The Fanniidae and Muscidae (Diptera) described ...

Zoosyst. Evol. 89 (1) 2013, 31– 166 / DOI 10.1002/zoos.201300004

The Fanniidae and Muscidae (Diptera) described by Paul Stein (1852–1921) Adrian C. Pont* Oxford University Museum of Natural History, Parks Road, Oxford OX1 3PW, United Kingdom

Abstract Received 7 November 2012 Accepted 20 November 2012 Published 18 March 2013

Key Words world type specimens type depositories

An overview is given of the species-group names in the families Fanniidae and Muscidae described by Paul Stein (1852–1921) together with their type specimens. Stein described 57 new species in the Fanniidae and 783 species in the Muscidae together with 5 replacement names, to which are added 6 unavailable species names (nomina nuda) in Fanniidae and 26 in Muscidae. A brief account of his life and career and of the various collections that he studied is given, followed by an alphabetical list of the species-group names within the Fanniidae and Muscidae. The type specimens (holotypes and syntypes) that have been located in museums and institutes throughout Europe and the United States are enumerated. The bibliography includes all the papers published by Stein on these families. The account concludes with three appendices: a systematic list of the species, a list of the species by zoogeographic region, and a list of the localities mentioned in Stein’s papers. Four lectotypes are designated (Homalomyia carbonella Stein, 1895; Homalomyia lineata Stein, 1895; Fannia nigra Stein, 1920; Mydaea pallidicornis Stein in Becker, 1910), and two replacement names for junior homonyms are given (Neodexiopsis simplicissima nom. nov. for Coenosia simplex Stein, 1920, and Helina steini nom. nov. for Aricia punctata Stein, 1898, both from the Nearctic Region).

Introduction – Stein’s life and achievement Paul Stein (1852–1921) was one of the most important workers on the muscoid families (Fanniidae, Anthomyiidae, Muscidae), and his papers and monographs laid the basis for the modern study of these families. He had a remarkable eye for species, and he wrote keys and descriptions that were concise, accurate and clear. In the family Fanniidae he described 57 new species and in the Muscidae 783 species plus 5 replacement names, to which should be added 6 unavailable species names (nomina nuda) in Fanniidae and 26 in Muscidae. Stein was born on 3 April 1852 in Wittenberg, Germany. His father, Carl Alexander Stein (1824–1902), was originally from Niemegk, and was composer, royal music director, professor, and organist and choir master at the two Wittenberg churches. He was a sufficiently eminent citizen to have had a plaque commemorating

* Corresponding author, e-mail: [email protected]

# 2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

his life fixed to the wall of the family home in what is now Jdenstrasse (Jews Street). His mother was Clara Julie Anna, ne Couard (1828–1912), daughter of a Huguenot family, and had been born in Berlin. The infant Stein, his parents’ first child, was baptised as Carl Ludwig Paul on 7 May 1852, by Dr Couard of St George’s church, Berlin. There were five further children: Elisabeth Stein (born 11 December 1854), Katharina Schmidt ne Stein (born 4 April 1856), Hermann Stein (born 9 December 1858), Ccilie Stein (born 4 June 1860), and Magdalene Rohrlach ne Stein (born 3 February 1865). Stein grew up in Wittenberg. He was the particular favourite of his grandfather Stein in Niemegk, with whom he usually spent the school holidays. He attended the local secondary school (“Gymnasium”) in Wittenberg and, in 1870 at the age of 18, enrolled at Berlin University to study mathematics and natural sciences, the subjects in which he had excelled at school. His parents were unable to support him, and so

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Pont, A. C.: Fanniidae and Muscidae (Diptera) described by Paul Stein

Figure 1. Paul Stein in a. 1905; b. 1909; c. 1919; d. 1920 or 1921; e. in the field, circa 1908.

