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Abb. 4: Komplex der Schafstelzen Motacilla flava sensu lato Nord-Eurasiens (männl., wbl.): flava, thunbergi, beema, zaissanensis, leucocephala, feldegg, melanogrisea, flavissima, lutea, plexa, tschutschensis, simillima, taivana und macronyx. – The Yellow Wagtail complex Motacilla flava sensu lato North Eurasia (m., f.): flava, thunbergi, beema, zaissanensis, leucocephala, feldegg, melanogrisea, flavissima, lutea, plexa, tschutschensis, simillima, taivana and macronyx.
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Species or not? Contentious taxonomic opinions on the birds of North Eurasia
The article presents the position of the leading Russian taxonomists in the current discussion of the
category ‘species’, as well as the biological and phylogenetic species concept (BSC, PSC). This alternative
draft concept differs significantly from the standpoint of West...
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Citations
... Zaissanensis is treated as synonymous with tschutschensis by some authorities, eg, AOU (Banks et al 2004) and IOC (following Alström & Mild 2003), while Red'kin (2000, 2001 groups it with Western Yellow Wagtail. It seems that it is geographically isolated from all Eastern Yellow Wagtail subspe cies, as suggested by Johansen (1944) and also in dicated in the map in Red'kin et al (2016). While the male head pattern in zaissanensis and tschutschensis is similar, zaissanensis differs in softer calls, westerntype songs, shorter hind claw length 501 Yellow wagtail / gele kwikstaart Motacilla flava sensu lato, male, Gornozavadovsk, Sakhalin, Russia, 9 July 1993 (John Klicka/UWBM). ...
To summarise the main findings of this study, the variation in sampled birds across Arctic regions suggests introgression between thunbergi and tschutschensis between Lower Ob and Lower Kolyma, challenging the division into two separate yellow wagtail species. Overlap between birds from different regions makes it impossible to identify geographic origins of out of range individuals based on measurements and plumage features examined here. However, average differences may provide clues, and when combined with recordings of series of typical contact calls and song, it is possible to assign birds with reasonable certainty to three areas: 1 Scandinavia and north-western Russia: soft calls that are often slightly disyllabic; song either single note type or slow variants of fast type; 2 The area between Yamal/Ob river basin and Taimyr: sharper calls without r-sound and calls with prominent frequency dip midway; song either single-note type or fast type at moderate pace; 3 The area east of Taimyr and in Alaska: sharp Citrine Wagtail-like calls with r-sound; song most likely only a fast type at high pace (more study of song needed in area between Eastern Taimyr and Lower Kolyma). Based on examined features, it is not possible to separate birds breeding east of Taimyr to Lower Kolyma, ie, within the alleged range of plexa, from tschutschensis with enough confidence to meet the 75% rule for subspecific recognition, although males without a supercilium are regular in Lower Kolyma and westwards while being rare in tschutschensis.
For the more southern taxa, the findings corroborate earlier studies showing only slight average differences between simillima and tschutschensis, and more pronounced and consistent average differences in plumage between macronyx and Arctic dark-headed birds. No convincing evidence of the existence of the alleged taxon angarensis has been found. Finally, the results suggest that zaissanensis is a fairly distinct form of Western Yellow Wagtail. Its consistent plumage and separate breeding range in the Altai mountains suggest that it is a valid subspecies (Yaroslav Red’kin in litt), although further study is needed to firmly rule out the possibility of an intergrade population.
From a European field identification perspective, the findings confirm that recordings of calls are essential when dealing with suspected vagrant eastern yellow wagtails. Actually, in Arctic populations, calls seem to provide a more precise indication of geographic origin than the mtDNA haplotypes upon which publication of Eastern Yellow Wagtails records in Europe has relied heavily so far. As long as the status of plexa is unresolved, assignment of north-eastern birds to a specific taxon will however remain problematic.
... L'averla maggiore Lanius excubitor Linnaeus, 1758 è una specie polimorfica alla quale attualmente vengono attribuite 12 sottospecie (Yosef & International Shrike Working Group, 2017). La sistematica di questa specie ha subìto profonde revisioni negli ultimi anni e sono state proposte diverse interpretazioni non sempre concordanti (Olsson et al., 2010;Poelstra, 2010;Red'kin et al., 2016); ciò ha portato a notevoli disaccordi nomenclatoriali. ...
[Lanius excubitor is a polymorphic species with 12 subspecies. In the last years the taxonomy of this species has been reviewed several times. According to the literature, in Italy Lanius excubitor is a regular migrator, wintering and irregular nesting, occurring with three subspecies: L. excubitor excubitor, L. e. homeyeri and L. e. sibiricus. This paper aims at reviewing the subspecies actually occurring in Italy by analyzing the specimens preserved in the Arrigoni degli Oddi’s collection at the Museo Civico di Zoologia in Rome (Italy). The morphometrics and colouring of 66 specimens were examined in relation to the most recent literature data. This review allowed to demonstrate that all specimens described as L. e. homeyeri in this collection have to be attributed to the nominate subspecies; only the specimen described as L. przelwaskii (now L. e. leucopterus) is a true L. e. homeyeri. According to these data, the ssp. homeyeri must be considered very rare in Italy, contrarily to some previously reported information. Specimens ascribed to L. borealis sibiricus do not show the characters of this subspecies, consequently they should not be included in the check list of Italian species.]
