With these “taxonomical notes” I want to launch a series of notes and thoughts about some problem related to the taxonomy of Western Palearctic birds, addressing some conundrums which have not yet been studied in detail or did not get enough attention. Many of these notes are simply reported in order to stimulate further in depth studies by taxonomist and researchers, with some being instead only a brief view of my researches in due course and in progress. Indeed, for what concern this “first case” here reported, my study started back in Autumn 2003 and the whole problem will be presented on a specific paper in preparation.
Taxonomical notes : Lesser Kestrel is really monotypic?
By Andrea Corso
During autumn 2003, I was at Chockpak Ringing Station, Kazakhstan, alongside with three Dutch birders and friend of mine (Arend Wassink, Justin J Jansen and Wim Nap), studying raptors and the other birds there, with the guiding of Professor Andrei Gavrilov. We visited as well many other sites of Eastern Kazakhstan. The trip was very interesting and fruitful for many aspects and the country was confirmed to be one of my favorite destination. Among the several remarkable observations, I was greatly intrigued by the pattern of the upperwing in most of the adult male Lesser Kestrel ringed and observed in the field. During the firsts observation days in Kazakhstan I was telling my fellow observers some tips about field characters of Lesser Kestrel. While talking about the upperwing grey panel in adult male I emphasized how this character is very often not visible under field condition and usually only at very close view, good light or in perched birds. Looking at the males flying around then, I was readily contradicted, as in all the males around, the grey on the upperwing was not only well visible, even at distance, but also very conspicuous and dark looking. This fact intrigued me very much so I started to check and study in details every male observed in the following days and eventually also all the birds caught to be ringed. It resulted therefore as almost 100% of the males observed shown more grey then I was used to see in European birds; this was confirmed by the many males caught during our stay at the ringing station. For what concern European birds, Corso (2000, 2001) report that to a certain degree, the upperwing grey panel could show a range of variability, among several other characters. However, not any of the European bird personally observed (some 20.000 adult males in total) ever shown an upperwing pattern that may look identical to the birds I observed in Kazakhstan. Rarely, birds within European population (chiefly from the eastern part of the range) may show a more extensive grey wing-panel, with all the GC (greater coverts) sooty-grey and in rare occasion some median coverts (MC) and tertials too (Corso, 2000, Corso, 2001; pers.obs.). However, I never observed any European bird with almost the whole upperwing coverts dark led-grey all the way to scapulars and tertials as in some Kazakhstan birds. None of the main references on European raptors either illustrate or describe birds like these (Cramp & Simmons, 1980; Snow & Perrins, 1998; Forsman, 1999; Clark, 1999; Ferguson-Lees & Christie, 2001). None of them even mention that the grey area (panel) appear to be greater in Eastern populations then in the Western populations. Only in Ferguson-Lees & Christie (2001) it is depicted a bird with more grey then usual but not as much as noticed in this study. In Forsman (1999), the only picture showing a male with much grey on upperwing has been taken in fact in Kazakhstan. I was therefore very excited as I thought to have found a yet un-described subspecies of the believed monotypic Lesser Kestrel. However, once back home, I started to search references on the taxonomy of this falcon and found that back in ‘800 a “variety” or race has been already described by Swinhoe.
