Parus

Eck, Siegfried, 2006, The Palaearctic Titmouse Species (Aves: Paridae: Parus sensu lato) — A current survey *, Zootaxa 1325, pp. 7-54 : 34-39

5. Parus [a.] montanus

P. montanus , montanus group

This group is here distinguished by the regiolect “alpine” (whistled song). It inhabits the Alps, as far as Slovenia (schiebeli), lives in the Danube lowlands around Vienna and in Hungary (Thönen 1996: 6), and its range extends to both the Bohemian Forest (Eck 1987) and the Altvater Mts. in Mähren (where modulated “lowland” song is also heard), all of Slovakia and at least S Poland as well as the entire Carpathian arc (Western, Wooded, Eastern and Southern Carpathians). The complexity of the differences between some populations in the Alps unfortunately caused all of them to be combined under montanus CONRAD (1827). Kleinschmidt wanted to reject this name and replace it by baldensteini, but was unsuccessful; cf. under P 34.

The list of alpine Willow Tits that follows must surely be further extended by western alpine forms. — The “forms” — if they exist — on Corsica (Kleinschmidt 1937: 37; Thibault 1983; Quaisser & Eck 2003: 4) remain entirely unclear, as also in the Abruzzi Mts., where Eck, Feiler and Quaisser recently searched for alpine tits without success.

P. montanus montanus CONRAD, 1827 [Graubünden; syn. baldensteini SALIS, 1861, St. Moritz; styriacus KLEINSCHMIDT, 1937, St. Georges above Judenburg, S Austria]; P. montanus supermontanus KLEINSCHMIDT, 1921 [Trent]; P. montanus schiebeli KLEINSCHMIDT, 1937 [Zirknitz/Krain, Slovenia];? P. montanus transsylvanicus KLEINSCHMIDT, 1921 [Türkös near Kronstadt/ Brašov]; P. montanus assimilis C.L. BREHM, 1855 [Carpathians in E Galicia, see above].

Parus montanus , salicarius group

This group is here distinguished by the regiolect “lowland” (modulated song), so that the Balkan rhodopeus and in SE France alpestris are included in it. The name alpestris, considered synonymous with montanus by authors from Hartert to Vaurie, belongs to birds that are distinctly greyer than salicarius but sing similarly! Thönen (1996: 12–15) discusses this situation in detail; see also his map in Fig. 3 p. 5. To me it seems unlikely that they belong to the ssp. salicarius. Grouping according to vocalizations is preferred here, because no reasonable discrimination criteria based on either colour or structure are available.

P. montanus kleinschmidti HELLMAYR, 1900 [Coalfall Wood at Finchley, near London]; P. montanus rhenanus KLEINSCHMIDT, 1900 [Rhine at Darmstadt; syn. subrhenanus KLEINSCHMIDT & V.JORDANS, 1916, Bonn]; P. montanus alpestris BAILLY, 1852 [Chambéry, cf. Jouard 1936; Thönen 1996: 5]; P. montanus ssp. [Harz, see Eck 1980a: 28–29; 1994: 113); P. montanus salicarius C.L. BREHM, 1831 [Renthendorf, Thuringia according to Kleinschmidt 1897: 79; syn. natorpi KLEINSCHMIDT, 1917, Emanuelssegen / Upper Silesia]; P. montanus rhodopeus HARRISON & PATEFF, 1937 [Beglik / Rhodope Mts.].

Parus montanus , borealis group

The borealis group is distinguished substantially by the “Siberian song” (Martens & Nazarenko 1993), documentation also in Cramp and Perrins (1993, VII: 179–180) and Thönen (1996: 8–9).

“ Poecile baicalensis suschkini ”, according to the measurements given by Hachlow (who described his measurement technique in his paper on Redpolls in “Uragus” IV, No. 3: 2), definitely has the same relative dimensions as borealis, which is also confirmed by two skins from the Saur Mts. The author emphasizes the gigantic throat patches of his new form (June and July birds, but also one from March!), and the underside is said to have a “tinge of cream colour”! The bird from March is said to show an “ochre­brownish tone” on the shoulder feathers. In fact, this raises the question of a hybridisation with songarus!