museum-zoosyst.evol.wiley-vch.de


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he financed his studies by giving private lessons and by working for the eminent naturalist Alfred Brehm (1829–1884): between 6 and 7 am he would take dictation from Brehm in shorthand and later in the day write out the scripts in full. At the conclusion of his studies he spent a year and a half teaching at a private school in Falkenberg (Mark) during which period, in 1875, he passed his state examination. From 1877 to 1880 he was a probationary teacher (“Probandus”) and then assistant teacher (“Hilfslehrer”) in the Realschule in Aschersleben. In 1880 he accepted a post as senior teacher (“Oberlehrer”) at the newly-founded secondary school (“Progymnasium”) in Genthin, where he remained until 1907. It was whilst on holiday in Bliesendorf near Werder (Havel) that he met the woman who would become his life’s companion: Luise Rietz was born on 8 November 1860, the daughter of the Berlin postal director. They were married in Berlin on 4 October 1881, and there were three sons from the marriage. Luise survived her husband by over two decades, and died on 5 May 1945 whilst fleeing westwards from the advancing Russian army with other German refugees from East Prussia and Pomerania.

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Stein’s three sons were: Kurt, born 9 September 1882, died 28 December 1965 in Chemnitz. He qualified as a civil engineer in Berlin, but was unable to find work and moved to Chemnitz, where he was involved in various water management projects and where he remained for the rest of his life. Kurt was the family chronicler, and in the 1920s and 1930s wrote up and edited a series of volumes detailing the history and activities of the Stein and Couard families. A man of principle, he suffered through his refusal to join the Nazi party and, later, the Communist party. Peterpaul, born 10 April 1885, died in 1945. A problem child for his parents, he was not academic and would not study, and he worked as a customs secretary in Stettin. In 1945 he fled to the family home in Wittenberg and from there to Chemnitz where he died shortly afterwards. Walter, born 11 June 1893, died 9 October 1918. A very popular poet and painter, he died in Berlin from influenza. The family home in Genthin was at Knigsstrasse 2, the modern Poststrasse. However, at Easter 1907 the Progymnasium in Genthin was closed down, and Stein

Figure 2. The Stein family. a. A family holiday at Karlshagen, circa 1897, Paul and Luise Stein, with their sons Peterpaul (standing, centre), Walter (seated, centre), and Kurt (right). b. Luise Stein, with Peterpaul (centre) and Kurt (right). c. Paul and Luise Stein, with Kurt, his wife Elsa and daughter Hilde, circa 1920. d. Luise Stein, with Peterpaul (centre) and Kurt (right).



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took a teaching position as one of the “Rte vierter Klasse” (councillor fourth class) in the Royal Bugenhagen-Gymnasium at Treptow an der Rega. This is now the Public Gymnasium (Publiczne Gimnazjum) at Trzebiatow, on the Baltic coast of Poland, province Zachodnio-Pomorskie. The old Genthin school eventually reopened and is now the Bismarck-Gymnasium at 5 Grosse Schulstrasse. Stein remained in Treptow until his retirement at Easter 1921. His health had been declining for a decade, with hardening of the arteries (“Verkalkung der Arterien”), but at the beginning of 1921 he became much weaker. On 23 August 1921 a stroke partially paralysed him, and he died in his sleep in the night of 6–7 September 1921. The family gathered in Treptow on 9 September, when the coffin was taken from the house to the church and then opened for the final farewells. Stein’s sister Magdalena wrote: “Unser lieber Paul sah wunderhbsch aus, unverndert und ganz verklrt” [“Our dear Paul looked absolutely beautiful, unchanged, and wholly transfigured”]. He was buried in the German cemetery in Treptow on 10 September. The funeral service included his favourite hymn “Jerusalem” and an anthem performed by the school


choir. The three houses in which the Stein family lived during their life in Trzebiatow are still standing, but the German cemetery has been destroyed and Stein’s grave can no longer be found. His son Kurt (K. Stein 1922) has described his father’s appearance and temperament. He was of medium height, well-built, and until his final years his health was robust. Taciturn by temperament, he was nevertheless physically restless and unable to remain sitting or still for long. As a father he was loving but strict, demanding at all times unconditional obedience and a strict adherence to truth. Typically, the children had to remain at the table at mealtimes until every scrap of food on their plates had been consumed. Yet Kurt remembers the freedom and the time they had to play, although he remarks somewhat ruefully that this was because his father was always spending so much time with his Diptera rather than with them. Stein taught mathematics, natural sciences and French. In the classroom he was a stern disciplinarian, and in those days corporal punishment was the norm. Kurt recalls that as he passed through his father’s classes he was given double the number of beatings or

