Golden Chicken Warbler

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Golden Chicken Warbler
20070304 贵阳 - 黄 腰 柳莺 .jpg

Golden-winged warbler ( Phylloscopus proregulus )

Systematics
Order : Passerines (Passeriformes)
Subordination : Songbirds (passeri)
Superfamily : Sylvioidea
Family : Warbler-like (Phylloscopidae)
Genre : Warbler ( Phylloscopus )
Type : Golden Chicken Warbler
Scientific name
Phylloscopus proregulus
( Pallas , 1811)
Distribution of the golden-winged warbler ( Phylloscopus proregulus ). Yellow: breeding area. Blue: wintering area.

The Firecrest-warbler ( Phylloscopus proregulus , Syn. : Abrornis proregulus ) is a Central and East Asian Singvogelart from the family of warbler-like . Its breeding area extends through the South Siberian coniferous forest taiga and local mountain forests to northern Mongolia and northeast China. The birds are very long-distance migrants that overwinter in southern China and neighboring countries. In recent decades, in Europe, too, growing numbers of golden-winged warblers have been observed as wanderers , especially during the autumn migration .

The name of the species is a certain physical resemblance with the Goldcrest due, especially since the Firecrest-warbler one of the smallest Eurasian birds from "Warblers type" (d. E. The subfamily Sylvioidea ) is.

The birds have a relatively large head and short tail for their size. The upper side of the plumage is generally olive-green, the underside rather light. Noticeable are the light over-eye stripes and the dark rein-eye stripes, as well as the two pale yellowish wing stripes. However, in these characteristics it is similar to the yellow-browed warbler found in the same distribution area . Characteristic are therefore the yellow apex and a lemon-yellow rump band that it only shares with some closely related Southeast Asian warbler species, which were previously considered a subspecies of the golden-winged warbler. A sure distinguishing feature of these species, however, is the very unusual song.

Golden-winged warbler are insectivores that catch their prey in trees and bushes in short, whirring hunting flights. Since the species has a very large distribution area and no negative population trends are known, the IUCN does not classify it as endangered ( least concern ).

description

Features: crown stripe, over eye stripe with headband and rein eye stripe. Only one of the two wing bars is clearly visible here.

With a length of 9–10 cm and a weight of 4–7 g, the golden-winged warbler is one of the smallest twig warbler, even smaller than the yellow-browed warbler and hardly larger than the winter golden-winged warbler. For such a small bird, the head still looks relatively large and the tail extremely short.

The top of the bird is generally dark olive green. The yellow tips of the rear feathers form a striking striped pattern. The dark medium and large arm covers with their sulfur-yellow tips form two conspicuous wing bands on both sides of the resting bird - one of the most important characteristics of the species. The typical, light, approximately 10 mm wide rump field is often recognizable in the fighter flight in particular. The otherwise dark top of the head is marked by a clear, light yellow parting. The side of the head is marked by a dark rein-eye stripe and a lemon-yellow over-eye stripe that extends to the neck. Above the beak, these over-eye stripes merge into a blurred light headband. The underside is light gray to white and partly light yellow in the chest area, especially towards the armpits.

The bird's beak is rather short. The upper bill is blackish-brown, the lower bill is more yellowish to light brown, especially towards the root, but towards the tip it is dark like the upper bill. The iris is brown. The legs are dark brown with a touch of green or gray.

Very similar: the Yunnan warbler ( Phylloscopus yunnanensis ) with a less distinctive yellow head and wing stripes

In its Asian distribution area, the golden-winged warbler can be distinguished from the very similar species, which were previously regarded as its subspecies, by the somewhat more yellowish markings mentioned above, but above all by its different song. But other Asian warblers are also quite similar: Golden-banded warbler ( Phylloscopus pulcher ) and Brooks warbler ( Phylloscopus subviridis ) are somewhat larger, lighter green on the upper side and less prominently patterned with more yellow-brown or white wing bands. The gray-throated warbler ( Phylloscopus maculipennis ) has a rather grayish head markings, face and neck, and a light yellow underside.