I examined about a dozen skins from various regions of the Altai, the wing and tail dimensions of which are scattered broadly across the clearly differentiated borealis and baicalensis dimensions (mainly borealis). The cap is also not as deep a black as in baicalensis . Ernst (1991) described several song forms occurring together in the same region (a mixing region for songs?). Is the only song known from Mongolia the “lowland” type? When a regiolect is used to characterize the group, as has so far been done for the Willow Tits, it becomes complicated in the Altai (Ernst 1991; Martens et al. 1995; Ernst & Hering 2000).

The Ussurian shulpini is dorsally a nuance more yellow than baicalensis , also smaller and relatively short­tailed like borealis (I examined the type specimen and additional skins in St. Petersburg); Martens’ far­eastern song samples refer to P. m. shulpini . — The broad interpretation of a ssp. borealis by Stepanjan (1990: 565) is incomprehensible.

P. montanus colletti STEJNEGER, 1888 [Bergen, Norway]; P. montanus borealis SELYS­ LONGCHAMPS, 1843 [corrected: Norway,? syn. uralensis GROTE, 1927, Ufa;]; P. montanus tischleri KLEINSCHMIDT, 1917 [Losgehnen at Bartenstein, E Prussia = Lusiny pow Bartoszyce, Poland, see Kaemmerer 2004]; P. montanus loennbergi ZEDLITZ, 1925 [Kuhmoniemi, Finland]; P. montanus baicalensis (SWINHOE, 1871) [Kultuk]; P. montanus shulpini (PORTENKO, 1954) [Sutschan, S Ussuria]. —

? P. montanus suschkini (HACHLOW, 1912), mistakenly written as “HACHLOR” [river Temir­Su in the Saur Mts.], see above.

Parus montanus , kamtschatkensis group

The forms in the far eastern Palaearctic, from the Koryak highlands through Kamchatka, Sakhalin and Japan, are coherent by way of a character gradation (Eck 1982: 139–140). A mixture between anadyrensis and baicalensis was also merely supposed by Stepanjan (1990: 566; see Eck l.c.: 139, footnote). — What is now especially problematic is the distribution of the songs: Portenko (1939, I: 95) represented that of anadyrensis as “tju­tju­tju”, which is hard to translate. In any case, each whistle seems to start with a “t” (“lowland”?). — The adjacent form to the south, kamtschatkensis, sings “alpine”, sachalinensis has the “Siberian song”, and on Honshu, for restrictus, the “Sino­Japanese frequency­jump song” is known. — The Willow Tits of Shikoku (abei) have been described according to features of coloration, but not recognized as ssp. I have not seen birds from Shikoku and Hokkaido.

The taxonomic position of the Willow Tits on Hokkaido (Jesso) is unclear, since their “alpine” song type became known (Thönen 1996: 18). Hartert in 1905 declared that the Willow Tits there did not belong to restrictus, but in 1934 he asserted without further commentary that restrictus lives on Hokkaido. According to the “Hand­list of Japanese birds” (1922: 157) Hokkaido is inhabited by sachalinensis, and according to Vaurie (1959) and the “Check­list of Japanese birds” (1975: 283, see also 2000: 237) also by restrictus. Abe & Kurosawa (1984) published measurements of 123 Hokkaido Willow Tits, in which the wing dimensions were taken as “unflattened natural chord” (maximally 65 mm), but the tail length varies widely, between 51 and 62 mm “from the base of central tail­feathers ...”. In view of the different song form, these birds cannot belong to sachalinensis.

Thönen (1996: 18 ff.) reconstructs the phylogenesis of the far eastern island populations on the basis of their song, and links the Hokkaido Willow Tits to the “Euro­ Siberian populations”, the Hondo Willow Tits to weigoldicus . This is an impressive demonstration that biosystematics cannot be based exclusively on bioacoustic or morphological­geographic data, nor (see below under weigoldicus ) exclusively on molecular genetics.

P. montanus anadyrensis BELOPOLSKI, 1932 [Markowo, Anadyr region]; P. montanus kamtschatkensis (BONAPARTE, 1850) [Kamchatka]; P. montanus sachalinensis Lö NNBERG, 1908 [Sakhalin]; P. montanus ssp. [Hokkaido]; P. montanus restrictus HELLMAYR, 1900 [Shimotsuke, Hondo; syn. abei MISHIMA, 1961, Takagoshi­yama, Tokushima/Shikoku].