Figure 3. Some of the Stein residences. a. Genthin, Knigsstrasse 2. b. Treptow. c. Paul Stein’s writing desk in Treptow. d. Paul Stein’s second house in Treptow.



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clouts, to avoid any accusations of favouritism. But Stein was a conscientious teacher, and his instruction was clear, factual and to the point. It was whilst on a visit to his cousin, the hymenopterist and Symphyta specialist Richard Ritter von Stein (1847–1933), a medical doctor in Chodau (now Chodov in the Czech Republic), that Stein was encouraged to take up the Diptera. This he did, with enormous enthusiasm and dedication, and rapidly focussed on the Calyptrata. At the time, the problematic and difficult “Anthomyidae” included most of the modern Muscidae and Fanniidae in addition to the Anthomyiidae, and he was soon known as “Anthomyid-Stein”. Richard Ritter von Stein’s father, the brother of Paul Stein’s father, was Friedrich Ritter von Stein, born on 3 November 1818 at Niemegk, died 9 January 1885 in Prague. He was professor of zoology at the University of Prague and was ennobled with a hereditary title because of his great contributions to zoology and his great four-volume work of 1859–1863, Der Organismus der Infusionsthiere. Incidentally, after graduation, he spent the year 1849 as a curator in the Zoologisches Museum Berlin. His wife was also a member of the Couard family. Stein was an avid and skilful collector, and rapidly built up a large and comprehensive collection. He carried his net everywhere and was completely in his element in the field. His children would follow him – at a distance, so that they did not disturb the flies – and often he would simply disappear from sight and they would only know where he was by the grunts of satisfaction that signalled the capture of an interesting specimen. One of his most productive localities was his own garden, and he refers to a species “. . . zu Genthin in meinem mitten in der Stadt gelegenen Garten an einer denselben durchziehenden Gosse gefangen” (“caught at Genthin in my garden, situated in the middle of the town, by a ditch running through the garden”) (1910, Wiener entomologische Zeitung 29: 19). His collection consisted of some 15000 specimens of 3000 species, stored in over 80 boxes. Like many an entomologist, his study and in particular his work table was a noli me tangere for the whole family. The table was covered with boxes, books, papers, tubes, hand lenses and equipment of all kinds. Stein was a modest man, and when honours eventually came his way he accepted them with pleasure but kept them to himself. What was important to him and what gave him the greatest pleasure and satisfaction in life was the study of flies, and he was totally uninterested in making money from these studies. His travels, to Paris, London, Lund, Vienna, were undertaken at his own expense. The life of the intellect was all-important for him, and his son Kurt regretted that his father spent so little time in “normal” family activities and conversations. He was a competent and largely self-taught linguist, being proficient in French and English, with a good knowledge of Latin, Greek and Hebrew, and a working knowledge of Russian, Swedish,