Nowadays, Lesser Kestrel Falco naumanni (Fleischer, 1818) is considered monotypic, with no subspecies recognized (Cramp & Simmons, 1980; Snow & Perrins, 1998; Forsman, 1999; Clark, 1999; Ferguson-Lees & Christie, 2001; Corso, 2000, 2001). However, Swinhoe described in 1870 a new race or “variety” (according to his given name) of Lesser Kestrel from Pekin, China which he named Falco cenchris var. pekinensis (Swinhoe, 1870). He described the holotype bird, collected on 18th October 1868, as following : “Large numbers of Kestrels were flying and hovering about. Their movement struck me as peculiar; and on shooting a male we found the species to be a race of Falco cenchris, Naumann. We procured on this occasion an adult male, and in the Western Hills a young male. They agree in size and form with Falco cencrhis of Europe; but the adult male has all the wing-coverts grey right up to the scapulars, most of them narrowly edged with rufous. The adult has the inner or short primaries broadly bordered at their tips with whitish, rufous in the immature, and wanting in the European bird. Both adult and immature have the white on the under quills 3 ¼ inches short of their tips; in the European bird it advances one inch nearer the tips. I will note this Eastern race as var. pekinensis. It will probably be the bird that winters in India.” (Swinhoe, 1870). Later on, Swinhoe (1871) on his “Revised Catalogue of the Birds of China” so reports “Breeds on the western hills of Pekin, and assembles in large numbers in September. Mr. Hodgson’s drawing of the Nepal bird (in the British Museum) shows that it is this race of Tichornis cenchris that resorts to India. The adult male has all the wing-coverts grey right up to the scapulars ; the inner or short primaries are broadly bordered at their tips with whitish, rufous in the immature ; the white on the under quills is 3^inches short of their tips. Subsequently, Jerdon (1871) report this taxon for the Birds of India. It is also mentioned later by Sharpe (1874) in his “Catalogue of the Accipiters, or Diurnal Birds of Prey, in the collection of the British Museum” and so described: “Very similar to C.naumanni, but darker and more vinous red above; underneath also darker-coloured and unspotted when adult. The principal distinction is the wing-coverts, which are almost entirely blue-grey, only the very innermost being slightly washed with rufous. Total length 12 inches, culmen 0-8, wing9-6, tail 5-8, tarsus 1-45. Hab. N: China; Himalayas.” Hodgson (1844, 1845a,b, 1855) mention too the race pekinensis. Hartert (1913) and Hartert & Steinbacher (1933) mention this race as well giving again a description fitting with the original one by Swinhoe and considering valid this taxon (though with some overlapping characters sometimes with the nominate naumanni and reporting some clines). Warren (1966) report : “Syntype, Adult male. Rea. no. 18184.108.40.2062. Near Ming Tombs, north of Peking, 18 Oct. 1868. Collected by R. Swinhoe. Seebohm Collection. Proc. zool. Soc. Lond., 1870 : 442. Also Warren & Harrison (1971, 1973) reported again pekinensis.
More recently, authors has variously faced with the Chinese population from simply ignoring it, to mentioning and describing it as Falco naumanni pekinensis with the given English name of Eastern Lesser Kestrel or synonymizing it with naumanni (lumping together the two taxa): so, for ex. Brown & Amadon (1968) and Brown, et al. (1983) mention among the African raptors also Falco naumanni pekinensis reporting that this race is distributed from Turkestan to northern China, and in Africa it is observed outside the breeding season in Eastern Africa with “Most of the Lesser Kestrels passing through Somaliland appear to belong to this race”. These authors describe pekinensis as so “Adult male differs from the preceding race (naumanni) in having the grey of the wings extending up to the scapulars; secondaries broadly tipped with whitish and a greater amount of blackish on the ends of the primaries. The female and young differ on the latter character only.” Of the same opinion are Etchécopar & Hüe (1967) which report “F.n.pekinensis Swinhoe (Généralment plus gris, sur les moyennes et petites couvertures alaires); Migratrice : Venant d’Asie, accidentelle en Ègypte”. For the Avifauna of China, Etchécopar & Hüe (1978) report again F.n.pekinensis while more recently, Tso-Hsin (1987, 1994) in his synopsis, synonymize pekinensis with naumanni which is reported to be monotypic. Vaurie (1965) mention Falco cenchris var. pekinensis only as a synonym of Falco naumanni, monotypic. Lesser Kestrel in Asia (Pakistan, South Asia, India, China) is considered monotypic also by Ali & Ripley (1978), Ripley (1982), Inskipp, et al. (1996), Grimmett, et al. (1998) while in the recent Rasmussen & Anderton (2005) no mention at all is given about pekinensis.
Was therefore with no clear idea that I went to Tring, the Natural History Museum (NHM) (alongside with my friend and MISC member Ottavio Janni) to check the syntypes of Swinhoe, but eventually what I found were birds clearly different from any Western Lesser Kestrel I knew before. All the specimens labeled as pekinensis preserved into the collections held at Tring and at Almaty Nature Museum of the Zoology Research Institute of the Academy of Science (IZA) show obvious differences to all the other skins of birds collected in the breeding grounds within the Western Palearctic, while some birds collected in the African and even more in the Indian wintering grounds appear indeed similar. I divided the skins into A) birds collected in China during breeding season; B) birds of unknown breeding ground origin, collected elsewhere in Asia; C) birds which are typical naumanni, collected mostly in the breeding grounds of Europe and N Africa; D) birds collected during migration or wintering grounds in Africa, the Middle East and Arabian Peninsula; E) birds collected in India, during migration or winter time.