Parus montanus , songarus group

When songarus is compared only with baicalensis (or borealis), the external difference is very obvious, so it is understandable that (predominantly Russian) authors have considered it a distinct species. This view is shared by Roselaar (In: Cramp & Perrins 1993, VII: 185) and Harrap and Quinn (1996), but contradicted by Thönen (1996: 17–18). The song form of the ssp. songarus is a special “Siberian song” (Martens et al. 1995: 375–377). P. m. affinis evidently has two spatially distinct variants of the “alpine” song, and only the “alpine” song is known for P. m. stoetzneri (but see Martens et al. 1995: 378).

P. montanus songarus SEWERTZOW, 1872 [Tian Shan]; P. montanus affinis (PRZEWALSKI, 1876) [Ala Shan and Gansu]; P. montanus stoetzneri KLEINSCHMIDT, 1921 [30 km N of Balihandia, NE of Jehol, NE Tschili; NE China].

The population groups: P 34 montanus , supermontanus, schiebeli, transsylvanicus, assimilis; Alps to Bohemian Forest, Tatra and Balkans (alpine tits). — Figs. Eck 1980b: Plates IIIa (supermontanus), VII (schiebeli).

* Subtle alterations in back coloration, wing length and relative tail length. At present

the following subdivision can be proposed:

The southern Alps, from Tessin to Slovenia, are inhabited by the supermontanus

described from Trent, a nuance greyer than more northern alpine tits and larger, with wing length up to 72 mm; found in Tessin by G. v. Burg and in “Unterkrain (Warmberg bei Ainödt)” by O. Kleinschmidt. Further north is the somewhat browner and smaller (wing length up to 69 mm) montanus , which here must be considered to include the birds of the Bohemian Forest with “alpine” song. The region east of supermontanus in Slovenia is inhabited by schiebeli (warm brown, wing length up to 70 mm, cf. Kleinschmidt 1937: 36). — Carpathians: in the Southern Carpathians we find transsylvanicus (very light brown coloration, wings up to 69 mm). The birds of the Eastern to Western Carpathians, first known as alpine tits in 1977 (Eck & Geidel 1977; Königstedt 1983), reach wing lengths of 68 mm and tend to have longer tails than montanus s.str. In a scatter diagram combining wing and tail lengths, the same position is occupied by montanus (syn. styriacus), with the alpine tits from the Czech Bohemian Forest (on the opposite side, around Frauenau in Bavaria, the Willow Tits have “lowland” song according to recordings made by Eck in March 2001), and the larger supermontanus. Birds from E Slovakia and the High Tatra (= Western Carpathians) are clearly disparate (tending to have relatively long tails) and should be distinguished as P. m. assimilis . The dimensions of the type specimen of assimilis (Mus. New York 682238: wing 65, tail 60 mm) are consistent with those of Slovakian alpine tits and with borealis. However, the assimilis type specimen, collected in April, according to Kleinschmidt (1897: 88), had a “weak brownish­grey shade” on the back that was reminiscent of montanus , and are thought to come from the Carpathians in eastern Galicia, where the song is “alpine”. If it turns out that the separation of the Slovakian and Eastern Carpathian alpine tits from transsylvanicus (Southern Carpathians) is justified, the designation assimilis would have to be used (cf. Stresemann & Sachtleben 1920: 257, footnote 2). Otherwise (as I suspect) transsylvanicus would be a synonym of assimilis. However, this would initiate a complicated nomenclatural­technical procedure, because the name assimilis has been interpreted in various ways! — “Alpine”­singing Willow Tits.

P 35 kleinschmidti, rhenanus, salicarius, alpestris, rhodopeus; W and central Europe. —

Figs. Quinn: Plate 16: 50a (kleinschmidti); Kleinschmidt 1897: Plate III below (rhenanus). * Is darkest and smallest (wing length 56–62 mm) on the British Isles (kleinschmidti); on the continent wing length reaches 63 mm (rhenanus) at the Rhine and 66 mm (salicarius) in the Ore Mountains of Saxony. The isolated Balkan birds (rhodopeus) are very dark; wing length according to v. Jordans (1940: 94), Harrison (1933), Harrison & Pateff (1937: 605) and Eck up to 68 mm. Relative tail length in England x = 84% (n=21), Rhineland as far as Lusatia x = 88% (n=123). — Willow Tits with “lowland” song.

P 36 colletti, borealis (with uralensis?), tischleri, loennbergi, baicalensis , shulpini;

? suschkini; N Palaearctic. — Figs. Kleinschmidt 1897: Plate IV lower Fig. (tischleri); Eck

1980 b: Plate IIIa (baicalensis).