The sexual dimorphism is weak, in the plain dress the birds are a little lighter greenish on the upper side, have slightly lighter edges of the feathers and almost whitish markings. The young birds are very similar to the adult animals, with the upper side having a slight brownish tinge and the lower side being rather light gray. The stripe above the eyes is less clear. The adult birds go through a full moult after the breeding season in August to September before they move to winter quarters. The juveniles moult only partially for the first time and a little earlier (July – August), juveniles and adult birds go through a partial moult before the breeding phase in March or April, during which they replace most of the body feathers and some tail feathers.

The song of the golden-winged warbler is usually performed from a covert singing point near the top of a tall tree. The singing is loud, varied and melodic. It can be described, for example, as tirrit-tirrt-tirritt-tertschie-terschie-tertschie-tschu-tschu-tschi-tschi-tschu-tschu , but it is very variable and often contains trill elements. It is reminiscent of the Kanarengirlitz , overall in the melody but also of the forest pipit , which also has a similar distribution area. The singing is divided into stanzas of 2-4 seconds, but is often performed in long series of up to 10 minutes. It is performed both in the breeding and wintering areas and on the train. The call is a short, two-syllable, hard, but melodic dschu-ie . In contrast, the former subspecies have very different chants, which often sound more metallic, monotonous and harsher. The calls of these species are typically sharp and monosyllabic.

Habitat and Distribution

Typical breeding area: forest area in the Siberian taiga

The golden-winged warbler breeds in the coniferous forests of the taiga, which contain fir , spruce , pine and larch , but also in mixed mountain forests with rhododendrons and a high proportion of coniferous wood. In mountains (. Eg in Stanovoy Range and in Kolyma ) also be partially crippled forests inhabited to the tree line. In southern Russia breeding areas up to an altitude of 1500–1700 m have been established. In winter quarters, the range of habitats is greater and includes deciduous and coniferous forests, evergreen oak and laurel forests, but also bushes and even gardens.

The breeding area expanded significantly to the west in the 19th century. While Irkutsk was still considered to be the Siberian western border at the turn of the last century , the species is now common in the area around Krasnoyarsk and also occurs regularly in Tomsk and areas west of the Ob . The Altai is also populated. The southern limit of distribution extends over northern Mongolia to the Amur region , northeastern China and the Sea of ​​Okhotsk . The island of Sakhalin is also part of the breeding area, possibly also parts of North Korea . To the north, the breeding area in central Siberia extends partially to the 64th parallel north, and in eastern Siberia to the northeast of Magadan .

The winter quarters are mainly in the subtropical southern China south of the Yangtze River including the island of Hainan , northern Vietnam and Laos and also northern Thailand .

Settlement density and status

In large parts of its range, the species is widespread and quite common. Settlement densities of up to 35–50 pairs / km 2 were found in south- east Russia , although densities of 2–6 pairs / km 2 , exceptionally up to 15 pairs / km 2 , were found in west-central and north Siberian areas . The species is also quite common locally in parts of the wintering area in Southeast Asia.

In the absence of any evidence to the contrary, the population size is considered stable, although more accurate data are not available. Based on this assessment and the large distribution area, the species is classified by the IUCN as least concern , as a higher risk level would require a reduction of 30 percent within a decade or three generations.

Migratory behavior

John Gould's drawing from 1837 of the "Damatin Gold Cockerel" ( Original: "Dalmatian Regulus" )

The main migration route of the golden-winged leaf warbler leads through Mongolia, Manchuria , Korea and above all East China. Although the species breeds on Sakhalin and overwinters in the described Southeast Asian area, only a few birds are observed on the migration in Japan. Before the turn of the millennium there was an extreme exception (an observation in 1967), but individuals are now detected there every year.

As the species has spread to the west, golden-winged warblers are now seen as wandering visitors, especially in autumn in Europe. The first evidence in Europe goes back to 1829, when the Austrian officer and ornithologist Christoph Fellner von Feldegg shot a specimen of the species in Dalmatia . John Gould , who examined the specimen and described it formally, did not recognize that it was a species already described in Asia, and called it "Dalmatian golden cockerel " (English "Dalmatian Regulus", Regulus modestus ). The German ornithologist Heinrich Gätke , who had moved to the then British island of Heligoland in 1837 , repeatedly found Asian leaf warblers there, including golden-cockerel leaf warblers in 1845 and 1875. In the 80s of the 19th century some specimens were found near Orenburg in the foothills of the Urals and since then in more and more regular numbers in various places in Eastern, Central and Northern Europe.