As shown in the here reported photos, all the birds coming from China (A), most of them labeled originally as pekinensis, show according to the original description of the holotype (called syntype by Warren, 1966 – why?) the whole upperwing coverts grey with also the tertials grey or partially grey. Additionally, the grey tinge is not only reaching the lower scapulars in some birds, but is also of a darker led grey, more sooty (less pure and cold bluish-grey or cerulean-grey), then the birds from Europe (C); on the same way, also the grey on the head and tail is darker and more sooty. The mantle, as correctly reported by Sharpe (1874) is of a deeper and darker vinous rusty-red colour, and the underparts are darker and more satured as well. Birds from other Asian countries (B), such as Kazakhstan, have similarly patterned wing-coverts, though in some birds less extensive and with in most birds part of the lesser and the leading edge rusty tinged (mostly as a rusty “patch” on the inner “harm”), the grey colour of head, tail and coverts being in some not so dark led grey as the Chinese birds (but in some birds being similar) and the underparts appearing less saturated as the mantle less vinous in many birds (but similar in some); indeed, these birds appear intermediate and could be called “cline”. Birds collected in the Middle East, Arabian Peninsula (as in UAE) and Eastern Africa during migration period or winter time have typical plumage (as C) or appear intermediate (as B), but their origin is not known so they could simply be taken into account as “intermediate” birds of unknown origin, while some birds collected in India (E) indeed look like “pekinensis” (as A). On the web and in ornithological books, several birds from India, Arabian Peninsula and Eastern Africa show either an intermediate plumage like birds of the group (B) or a “pekinensis-type” like plumage. The birds from Turkestan, described by Zarudny (1912) as Cerchneis naumanni turkestanicus and reported to have paler plumage than pekinensis with less grey on upperwing but more than on European birds, could be included into the group (B) as intermediate birds of clinal population, while the taxon Cerchneis naumanni sarmaticus (Domaniewski, 1917) is of no taxonomic value at all.
NB: For a comparison study, only birds of the group (A) and (C) could be taken into account, while birds of the group (B) in the middle could only be taken into account as clines or intermediate birds, which are normally found in every subspecies.
From a first and preliminary analysis, it seems that birds of the species’ range show an increasing amount of grey to coverts and saturation in colors moving west to east, with some birds from the Spanish population showing almost no grey on coverts (only a tinges or a tip to the greater coverts), moving east the coverts showing an increasingly wider amount of coverts grey tinged, almost gradually, as well a more saturated plumage. Therefore, it seems likely that there is a cline in both the extension of the grey on the coverts and the saturation of the pigmentation, with however the Chinese birds being constantly different from all the other Lesser Kestrels and with the differences strongest and most visible. This is only a brief overview of the results obtained and a simple and basic summary, for a better and more in depth treatment see Corso, et al. (in prep.). However, from the preliminary result, not only it seems that the pekinensis taxon is valid, as being readily identifiable and obviously different, but that a genetic study of the Chinese population is surely advisable and should be taken into account (could be that result would indicate something like Red-footed Falcon and Amur Falcon, in the past considered conspecific and often called Western and Eastern RFF ?).
In any case, whatever the validity of pekinensis would be confirmed or not, and accepted or rejected, still the differences in the plumage of the eastern populations should finally be acknowledged and taken into account, as up to date these have never been considered and illustrated in any field guide, handbook or reference dealing with Asian birds in last decades and always Lesser Kestrel has been only depicted with the same wing pattern, therefore with a limited amount of grey on the greater coverts, often hard to be seen in flying birds.
I wish to thanks as always the Tring, NHM staff to which I am much indebted for the most important help for any of my birds plumages studies. So a warm thanks goes to Katrina Kook, Robert-Pries Johanes, Mark Adams and the others working at Tring and that helped me in various way. On the same way, a warm thanks goes to Dr. Carla Marangoni, curator of the ornithological section at Museo Civico di Zoologia in Roma (MCZR) for the countless hours of help while I was at the museum among hundreds of skins and specimens. Thanks also to the always kind and helpful Anita Gamauf, curator at the Wien Museum (NMW) and to Prof. Andrei Gavrilov who assisted during the skins study at Almaty Nature Museum of the Zoology Research Institute of the Academy of Science (IZA) and thanks to all the curators of all the other museums where I studied the skins collections in all my visits.