[Article in Italian]
... L'averla maggiore Lanius excubitor Linnaeus, 1758 è una specie polimorfica alla quale attualmente vengono attribuite 12 sottospecie (Yosef & International Shrike Working Group, 2017). La sistematica di questa specie ha subìto profonde revisioni negli ultimi anni e sono state proposte diverse interpretazioni non sempre concordanti (Olsson et al., 2010;Poelstra, 2010;Red'kin et al., 2016); ciò ha portato a notevoli disaccordi nomenclatoriali. ...
The Great grey Shrike Lanius excubitor L. of the Arrigoni degli Oddi's ornithological collection at the Museo Civico di Zoologia, Rome (Italy). Lanius excubitor is a polymorphic species with 12 subspecies. In the last years the taxonomy of this species has been reviewed several times. According to the literature, in Italy Lanius excubitor is a regular migra-tor, wintering and irregular nesting, occurring with three subspecies: L. excubitor excubitor, L. e. homeyeri and L. e. sibiricus. This paper aims at reviewing the subspecies actually occurring in Italy by analyzing the specimens preserved in the Arrigoni degli Oddi's collection at the Museo Civico di Zoologia in Rome (Italy). The morphometrics and colouring of 66 specimens were examined in relation to the most recent literature data. This review allowed to demonstrate that all specimens described as L. e. homeyeri in this collection have to be attributed to the nominate subspecies; only the specimen described as L. przelwaskii (now L. e. leu-copterus) is a true L. e. homeyeri. According to these data, the ssp. hom-eyeri must be considered very rare in Italy, contrarily to some previously reported information. Specimens ascribed to L. borealis sibiricus do not show the characters of this subspecies, consequently they should not be included in the check list of Italian species. Riassunto-Lanius excubitor è una specie polimorfica con 12 sot-tospecie, la cui tassonomia è stata più volte rivista negli ultimi anni. Secondo la letteratura, in Italia Lanius excubitor è un migratore rego-lare, svernante e nidificante irregolare, che compare con tre sottospecie: L. excubitor excubitor, L. e. homeyeri e L. e. sibiricus. Questo lavoro intende rivedere le sottospecie effettivamente presenti in Italia analiz-zando i reperti conservati nella collezione Arrigoni degli Oddi presso il Museo Civico di Zoologia di Roma. La morfometria e la colorazione di 66 campioni sono stati esaminati in relazione ai dati di letteratura più recenti. Questa revisione ha permesso di dimostrare che tutti gli esemplari descritti come L. e. homeyeri in questa raccolta devono essere attribuiti alla sottospecie nominale; solo l'esemplare descritto come L. przelwaskii (ora L. e. leucopterus) è un vero L. e. homeyeri. Secondo questi dati, la ssp. homeyeri deve essere considerata molto rara in Italia, contrariamente ad alcune informazioni precedentemente riportate. I campioni ascritti a L. borealis sibiricus non mostrano i caratteri di questa sottospecie, pertanto non dovrebbero essere inclusi nell'elenco delle specie italiane. Parole chiave: averla maggiore, Lanius excubitor L., collezione ornitologica Arrigoni degli Oddi.
Background
Field studies from 2011 onwards have demonstrated the presence of a breeding population of Yellow Wagtails ( Motacilla flava ) in the Xinjiang Uygur Autonomous Region, China that is phenotypically distinct from known subspecies occurring in Asia. Here we describe the plumages and vocalisations of this population and discuss its taxonomic status.
Methods
The analysis of plumage is based on field studies and photos available online. Recordings of vocalisations are compared with recordings from other Yellow Wagtail populations, and differences are analysed based on sonograms. Mitochondrial DNA from one individual is compared to other Yellow Wagtail taxa.
Results
Unlike M. flava subspecies breeding in or near Xinjiang, males in the studied population show a blue-grey head without prominent white supercilium, being most similar to the widely disjunct M. f. cinereocapilla . They differ from the similarly widely allopatric M. f. thunbergi , which might occur as a migrant or vagrant in Xinjiang, by on average cleaner yellow breast and more extensive white on the throat, and from the widely disjunct M. f. plexa and M. f. macronyx , which might also occur on migration in that area, by softer contact calls and slower pace of song. Females are similar to female M. f. feldegg in plumage. The mitochondrial ND2 tree shows the single sample from Xinjiang to be nested in the clade of western Yellow Wagtail taxa.
Conclusion
We discuss whether the Xinjiang breeding population could represent an intergrade between subspecies breeding nearby, or whether it is better regarded as a separate as yet unrecognized subspecies. We argue that the localization of its apparent range in relation to other subspecies along with fairly consistent male and female plumages suggest that it is more likely to represent an undescribed taxon, but conclude that more research is needed to firmly establish its status.