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Danish, Dutch and Italian. His knowledge of classical languages is apparent in the species-group epithets that he coined, which are almost invariably well-chosen, well-formed, euphonious and appropriate. He was a voracious reader but an unwilling speaker, and the speeches which he had to deliver on special occasions at school caused him much concern. His handwriting, as can be seen in his letters and labels, was small, neat, but barely legible. He was a competent pianist but otherwise inherited little of his father’s musical talent. His life was disciplined and focussed. For example, he got up and went to bed at the same time every day, summer and winter, schooldays and holidays. The Prussian virtues of duty, punctuality and economy ruled the household – how else could he have achieved so much in what was for him a spare time activity, a hobby. But life was not always serious, and within the broad family circle he was a lively companion and enjoyed meeting up with the several generations of the Stein clan in Germany. The one great tragedy of his life was the early death of his youngest son, a blow from which he never fully recovered. Nevertheless, he was fortunate that his other two sons, Kurt and Peterpaul, served in the First World War and both survived. Stein was a gifted and, for the time, an unusually conscientious taxonomist. He had an excellent eye for species, and his strong point was the recognition, description and accurate naming of species, rather than the generic or higher classifications. Kramer (1922) offered some gentle criticism of him for not taking up the work on the genitalic structures by other contemporary taxonomists such as Girschner and Schnabl. However, through his great monographic works, his detailed descriptions, his keys, his catalogues, and his study of types and enumeration of synonyms, he left the muscoid families in an infinitely better state than that in which he found them, bringing a semblance of order into the chaotic plethora of names that had accumulated during the 19th century. As Lichtwardt (1905: 28) wrote: “Stein hat wie wenige das Vermgen, eine schwierige Materie zu entwirren und einzelne Teile greifbar klar herauszuziehen . . . Trotz Meade, Strobl, Pokorny, Schnabl etc. ist die Bestimmung einer Anthomyide immer noch eine Art Rtselaufgabe, deren Lsung oft nach stundenlanger Mhe aufgegeben wird, um das betreffende Tier – an ‘Stein’ zu schicken” [“Like few others, Stein has the ability to disentangle difficult material and to extract individual parts in a tangibly clear manner . . . In spite of Meade, Strobl, Pokorny, Schnabl etc., it is still always a puzzling task to identify an anthomyid, and often after hour-long efforts the attempt has to be given up so that the specimen in question – may be sent to ‘Stein’.”]. One only needs to glance at some of his major works (e.g. Stein 1914a, 1916, 1919a) and compare them with the monograph by his contemporary Schnabl (Schnabl & Dziedzicki 1911) to see the respective merits of both. Schnabl’s work was ground-breaking in many ways, with its numerous drawings and micrographs of the male termina-


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Figure 4. Paul Stein’s handwriting. a–d. Samples of Stein’s correspondence. a. A card to the English dipterist George Henry Verrall (1848–1911), dated 13.ii.1907 and from Genthin. b. Part of a letter to Verrall, dated 11.ii.1909 and from Treptow. c. A card to the English dipterist James Edward Collin (1876–1968), nephew of Verrall, dated 21.ix.1909 and from Treptow. d. A card to Collin, dated 15.xi.1912. e–g. Examples of Stein’s data labels. e. 20.v.1891, locality illegible but in the Genthin district, Germany. f. 6.vi.1906, locality “W” is Wittenberg, Germany. g. “Phaonia debilis sp n” (Muscidae), determination label on a syntype. Scale line 5 mm.