Birds studied in the field:
60.000+ birds (MM and FF): in Italy, Spain, Greece, Tunisia, Morocco, Egypt, Sinai, France, Greece, Israel, Turkey, Georgia, Kazakhstan.
Birds studied in the hand (combined skins and ringed birds):
(A)“Falco naumanni pekinensis”: adult ♂. Rea. no. 18220.127.116.112. Near Ming Tombs, north of Peking, 18 Oct. 1868. Collected by R. Swinhoe. For comparison – 40♂♂- from China. (Tring, NHM; Wien, WNHM; Almaty Nature Museum, IZA); birds in the group (E) were not considered for the description of the characters of pekinensis
(B) Falco naumanni ssp. (showing intermediate characters): 75♂♂ ; 27♀♀ – Kazakhstan, Azerbaijan, Turkmenistan.
(C) Falco naumanni (typical plumage): 270♂♂ ; 82♀♀ – coming from Spain, Italy, Greece, Tunisia, Morocco, Albania, France;
(D) Falco naumanni (both typical naumanni and intermediate birds): 79♂♂ ; 90♀♀ – Turkey, Israel, Egypt, Senegal, Niger, Mauritania, Iraq, Iran, Tanzania, Syria, Lebanon, Jordan, Palestine, Kirgizstan, Slovenia, South Africa, Check Republic, Georgia.
(E) Falco naumanni “pekinensis-type”: 10♂♂ from India.
Skins studied preserved at the following museums: Natural History Museum, Tring, England (NHM); Institute of Zoology, Almaty, Kazakhstan (IZA); Museo Civico di Storia Naturale di Milano, Milan, Italy (MCSM); Museo Civico di Terrasini, Palermo, Italy (MCT); Museo Civico dell’Università di Scienze Naturali di Catania, Catania, Italy (MCUCT); Museo Civico di Zoologia di Roma, Rome, Italy (MCZR); Museo Regionale di Scienze Naturali di Torino, Torino, Italy (MRSN); Museo di Storia Naturale “Giacomo Doria”, Genova, Italy (MSNGD); Museo di Storia Naturale “La Specola”, Firenze, Italy (MSNLS); Museo Civico di Storia Naturale di Carmagnola (); Museo Civico di Scienze Naturali “Angelo Priolo”, Randazzo, Italy (MCR); Naturhistorisches Museum Wien, Vienna, Austria (NMW); Nationaal Natuurhistorisch Museum Naturalis, Leiden, the Netherlands (NNM).
Ali, S. & S.D. Ripley, 1978. Handbook of the Birds of India and Pakistan. 9: i-xvi, 1-306.— Bombay.
Brown, L. H., and Amadon, D., 1968. Eagles, Hawks and Falcons of the World, vol.1 and 2. Country Life Books, London.
Brown, L. H., Urban, E. K.& Newman, K. 1982. The Birds of Africa. Vol. I. London, UK: Academic Press.
Cheng, Tso-hsin, 1987. A synopsis of the Avifauna of China. i-xvi, 1-1223.— Beijing.
Cheng, Tso-hsin, 1994. A complete checklist of species and subspecies of the Chinese birds. i-xx, 1-
Clark W.S., 1999. A Field Guide to Raptors of Europe, The Middle East and North Africa. Oxford University Press, Oxford.
Corso A., 2000. Less is More: British vagrants, Lesser Kestrel. Birdwatch 91: 29-33.
Corso A., 2001. Notes on the moult and plumages of Lesser Kestrel. British Birds 94: 409-418.
Cramp S & Simmons K.E.L. eds., 1980. The Birds of the Western Palearctic. Vol. 2: Hawks to Bustards. Oxford University Press, Oxford.
Domaniewski J., 1917: Przyczynek do znajomości form geograficznych Cerchneis naumanni (Fleisch.) [A
contribution to the knowledge of geographic forms of Cerchneis naumanni (Fleisch.)]. – Comptes Rendus
de la Société des Sciences de Varsovie 10 (9):1043-1047. [In Polish.]