In Western Europe, the first record in the British Isles dates back to 1896, where a bird was shot in Norfolk . The next record, however, comes from 1951. Since then, the species has been spotted frequently as a stray visitor and has no longer been a rarity there since 1990. In 2003, as many as 313 specimens were counted in the United Kingdom. The bird now appears annually in the Nordic countries of Sweden, Finland and Denmark. Since the turn of the millennium, individuals have also been found almost every year on the north German coasts, especially on Heligoland, but also on other offshore islands such as Greifswalder Oie .

Most of the European records of the golden-winged warbler are birds in their first year of life. Various reasons for the steady increase in the number of records in Western Europe are discussed. Classically, the birds were seen as random visitors who deviated from normal behavior. The abundance of species from the Central Siberian, especially this species, ultimately led to the theory of the reverse migration , since the young birds migrate practically exactly in the opposite direction to their actual migration route on their way to Western Europe . Recently, the establishment of a new migration route has also been suspected, which takes advantage of the increasingly mild climatic conditions in wintry Western Europe. Criticism of this idea arises from the fact that many of these birds should overwinter in northwestern Spain, but little in Spain as a whole, and when they are observed in the east. It has therefore been suggested that the birds continue their migration directly in a south-easterly direction from the frequent central and western European observation sites.

Outside Europe, the species has also been detected as an errant in North Africa (Tunisia and Morocco), the Middle East (Israel, Turkey and Iran), Central Asia (Uzbekistan and Tajikistan), South and Southeast Asia (Bangladesh and Taiwan), as well as Alaska .

behavior

The golden-winged warbler is not particularly shy, but its inconspicuous way of life in the crown area makes it difficult to watch. The bird is practically constantly in motion, flapping and shaking like the golden cockerel , but much more often. Sometimes the bird hangs overhead like tit-like.

nutrition

Like other warblers, the golden-winged warbler is insectivorous and feeds on the adults , larvae and pupae of small insects such as flies , moths and aphids, as well as arachnids . The food is picked up from leaves and twigs or captured like a flycatcher in short fighter flights . Outside the breeding season, they also join mixed flocks of small birds, including titmice , golden cockerel and other warblers, to forage. In the winter quarters, the spectrum of species with which foraging is done collectively also includes spectacled birds , prickly-bred pods and timales .

Reproduction

The breeding season takes place from June to July, the eggs are laid from mid-June. The nest is only built by the wilt, usually near the trunk of a conifer at a height of between 2 and 5 m, in some cases up to 13 height. Exceptionally, you can also breed in bushes or crippled wood. The nest is a spherical structure about 8 cm in size made of moss, lichen, needles and stalks with an inner padding of feathers, grass and animal hair. Four to six eggs are laid, which are only incubated by the female. The eggs are whitish with dark colored dots that condense towards the blunt pole. The young hatch after 12–13 days and fledge after a further 12–14 days. During this time they are cared for exclusively by the female, and then by both parents for about a week. In the south of the distribution area a second brood may sometimes also be carried out, but more detailed information on this topic is not yet available. The breeding area is usually 3–5 hectares in size, exceptionally up to 10 hectares.

The Firecrest-warbler is host bird which specializes in warblers and Pieper Hopfkuckucks , like its close relative, the (ordinary) Cuckoo , a brood parasite is. The eggs of this cuckoo are similarly colored, but slightly larger than those of the host bird.

Systematics

The first to describe the species is the German zoologist Peter Simon Pallas , who found the species on the Ingoda River in Siberia in May 1772. At that time, Pallas led a scientific expedition to Siberia on behalf of Tsarina Catherine the Great , which took place between 1768 and 1774. When published in 1811 he named the species as Motacilla proregulus . The English name of the golden-winged leaf warbler is " Pallas's leaf warbler " in honor of the person who first described it .

The current scientific genus name Phylloscopus is derived from the ancient Greek φύλλον (phýllon) , "leaf", and skopos , "seeker" (from σκοπέω , "peeking, looking around, looking at"). The specific epithet proregulus comes from the Greek pro , "near", and Latin regulus , "little king", which refers to the golden cockerel , Regulus regulus .

The genus Phylloscopus , first described in 1826 by the German zoologist Friedrich Boie , currently contains around 78 species of small songbirds of the twig-singing type and bears the German name Laubsänger . Formerly part of the then larger family of Sylviidae , the genus now forms its own family, the Phylloscopidae .