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lia, but the text is verbose, impenetrable, complicated and now virtually unusable. Stein’s papers however are clear and concise, to the point, and still usable. His classification and keys were superseded in the first decades after his death, but his descriptions are excellent and are still helpful a century or more later. In keeping with his modesty and devotion to truth, Stein was completely without the “mihi itch” so often found among 19th century entomologists. He was a living exemplar of Hermann Hesse’s aphorism, “The truth is lived, not taught”. If he found that one of his species was a synonym, he published the synonymy without more ado. In fact, he had to sink one of the first species that he described, Hydrotaea eximia (Stein 1888b), within two years of its description (Stein 1890a). When publishing his first paper on North American species, he complained about the poor quality of Walker’s descriptions and, being unable to recognise the species and being informed by the British Museum (Natural History) that the types could not be loaned to him (“. . . ich wurde aber auf eine diesbezgliche Anfrage belehrt, dass, wie der Ausdruck lautete, they are not allowed to leave the building” [“. . . but in response to such an enquiry I was informed that, as the expression goes, they are not allowed to leave the building”] (Stein 1898: 162)), he proceeded with his own descriptions in order to open up the field of North American “Anthomyidae”. But, unusually for the time, he pursued the study of types whenever he had the opportunity, and within five years had visited London, revised the Walker muscoid types, and along the way synonymised a number of his own North American species (Stein 1901). Over the years he studied the types of almost all the major workers on “Anthomyidae” such as Wiedemann, Meigen, Zetterstedt, Walker, Jaennicke, Schiner, Thomson, Boheman and Holmgren (see below). In a review of Stein’s (1924) posthumously-published work on the Tachinidae of Europe, the French dipterist Joseph Villeneuve de Janti gave a most sympathetic and heartfelt appreciation of Stein the dipterist and of Stein the man (Villeneuve 1931): “Le mmoire du Prof. P. Stein sur les Tachinides europens est, comme tous les travaux du savant diptrologue, une oeuvre d’un prcieux intrt et d’une longue patience. Ecrit avec un sens aigu des bons caractres, dont le choix judicieux donne aux descriptions cette limpidit et cette clart familires aussi l’auteur, ce mmoire reflte admirablement l’tat actuel de nos connaissances sur les Tachinides et surtout l’esprit qui, selon moi, doit prsider leur tude. Stein se montra toujours, en effet, le naturaliste-n, au sens linnen du mot. Il resta fidle la grande espce, peut-tre parce qu’il dlaissa l’tude des pieces genitales au moindre detail desquelles trop de nos contemporains prtent inconsidrment une valeur absolue, prpondrante dans tous les cas . . . J’tais li avec le Prof. Stein par une profonde amiti, ne le premier jour o nous nous conn mes. Rien n’branla jamais cet attachement rciproque qui reposait tout entier sur un gal amour de la


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Vrit et de toutes les lumires susceptibles de la faire jaillir. Il nous arriva souvent de discuter, et de disputer, avec la meme tnacit, avec le meme acharnement. C’tait toujours courtois et loyal. Tel fut Stein, tel je le retrouve dans son mmoire sur les Tachinides o il ne me mnage pas ses critiques quand il les croit justes.” [“Professor Stein’s memoir on the European Tachinidae is, like all the papers by this scholarly dipterist, a work of invaluable interest and prolonged patience. Written with an acute feeling for good characters, the judicious selection of which lends to the descriptions that limpidity and clarity that are familiar also to the author, this memoir reflects admirably the current state of our knowledge of the Tachinidae and above all the spirit which, in my opinion, should prevail in their study. In fact, Stein always showed himself to be a born naturalist, in the Linnaean sense of the word. He remained true to the large species, perhaps because he did not take up the study of the genital structures the smallest detail of which is unthinkingly given an absolute value, or a dominant one in every case, by too many of our contemporaries . . . I was united with Prof. Stein by a deep friendship that was born on the first day that we met. Nothing ever shook this reciprocal attachment which was based entirely on an equal love of the Truth and of all the illuminations that might reveal it. We were often in the situation of discussing, of arguing, with the same tenacity, the same determination, but this was always courteous and loyal. Such was Stein, and so do I find him in his memoir on the Tachinidae where he does not spare me from his criticisms when he believes them to be just.”] Stein joined the Entomologischer Verein in Berlin in the second half of 1887 (Berliner Entomologische Zeitschrift 31: xxix) and is in the list of members (op.cit.: lii) simply as “Stein, P., Gymnasial-Lehrer, Genthin”. He left the society in 1907 (Berliner Entomologische Zeitschrift 51: under the unpaginated “Vereinsangelegenheiten II”), following his move to Treptow. He then joined the Deutsche Entomologische Gesellschaft in 1909 (Deutsche Entomologische Zeitschrift 1909: 12), where he was entered as “Stein, Prof. P., Treptow a.R.”. He remained a member of this Society until his death, which was recorded in Deutsche Entomologische Zeitschrift 1921: 303. Collections received and their fate Stein’s personal collection Stein worked on collections of “Anthomyidae” from all regions of the world, describing 57 new species in the family Fanniidae, 783 in the Muscidae and 195 in the Anthomyiidae (1035 species; other unavailable names excluded). His personal collection contained the material that he collected himself, material that was given to him or exchanged with his contemporaries in Europe (especially Becker, Oldenberg, Lichtwardt, Bezzi, Riedel, Schnuse, Kuntze, Villeneuve and, later, Karl and