Etchécopar, R. D. & Hüe, F., 1978. Les oiseaux de Chine, de Mongolie et de Corée – Vol. I, Non passereaux. Papeete, Tahiti, Éditions du Pacifique.
Etchécopar, R. D. & Hüe, F. 1964. Les Oiseaux du Nord de l’Afrique, de la Mer Rouge aux Canaries. Paris, France: Editions N. Boubée & Cie.
Ferguson-Lees J. and Christie D.A., 2001. Raptors of the World. Christopher Helm, London.
Forsman D., 1999. The Raptors of Europe and the Middle East. A Handbook of Field Identification. L.T & A.D. Poyser, London.
Grimmett, R., C. Inskipp & T. Inskipp, 1998. Birds of the Indian Subcontinent. 1-888.— London.
Hartert, E., 1913. Die Vögel der paläarktischen Fauna, Heft VIII- p.1081.
Hartert, E. & F. Steinbacher, 1933. Die Vögel der Paläarktischen Fauna. Ergänzungsband. 2: 97-192.— Berlin.
Hodgson, B.H., 1844. Catalogue of Nipalese Birds collected between 1824 and 1844.— [Gray’s] Zoological
Hodgson, B.H., 1845a. Characters of six new species of Nepalese birds.— Ann. Mag. Nat. Hist., (1) 15 (99): 326-327.
Hodgson, B.H., 1845b. [On Nepalese birds.].— Proc. Zool. Soc. Lond.: 22-37.
Hodgson, B.H., 1855. Catalogue of Nipalese Birds, collected between 1824 and 1844.— J. Asiatic Soc.
Bengal, 24 (7): 572-582.
Jerdon,T. C., 1871. Supplementary Notes to ‘The Birds of India’. Ibis 13: 335–356.
Inskipp, T.P., N. Lindsey & J.W. Duckworth, 1996. An Annotated Checklist of the Birds of the Oriental
Region. [i-x], 1-294.— Sandy, Beds., UK.
Rasmussen, P. C. & Anderton, J. C., 2005. Birds of South Asia: the Ripley guide. Barcelona, Lynx Editions.
Ripley, S.D., 1982. A synopsis of the birds of India and Pakistan together with those of Nepal, Bhutan,
Bangladesh and Sri Lanka. i-xxvi, 1-653.— Bombay.
Sharpe, R.B., 1874. Catalogue of the birds in the British Museum. I. Catalogue of the Accipitres or diurnal birds of prey in the collection of the British Museum.— London, Taylor & Francis.
Snow D.W. & Perrins C.M., 1998. Birds of the Western Palearctic: concise edition. Vol 1 – Non-Passerines. Oxford University Press, Oxford.
Swinhoe, R., 1870. Zoological notes of a journey from Canton to Peking and Kalgan.— Proc. Zool. Soc.
Swinhoe, R. 1871. A Revised Catalogue of the Birds of China and its Islands, with Descriptions of New Species, References to former Notes, and occasional Remarks. Proc. Zool. Soc. Lond.: 337-423.
Vaurie, C., 1965. The Birds of the Palearctic Fauna. A systematic reference. Non-Passeriformes: i-xxi, 1-
Zarudny, N.A., 1912. On the Ornithology of Turkestan – Urinator arcticus suschkini and Cerchneis naumanni turkestanicus subspp. nov. Ornithologische Mitteilungen : 114.
Warren, R.L.M., 1966. Type-specimens of birds in the British Museum (Natural History). 1: i-x, 1-
Warren, R.L.M. & C.J.O. Harrison, 1971. Type-specimens of birds in the British Museum (Natural History).
2: i-vi, 1-628.— London.
Warren, R.L.M. & C.J.O. Harrison, 1973. Type-specimens of birds in the British Museum (Natural History).
3: i-xii, 1-76.— London.