Within the genus, the golden-winged warbler is part of a group of similar, very small, Asian warbler species with similar characteristics, such as yellowish-white underside, clear stripes above the eyes, crown stripes and wing bands, which are also genetically related and form a clade within the genus . In the past, the birds of this group were listed as a separate genus Abrornis (from the Greek αβρος, abros: pretty and ορνις, ornis: bird), which, however, still contained some not closely related species. Today this genus is again used by some authors in the sense of the clade comprising about ten species.

Four of the species in the Abrornis clade were previously listed as subspecies of the golden- winged warbler . The species as it is today was used as the nominate form P. p. proregulus led. The other four subspecies have breeding areas that are further south and often at higher altitudes, for example in the western Himalayas , in Yunnan and north to Gansu and Hebei (only here with contact with the breeding area of ​​the nominate form).

Although field ornithologists such as Gilbert White and William Edwin Brooks underlined the importance of vocalizations to distinguish the very similar-looking leaf warblers as early as the 19th century , this view was viewed with skepticism by experts for a long time. In recent times, vocalization has become increasingly important for systematics. In the case of the golden-winged warbler, this has led to the southern subspecies being separated off as separate species because of their distinctly different calls and chants, although the plumage characteristics differ only slightly. This “split” was confirmed by molecular genetic studies by Per Alström from the year, so that the golden-winged warbler is nowadays (2018) generally regarded as a monotypical species. The subspecies separated as new species are:

  • Yellow-rump warbler ( Phylloscopus chloronotus ): Along the Himalayas from Pakistan to Nepal and Assam , bordering there in southwestern China. Two subspecies: P. c. chloronotus and P. c. simlaensis .
  • Sichuan warbler ( Phylloscopus forresti ): South China ( Tibet , Shaanxi , Shanxi , south to Yunnan and western Sichuan ), possibly western and northern Myanmar
  • Cansar warbler ( Phylloscopus kansuensis ): Central China ( Qinghai and Gansu), monotypical.
  • Yunnan warbler ( Phylloscopus yunnanensis ): Central to Northeast China (Sichuan, Qinghai, South Gansu, Shaanxi and Hebei), monotypical.

The total of five species thus form a species complex or a super species . The golden-winged warbler seems to have genetically separated from the Yunnan warbler 4.1–5.5 million years ago, from the other species around 1.7–3.2 million years ago.

literature

  • Kevin Baker: Warblers of Europe, Asia and North Africa (Helm Identification Guides) . Helm, London 1997, ISBN 978-0-7136-3971-1 .
  • Urs N. Glutz von Blotzheim & Kurt M. Bauer: Handbook of the birds of Central Europe . Passeriformes (3rd part): Sylviidae, licensed eBook edition. Ed .: Urs N. Glutz von Blotzheim. tape 12 . Aula-Verlag, 2001, ISBN 3-923527-00-4 .
  • P. Alström, P. Clement & GM Kirwan: Pallas's Leaf-warbler ( Phylloscopus proregulus ) . In: J. del Hoyo, A. Elliott, J. Sargatal, DA Christie & E. de Juana, E. (Eds.): Handbook of the Birds of the World Alive . Lynx Edicions, Barcelona 2018 ( hbw.com [accessed September 7, 2018]).
  • Andy M Stoddart: Siberia's Sprite: A history of fascination and desire . Self-published, 2016, ISBN 978-1-5327-6903-0 .
  • Lars Svensson , Killian Mullarney & Dan Zetterström: The Cosmos Bird Guide . Kosmos, Stuttgart 2011, ISBN 978-3-440-12384-3 .

Web links

Commons : Golden Grouse Warbler ( Phylloscopus proregulus )  - Collection of images, videos and audio files