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Ringdahl), and also duplicates that he kept from the collections that were sent to him for study. His collection was thus of exceptional importance, and was purchased by the then Zoologisches Museum, Berlin (now: Museum fr Naturkunde, Leibniz Institut fr Evolutions- und Biodiversittsforschung an der HumboldtUniversitt zu Berlin) for 3000 Marks. This sale is recorded in a letter dated 29 November 1921 from Stein’s son Kurt to the Diptera curator at the Zoologisches Museum, Gnther Enderlein: “Im Auftrag meiner Mutter biete ich dem Museum die Dipteren-Sammlung meines verstorbenen Vaters, Prof. P. Stein – Treptow (Rega), zum Preise von dreitausend Mark zum Kauf an und zwar nach dem Vorschlag des Herrn Direktor Geheimrat Prof. Kkenthal unter Ausschluss derjenigen Objekte, die das Museum nicht bentigt, deren Auswahl dem Museum freisteht und dem Verkauf im Interesse meiner Mutter noch erfolgen soll. “Wegen der Preisforderung verweise ich auf das Angebot der Naturalien-Sammlung in Stuttgart in Hhe von 5–6000 M., das wir aber nicht bercksichtigen mchten, da uns das Museum in Berlin sympathischer erscheint.” [“On behalf of my mother, I am offering to the Museum the Diptera collection of my late father, Prof. P. Stein – Treptow (Rega), for the purchase price of three thousand Marks and, following the suggestion of Director and Privy Councillor Prof. Kkenthal, with the exclusion of those objects which the Museum does not require, the selection of which lies at the discretion of the Museum and the sale of which should be made in the interests of my mother. So far as the sale price is concerned, I refer you to the offer from the Natural History collection in Stuttgart for as much as 5–6000 Marks which, however, we would not like to consider as the Museum is Berlin appears to us to be more sympathetic.”] The collection was packed and moved to the Berlin Museum by Enderlein himself, and he recorded the receipt of the collection in the Museum’s Accessions Register as follows: “No. 916 Sammlung Prof. Stein, Treptow, bes. Anthomyiiden mit zahlreichen Typen. Aus allen Erdteilen, besonders aus Deutschland. Jan. 1922. [Von] Frau Prof. Stein. [Wert] 3000 Fr 3000.– M gekauft. Wert viel hher.” “No. 916 Collection of Prof. Stein, Treptow, especially Anthomyiidae with numerous types. From all regions of the world, particularly from Germany Jan. 1922. [From] Frau Prof. Stein. [Value] 3000 Purchased for 3000.– Marks. Value much greater.”