– APPENDIX I
LESSER KESTREL NOMENCLATURE (synonym)
FALCO NAUMANNI (Fleischer, 1818)
– Falco naumanni [as Naumanni] J.G.Fleischer, 1818 – in Laurop & Fischer, Sylvan. Ein Jahrbuch für Forstmänner,Jäger und Jagdfreunde auf das Jahr 1818, p.174. (“spärlicher Gast im südl. Deutschland und Schweiz”; error for Sicily, fide Stresemann (MS))
Falco xantonyx [as Xantonyx] J.G.Fleischer(ex Natterer MS), 1818- in Laurop & Fischer, Sylvan. Ein Jahrbuch für Forstmänner, Jäger und Jagdfreunde auf das Jahr 1818, p.175. (= F.naumanni)
Falco tinnunculoides Temminck (ex Natterer MS) (1820) – Manuel d’ornithologie, ou Tableau systématique des oiseaux qui se trouvent en Europe…,2nd edn,1,p.30. (“Hongrie,Autriche-Naples-Sicile-Sardaigne-Espagne”). (= F.naumanni)
Falco cenchris Frisch (1820) – in J.F.Naumann, Johann Andreas Naumann’s mehrerer gelehrten Gesellschaften Mitgliede, Naturgeschichte der Vögel Deutschlands ,2nd edn,1,p.318,pl.29. (Italy,Austria,Tyrol,Switzerland,Savoy,Piedmont).
Falco xanthonyx Naumann (ex Natterer MS) (1822) – Johann Andreas Naumann’s mehrerer gelehrten Gesellschaften Mitgliede,Naturgeschichte der Vögel Deutschlands…,1,p.323. (= F.naumanni)
Falco tinnuncularius Roux,1825 – Ornithologie provençale;ou description…de tous les oiseaux qui habitent constamment la Provence,ou qui n’y sont de passage,1,p.60,pls.40,41. (Provence). (= F.naumanni)
Falco subtinnunculus C.L.Brehm (1827) – Ornis,3 Heft,p.12. (Egypt and southern European islands). (= F.naumanni)
Cerchneis cenchris C.L.Brehm (1831) – Handbuch der Naturgeschichte aller Vögel Deutschlands,p.74. (= F.naumanni)
Tinnunculus cenchris Bonaparte (1842) – Catalogo Metodico degli Uccelli di Europa, p.21. (= F.naumanni)
Tichornis cenchris Kaup (1844) – Classification der Säugethiere und Vögel,p.108. (= F.naumanni)
Poecilornis cenchris Kaup, 1850 in W.Jardine(ed.) – Contributions to Ornithology for 1850,p.53. (= F.naumanni)
Cerchneis paradoxa C.L.Brehm, 1855- Der Vollständige Vogelfang,p.29. (Greece). (= F.naumanni)
Cerchneis ruficeps C.L.Brehm,1855,Naumannia, p.269. (= F.naumanni)
Cerchneis ruficauda C.L.Brehm,1855, Naumannia, p.269. (= F.naumanni)
Erythropus cenchris Jerdon , 1862 – The Birds of India, 1, p.38. (= F.naumanni)
Falco naumanni naumanni – Hartert,1913 [“1921”], Die Vögel der paläarktischen Fauna,Heft VIII(Aug.),p.1080. (= F.naumanni)
Cerchneis naumanni – Sharpe (1874) – Catalogue of the Birds in the British Museum,1, Accipitres or Diurnal Birds of Prey, p.435. (= F.naumanni)
Falco cenchris var. pekinensis Swinhoe (1870). Proceedings of the Zoological Society of London,p.442. (near Peking).(= F. naumanni pekinensis)
Tichornis pekinensis Swinhoe (1871). Proceedings of the Zoological Society of London,p.341. (= F. naumanni pekinensis)
Cerchneis pekinensis in Sharpe (1874). Catalogue of the Birds in the British Museum,1, Accipitres or Diurnal Birds of Prey,p.437. (= F.naumanni pekinensis )
Erythropus pekinensis in Jerdon (1871). Ibis,p.242. (= F.naumanni pekinensis)
Falco naumanni pekinensis Hartert 1913 [“1921”], Die Vögel der paläarktischen Fauna, Heft VIII(Aug.),p.1081. (= F.naumanni pekinensis)
Cerchneis angolensis Bocage, 1876 – Jornal de Sciencias mathematicas,physicas e naturas, publicado sob os auspicos da Academia real das sciencias da Lisboa 5: 153. (Huilla in Angola). (= F.naumanni)
Cerchneis naumanni turkestanicus Zarudy, 1912 – Ornithologische Mitteilungen, p.114. (= F.naumanni ssp.)
Cerchneis naumanni sarmaticus Domaniewski, 1917 – Compt. Rend. Soc. Sci. Varsovie, X, p. 1044. (= F.naumanni)