Individual evidence

  1. ↑ a b c d e f g h i j k l Glutz von Blotzheim, p. 1086 ff.
  2. ↑ a b c Svensson (2011), pp. 334-335
  3. ↑ a b c d e f g h i Alström et al. in HBW alive, Phylloscopus proregulus
  4. ↑ a b H. Shirihai and L. Svensson: Handbook of Western Palearctic Birds Volume I Passerines: Larks to Warblers . Helm, London 2018, ISBN 978-1-4729-3757-5 , pp. 418-422 .
  5. ↑ a b c d e Baker (1997) pp. 283-285.
  6. ^ A b Mark Brazil: Birds of East Asia . A & C Black, London 2009, ISBN 0-7136-7040-1 , pp. 356 .
  7. ^ A b c Eric Simms: British Warblers (New Naturalist Series) . William Collins, Sons, London 1985, ISBN 0-00-219810-X , pp. 338-340 .
  8. ↑ Lemon-rumped Leaf-warbler ( Phylloscopus chloronotus ) . In: J. del Hoyo, A. Elliott, J. Sargatal, DA Christie & E. de Juana, E. (Eds.): Handbook of the Birds of the World Alive . Lynx Edicions, Barcelona ( hbw.com [accessed September 7, 2018]).
  9. ↑ Chinese Leaf-warbler ( Phylloscopus yunnanensis ) . In: J. del Hoyo, A. Elliott, J. Sargatal, DA Christie & E. de Juana, E. (Eds.): Handbook of the Birds of the World Alive . Lynx Edicions, Barcelona ( hbw.com [accessed September 7, 2018]).
  10. ↑ IUCN 2018.1 Red List of Endangered Species, see above
  11. ^ Alan C. Jenkins: The Naturalists: Pioneers of Natural History . Hamish Hamilton, London 1978, ISBN 0-241-89999-0 , pp. 84-86 .
  12. ^ John Gould: Birds of Europe . tape 2 . Self-published, London 1837, p. 149 ( archive.org ).
  13. ^ Henry Seebohm : On the Phylloscopi or Willow-Warblers . In: Ibis . tape 19 , no. 1 , 1877, p. 66-108 , doi : 10.1111 / j.1474-919X.1877.tb06167.x .
  14. ^ Henry Seebohm: Supplementary notes on the ornithology of Heligoland . In: Ibis . tape 19 , no. 2 , 1877, p. 156-165 , doi : 10.1111 / j.1474-919X.1877.tb06176.x ( biodiversitylibrary.org ).
  15. ^ Thomas Southwell: Occurrence of Phylloscopus proregulus in Norfolk . In: The Zoologist . tape 20 , 1896, pp. 466-467 ( biodiversitylibrary.org ).
  16. ↑ Eric Ennion: Pallas's Warbler at Monks' House, Northumberland . In: British Birds . tape 45 , no. 7 , 1952, pp. 258-260 ( britishbirds.co.uk [PDF]).
  17. ^ Michael J. Rogers & the Rarities Committee: Report on rare birds in Great Britain in 1991 . In: British Birds . tape 85 , no. 10 , 1992, pp. 507-555 ( britishbirds.co.uk [PDF]).
  18. ↑ Peter Fraser & Michael J. Rogers: Report on scarce migrant birds in Britain in 2003 Part 2: Short-toed Lark to Little Bunting . In: British Birds . tape 99 , no. 3 , 2006, p. 129-147 ( britishbirds.co.uk [PDF]).
  19. ^ A b David Snow & Christopher M. Perrins (eds.): The Birds of the Western Palearctic concise edition . tape 2 . Oxford University Press, Oxford 1998, ISBN 0-19-854099-X , pp. 1324-1325 .
  20. ↑ Kasper Thorup: Reverse migration as a cause of vagrancy . In: Bird Study . tape 51 , no. 3 , 2004, p. 228-238 , doi : 10.1080 / 00063650409461358 .
  21. ↑ James Gilroy & Alex Lees: Vagrancy theories: are autumn vagrants really reverse migrants? In: British Birds . tape 96 , no. 9 , 2003, p. 427-438 ( britishbirds.co.uk [PDF]).
  22. ↑ Eduardo de Juana: Where do Pallas's and Yellow-browed warblers ( Phylloscopus proregulus , Ph. Inornatus ) go after visiting northwest Europe in autumn? An iberian perspective . In: Ardeola . tape 55 , no. 2 , 2008, p. 179-192 ( ucm.es [PDF]).
  23. ↑ Qiang Zhang, Richou Han, Zhongliang Huang & Fasheng Zou: Linking vegetation structure and bird organization: response of mixed-species bird flocks to forest succession in subtropical China . In: Biodiversity and Conservation . tape 22 , no. 9 , 2013, p. 1965–1989 , doi : 10.1007 / s10531-013-0521-5 .
