed two very different mtDNA haplogroups (5% K2P distance). This divergence is comparable to that among many sister species; however, both haplogroups coexist and interbreed in Europe today. Herein, we describe the phylogeographic pattern of these lineages and test hypotheses for how such high diversity in mtDNA has evolved. We found no evidence for mitochondrial pseudogenes confirming that both haplotypes are of mitochondrial origin. When testing for possible reproductive barriers, we found no evidence for lineage‐specific assortative mating and no difference in sperm morphology, indicating that they are not examples of cryptic species, nor likely to reflect the early stages of speciation. A gene tree based on a short fragment of cytochrome c oxidase subunit 1 from the common redstart and 10 other Phoenicurus species, showed no introgression from any of the extant congenerics. However, introgression from an extinct congeneric cannot be excluded. Sequences from two nuclear introns did not show a similar differentiation into two distinct groups. Mismatch distributions indicated that the lineages have undergone similar demographic changes. Taken together, these results confirm that deeply divergent mitochondrial lineages can coexist in biological species. Sympatric mtDNA divergences are relatively rare in birds, but the fact that they occur argues against the use of threshold mtDNA divergences in species delineation.     

Begging in Common Redstart Nestlings: Scramble Competition or Signalling of Need?

Begging behaviour by the young affects parental food distribution among nestlings of altricial birds. We present an analysis of two types of begging behaviour (assuming the front nest positions and gaping) based on videotaped natural nestling feeding in European common redstart (Phoenicurus phoenicurus). We test whether these types of begging support the predictions of two mathematical models: scramble competition with competitive asymmetries between nestlings [Anim. Behav. 27 (1979) 1210] or honest signalling model [Nature 352 (1991) 328]. None of the measured variables of nestling or parental behaviour were affected by body weight differences between siblings. In contrast, both gaping and nest positioning were affected by individual differences in nestling hunger. In agreement with the honest signalling model, hungrier nestlings gaped with higher probability and started to gape sooner after the arrival of the parent than did their less hungry nestmates. Those nestlings with the shortest latency to gape also received food more often. Nest positioning was related to nestling hunger in a way unforeseen by the existing models. The intervals between nestling position changes were several times longer than the intervals between parental feeding visits, and parents preferred to feed nestlings in front positions, so nestlings in front positions were always less hungry than nestlings in back. Hence the pattern of movements influenced the feeding decision in favour of the more satiated nestlings and acted against the effect of gaping. Nestling movement seemed to be caused by the less hungry nestlings moving actively from front to rear positions. Low mortality of individual nestlings within broods that survived to fledging and small within‐brood variation in fledging weights indicated low competition among nestmates. We suggest that there are two behavioural mechanisms that contribute to the equalization of fledging weights in common redstart nestlings: the signalling of need through gaping and the regular turnover of nestlings at front positions.     

A survey for Yersinia pseudotuberculosis in migratory birds in coastal Japan

Yersinia pseudotuberculosis was isolated from three specimens of two species of birds, the black-faced bunting (Emberiza spodocephala) and pied wagtail (Motacilla alba), of 528 specimens of birds examined from coastal regions in Japan. The two isolated strains of Y. pseudotuberculosis were identified as serovar 4b and serovar 3. This is the first isolation of Y. pseudotuberculosis from birds in Japan. Yersinia enterocolitica was isolated from three specimens of the pied wagtail, one specimen of the reed bunting (Emberiza schoeniclus) and one specimen of the rustic bunting (Emberiza rustica). Yersinia frederiksenii was isolated from two specimens of the gray-rumped sandpiper (Heteroscelus brevipes). Yersinia intermedia was isolated from one specimen of the pied wagtail.     

Cuckoo parasitism in a cavity nesting host: near absent egg‐rejection in a northern redstart population under heavy apparent (but low effective) brood parasitism

Brood parasite – host systems continue to offer insights into species coevolution. A notable system is the redstart Phoenicurus phoenicurus parasitized by the ‘redstart‐cuckoo’ Cuculus canorus gens. Redstarts are the only regular cuckoo hosts that breed in cavities, which challenges adult cuckoos in egg laying and cuckoo chicks in host eviction. We investigated parasitism in this system and found high overall parasitism rates (31.1% of 360 redstart nests), but also that only 33.1% of parasitism events (49 of 148 eggs) were successful in laying eggs into redstart nest cups. The majority of cuckoo eggs were mislaid and found on the rim of the nest; outside the nest cup. All available evidence suggests these eggs were not ejected by hosts. The effective parasitism rate was therefore only 12.8% of redstart nests. Redstarts responded to natural parasitism by deserting their nests in 13.0% of cases, compared to desertion rates of 2.8% for non‐parasitized nests. Our egg parasitism experiments found low rates (12.2%) of rejection of artificial non‐mimetic cuckoo eggs. Artificial mimetic and real cuckoo eggs added to nests were rejected at even lower rates, and were always rejected via desertion. Under natural conditions, only 21 cuckoo chicks fledged of 150 cuckoo eggs laid. Adding to this low success, is that cuckoo chicks are sometimes unable to evict all host young, and were more likely to die as a result compared to cuckoo chicks reared alone. This low success seems to be mainly due to the cavity nesting strategy of the redstart which is a challenging obstacle for the cuckoo. The redstart‐cuckoo system appears to be a fruitful model system and we suggest much more emphasis should be placed on frontline defences such as nest site selection strategies when investigating brood parasite–host coevolution.     

Unusual finding of Trichobilharzia sp. in Motacilla alba in the Czech Republic

Adult male worms of Trichobilharzia sp. recovered from a pied wagtail (Motacilla alba) in the Czech Republic were found to belong to a new species of the genus. The finding of Trichobilharzia sp. in a passeriform bird in Europe represents an important discovery, as only anseriform birds have thus far been reported as final hosts of the European Trichobilharzia species.     

Genetic diversity and morphological variation of the common chaffinch Fringilla coelebs in the Azores

We present new insights into the genetic diversity and phylogeography of the common chaffinch Fringilla coelebs from the Azores, based on sequences of mitochondrial and nuclear genes from 44 individuals and an outgroup/comparison of 44 birds from Madeira, the Canary Islands and the Continental Western Palearctic. To understand the level of concordance between the genetic data and morphometric variability we analysed eight morphometric characters from 413 adult living birds from all the Azores islands and compared the population genetic distances with quantitative morphometric traits. Our results indicate the occurrence of gene flow among the common chaffinch populations in the archipelago revealing the lack of current genetic structure within it and the existence of two co‐occurring lineages. Results also indicate the existence of morphometric differences among islands that could be due to ecological features instead of island isolation. This study also confirms the genetic distance among the common chaffinch populations within Macaronesia and between these archipelagos and the Continental Western Palearctic.