* Regarding colour nuances on the upper side (colletti and tischleri browner, loennbergi greyer than borealis), wing length and relative tail length (Eck 1982): clinal variation. Cf. also the remarks under “ borealis ” group; relative tail length of 70 baicalensis x = 94.1% ± 2.08, of 10 (browner!) shulpini x = 92% ± 1.97. — “Siberian” song.

P 37 anadyrensis, kamtschatkensis, sachalinensis, ssp. on Hokkaido, restrictus; Anadyr

region to Japan. — Figs. Eck 1980b: IIIb (anadyrensis, restrictus); Quinn: Plate 16: 50f. * From north to south becomes greyer, smaller and relatively shorter­tailed. Problems are treated under the P. montanus , kamtschatkensis group. — anadyrensis wing length: 31 ♂♂ 64–70 mm, x = 67.4 mm ± 1.52; 17 ΨΨ 63–69.5 mm, x = 66.3 mm ± 1.75. Tail length: 27 ♂♂ 58–65 mm, x = 62.2 mm ± 1.95; 17 ΨΨ 58.5–66 mm, x = 62.0 mm ± 2.10. The absence of sexual dimorphism is also evident from Portenko’s (1939, I: 97) table of the tail measurements. Relative tail length (n=47) x = 92.8% ± 1.54; kamtschatkensis wing length: 17 ♂♂ 62–68 mm, x = 65.1 mm ± 1.67; 7 ΨΨ 62–65 mm, x = 63.8 mm ± 1.22. Tail length: 14 ♂♂ 56.5–64 mm, x = 59.8 mm ± 2.05; 8 ΨΨ 56–60 mm, x = 58.4 mm ± 1.6. TWI (n=30): x = 91.4% ± 1.60, of sachalinensis (n=14): x = 89.0% ± 1.42.

P. montanus restrictus from Honshu: Wing length of 17 ♂♂ 61–68 mm, x = 64.7 mm ± 1.92; of 10 ΨΨ 61–66 mm, x = 63.7 mm ± 1.72. Tail length of 17 ♂♂ 51–57.5 mm, x = 53.8 mm ± 1.81; of 10 ΨΨ 50.5–55 mm, x = 53.3 mm ± 1.75. TWI (n=29): x = 83.4% ± 1.85.

P 38 songarus; Tian Shan. – Figs. Pleske (1889–1905): Plate VIII: 1; Eck 1980b: Plate VI:

2nd from bottom; Quinn: 16: 51a.

* Conspicuous yellowish brown, wing length of ♂♂ (n=43) 67–73 mm, x = 69.7 mm ± 1.36, of ΨΨ (n=32) 66–71.5 mm, x = 68.2 mm ± 1.35; tail length of ♂♂ (n=42) 57–64 mm, x = 59.9 mm ± 1.57, of ΨΨ (n=31) 56–61.5 mm, x = 59.1 mm ± 1.23. Cf. also under P 40.

P 39 affinis, stoetzneri; N China. — Figs. Pleske (1889–1905), Plate VIII: 2 (affinis); Eck

1980 b: Plate VI: the two upper Figs.; Quinn: Plate 16: 51 b (stoetzneri).

* In comparison with P 38 yellowish brown on the back, in the east greyer birds with a head coloration that Hartert calls “seal brown” in the case of affinis according to Ridgway (1886, cf. Ridgway 1912: Plate XXXIX), and that is approximately the same in stoetzneri (according to the “Michel­Farbenführer”, 37th Ed., 2000, Plate 17 “blackish orange­brown”, Code 11­13­8). The two forms differ in size; stoetzneri is smaller but relatively long­tailed. Furthermore, stoetzneri in its area has a slightly different shade. The dimensions of affinis: wing length of ♂♂ (n=13) 65–68 mm, x = 66.4 mm ± 1.00; of ΨΨ (n=16) 62–66 mm, x = 64.3 mm ± 1.36. Tail length of ♂♂ (n= 13) 59.5–62.5 mm, x = 60.6 mm ± 0.94; of ΨΨ (n=16) 56.5–61.5 mm, x = 58.9 mm ± 1.59. Relative tail length (n=29) x = 91.5% ± 1.58.

stoetzneri: Wing length of ♂♂ (n=10) 62–66 mm, x = 63.7 mm ± 1.21; of ΨΨ (n= 6) 61–63 mm, x = 61.5 mm ± 1.22. Relative tail length (n=18) x = 93.8% ± 2.04.