As it has presently become evident, reproductive isolation can no longer be considered as the leading criterion for estimating the species limits, but it is still used for determining the status of the majority of morphs. The presence/absence of hybridization between individuals is not directly related to the degree of their evolutionary insularity. Hybridization can result in the origin of new morphs including those that have species status. The application of phylogenetic methods is justified in reconstructing the relation links within complex groups including the morphs of various evolutionary levels (from geographical races to "good" species), the relationships between which are exacerbated by hybridogenous polymorphism and/or the hybrid-ization origins of their populations. Taking into account both new data and new conceptions, we suggest genuine interrelated definitions of concepts such as species, subspecies, and semispecies in birds. The definitions are based on two main criteria: biological, i.e., an evaluation of the reproductive relations of the particular morphs with each other, and phylogenetic, i.e., an evaluation of their evolutionary age and kinship. The main feature of a species as an evolutionary entity should be considered through its stability in time even when its reproductive isolation is periodically broken. Geographic intraspecies races show sustainable variations of different degrees, but they have no reproductive isolation; they breed upon contact and form intergradation zones. They are taxonomically denoted as subspecies. Descriptions of new subspecies are viable to the limits that reflect the species' natural geographic structure to the fullest extent. For the young morphs that have reached the level of species insularity in the course of evolution, it seems appropriate to restore a semispecies category. Semispecies show significant morphological differences and distinguished ecological particulari-ties, as a rule, but they are connected to closely related morphs by gene flows in contact zones. Distinguishing this category is not regulated by the International Code of Zoological Nomenclature and, as splitter tendencies prevail in modern systematics, semispecies are more often equated to species; i.e., they have binominal names. We propose to denote the attribution of a semispecies to a particular species group (superspecies) in parentheses between genus and species names. Thus, it would become possible to outline natural complexes and avoid a groundless increase in the taxonomic statuses of morphs in the stage of development. Representatives of distant phylogenetic lines (morphs that separated historically long ago) are not to be considered as semispecies even in the cases of their reproductive isolation being broken and a steady hybridization existing between them.
As it has presently become evident, reproductive isolation can no longer be considered as the leading criterion for estimating the species limits, but it is still used for determining the status of the majority of morphs. The presence/absence of hybridization between individuals is not directly related to the degree of their evolutionary insularity. Hybridization can result in the origin of new morphs including those that have a species status. The application of phylogenetic methods is justified in reconstructing the relation links within complex groups including the morphs of various evolutionary levels (from geographical races to “good” species), the relationships between which are exacerbated by hybridogenous polymorphism and/or the hybridization origins of their populations. Taking into account both new data and conceptions we suggest genuine interrelated definitions of such concepts as species, subspecies and semispecies in birds. The definitions are based on two main criteria: biological, i.e. an evaluation of the reproductive relations of the particular morphs with each other, and phylogenetic, i.e. an evaluation of their evolutionary age and kinship. The main feature of a species as an evolutionary entity is to be considered through its stability in time even when its reproductive isolation is periodically broken. Geographic intraspecies races show sustainable variations of different degrees, but they have no reproductive isolation, they breed at contact and form intergradation zones. They are taxonomically denoted as subspecies. Descriptions of new subspecies are viable to the limits that reflect the species’ natural geographic structure to the fullest extent. For the young morphs that have reached the level of species insularity in the course of evolution, it seems appropriate to restore a semispecies category. Semispecies show significant morphological differences and distinguished ecological particularities, as a rule, but they are connected to closely related morphs by gene flows in contact zones. Distinguishing this category is not regulated by the International Code of Zoological Nomenclature and, as splitter tendencies prevail in modern systematics, semispecies are more often equated to species, i.e. they have binominal names. We propose to denote the attribution of a semispecies to a particular species group (superspecies) in parentheses between genus and species names. Thus it would become possible to outline natural complexes and avoid a groundless increase in taxonomic statuses of morphs in the stage of development. Representatives of distant phylogenetic lines (the morphs that separated historically long ago) are not to be considered as semispecies even in the cases of their reproductive isolation being broken and a steady hybridization existing between them.
The common stonechat Saxicola torquatus traditionally was considered as a polytypic species widely distributed in Africa, Europe and
Asia. Recently, several authors have suggested that this formerly single species needs to be split into several distinct species composing
the Saxicola torquatus complex based on mitochondrial markers. However, mitochondrial DNA alone is not sufficient for the evaluation
of species status. In this paper, we reviewed the taxonomy of Northern Eurasian taxa from the complex based on morphometrics, plumage,
song and alarm calls. The morphological and vocal data clearly matched the phylogroups reconstructed from mitochondrial DNA
sequences, and separated Northern Eurasian taxa of the Saxicola torquatus complex into three groups: rubicola, maurus and stejnegeri. We
proposed the species status for these three groups: European stonechat Saxicola rubicola, eastern stonechat Saxicola maurus and Japanese
stonechat Saxicola stejnegeri. Among them, S. stejnegeri is a cryptic species as it cannot be distinguished by morphometrics and by worn
spring plumage from S. maurus, but differs noticeably by male song.