  24. ^ Paul A. Johnsgard: The Avian Brood Parasites: Deception at the Nest . Oxford University Press, Oxford 1997, ISBN 0-19-535499-0 , pp. 196 .
  25. ↑ Phylloscopus proregulus , Pallas's Warbler, Cuculus saturatus, Cuculus optatus , Oriental Cuckoo. Zoological Museum of Moscow University, September 24, 2010, accessed September 20, 2016 (Russian, English).
  26. ↑ James R. Masterson & Helen Brower: Bering's Successors, 1745-1780. Contributions of Peter Simon Pallas to the History of Russian Exploration toward Alaska . In: Pacific Northwest Quarterly . tape 38 , no. 1 , 1947, p. 35-83 , JSTOR : 40486763 .
  27. ↑ Peter Simon Pallas: Zoographia Rosso-Asiatica . tape 1 . Russian Academy of Sciences , Saint Petersburg 1811, p. 499 (Latin, biodiversitylibrary.org ).
  28. ↑ James A. Jobling: The Helm Dictionary of Scientific Bird Names . Christopher Helm, London 2010, ISBN 978-1-4081-2501-4 , pp. 305, 318 .
  29. ↑ Per Alström, Per GP Ericson, Urban Olsson & Per Sundberg: Phylogeny and classification of the avian superfamily Sylvioidea . In: Molecular Phylogenetics and Evolution . tape 38 , no. 2 , 2006, p. 381-397 , doi : 10.1016 / j.ympev.2005.05.015 , PMID 16054402 .
  30. ↑ Baker (1997) pp. 252-253.
  31. ↑ Per Alström, FE Rheindt, R. Zhang, M. Zhao, J. Wang, X. Zhu, CY Gwee, Y. Hao, J. Ohlson, C. Jia, DM Prawiradilaga, PGP Ericson, F. Lei & U. Olsson: Complete species-level phylogeny of the leaf warbler (Aves: Phylloscopidae) radiation . In: Molecular Phylogenetics and Evolution . tape 126 , 2018, p. 141–152 , doi : 10.1016 / j.ympev.2018.03.031 .
  32. ^ E. Dickinson & L. Christides, The Howard and Moore Complete Checklist of the Birds of the World . Passerines. tape 4 , 2014.
  33. ↑ a b c Leaf-warblers (Phylloscopidae) . In: J. del Hoyo, A. Elliott, J. Sargatal, DA Christie & E. de Juana, E. (Eds.): Handbook of the Birds of the World Alive . Lynx Edicions, Barcelona ( hbw.com [accessed September 9, 2018]).
  34. ^ John Edward Gray: Catalog of the specimens and drawings of mammalia and birds of Nepal and Thibet. Presented by BH Hodgson to the British Museum. British Museum, London 1846, p. 66 ( biodiversitylibrary.org ).
  35. ↑ Baker (1997) p. 282.
  36. ↑ Kenneth Williamson: Identification for Ringers 2 . Field guide. tape 8 . British Trust for Ornithology, Tring, Herts 1976, pp. 8, 13-15 ( bto.org [PDF]).
  37. ^ Gilbert White: The natural history of Selborne: with observations on various parts of nature, and the naturalist's calendar . J. Chidley, London 1840, p. 38 .
  38. ^ William Edwin Brooks: A few observations on some species of Phylloscopus . In: Ibis . tape 6 , no. 22 , 1894, pp. 261-268 , doi : 10.1111 / j.1474-919X.1894.tb07751.x ( biodiversitylibrary.org ).
  39. ↑ Horace Gundry Alexander: Field notes on some Asian leaf-warblers . In: British Birds . tape 48 , no. 11 , 1955, pp. 293-299 ( britishbirds.co.uk [PDF]).
  40. ↑ Per Alström: Species concepts and their application: insights from the genera Seicercus and Phylloscopus . In: Acta Zoologica Sinica . 52 (Supplement), 2006, p. 429-434 ( actazool.org [PDF]).
  41. ↑ a b Jochen Martens, Dieter Thomas Tietze, Siegfried Eck & Michael Veith: Radiation and species limits in the Asian Pallas's Warbler complex ( Phylloscopus proregulus see left) . In: Journal of Ornithology . tape 145 , no. 3 , 2004, p. 206-222 , doi : 10.1007 / s10336-004-0042-9 .