In January 1922, just before the beginning of hyperinflation, 3000 German marks were worth around US$ 15.50, which had the modern buying power of almost US$ 200.00 – a derisory sum to pay for such a collection. Collections and types studied by Stein Stein collected intensively around his home towns of Wittenberg, Genthin and Treptow (Trzebiatow), and also on Rgen Island, Bornholm Island (Denmark), Lund (Sweden), Kissingen, and Chodau (the home of his cousin Richard von Stein). As his reputation as an anthomyid specialist grew, so did the number of his international contacts and colleagues. Whilst working all his life on his own collection and on the European collections that were sent to him, the mid-1890s saw an expansion in the range of his work. He was sent North American material by the American dipterists Hough, Aldrich, Johnson and Nason and published an extensive work on these (Stein 1898). One of his last papers was a second long paper on the Nearctic fauna (Stein 1920c). This was followed by requests from the Hungarian Natural History Museum, Budapest, and this gave rise to papers on “Anthomyidae” from all regions of the world (Stein 1900b, 1900d, 1904a, 1910c, 1913a, 1918). From Amsterdam he received material from the then Dutch colonies in Indonesia and New Guinea (Stein 1904b, 1909, 1919b, 1920a, 1920b). He also studied the unidentified “anthomyid” material of the Zoologisches Museum, Berlin (Stein 1906). Expedition material from other sources was also sent to him, for example from Central Asia, the Seychelles Islands, Sokotra, South-West Africa, “British East Africa”, Taiwan (Sauter), New Guinea, South America (Schnuse), etc. As discussed above, when faced with the huge number of names in the literature, many of which were unidentifiable, and realising that existing interpretations of older names was frequently a matter of guesswork, Stein undertook the study of types, either by personal visits to Museums or through loans. In this way he reviewed types in all the leading collections in Europe: – Lund: Falln and Zetterstedt types, following a visit to Lund in July 1892 (Stein 1892b). The Lund guestbook records his visit as 9 July 1892. – Paris: Meigen types (Stein 1900c). – Copenhagen: visited for several days in the 1890s (Stein 1895: 112). – London: Walker types (Stein 1901). – Stockholm: Zetterstedt, Holmgren, Boheman and some Falln types were sent to him on loan (Stein 1902a); Thomson types were also sent to him on loan (Stein 1910a). – Frankfurt: Wiedemann and Jaennicke types were sent to him on loan (Stein 1902b). – Vienna: the entire muscoid collection was sent on loan for revision, including types of Schiner’s “No-


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– –

– – –

vara Reise”, Brauer & Bergenstamm, and the collections of von Winthem and Wiedemann (Stein 1919a). Cambridge: Lamb’s New Zealand types (Stein 1919a). Newmarket: the private collection of G. H. Verrall, with types of Macquart and Bigot, through a visit and subsequent loans (Stein 1907a, 1907b). He visited Verrall on 29 January 1894. Amsterdam: van der Wulp types, sent on loan by de Meijere. Stuttgart: von Roser types sent on loan. Rambouillet: the private collection of J. Villeneuve, whom he visited in 1900.

Whilst Stein’s personal collection was sold after his death to the Zoologisches Museum in Berlin, the fate of several of the other collections that he studied requires some discussion. Some derived from private collections as well as from public Museums, and so material studied and/or described by him can be found in numerous Museums and other institutes scattered through Europe and the United States. Some, regrettably, have been destroyed, but it is fortunate that he always retained duplicates for his own collection when there were series of two or more specimens. At the conclusion of this section I list a selection of those individuals who contributed material recorded in Stein’s papers, which may have been returned to them or may have been kept by Stein, together with a note on the current location (where known) of their personal collections. Alluaud & Jeannel. Stein (1914b) worked on the material brought back by the 1911–1912 French expedition to the mountains of East Africa (see Alluaud & Jeannel 1912). However, this entire “anthomyid” collection has been lost, most probably destroyed during the conflict between Germany and France, 1914–1918. The late Loc Matile of the Musum national d’Histoire naturelle, Paris, told me that it had never been returned to Paris. I have not traced a single specimen. Becker, Theodor. One of Stein’s most fruitful collaborations was with Becker, who was also an assiduous collector. Stein worked on “Anthomyidae” for the various reports that Becker prepared on his own collecting and on collections sent to him for study. He collected in Egypt in 1898–1899; Canary Islands in 1901; and Canary Islands and Madeira in 1904 (see Stein 1903, 1908a, 1908b, 1910, 1913c, 1913d, 1922). Becker’s personal collection is also in the Zoologisches Museum, Berlin, and his material can be recognised by the very characteristic labels which are all handwritten and contain two lines of data: 1. Locality, e.g. “Odessa”. 2. Month (Roman) and code number, e.g. “IX. 50437”. There are four notebooks that give the key to the code numbers. Each number may refer to a single specimen


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or to a series of conspecific specimens. Entries under each number include the species and number of specimens, locality and sometimes precise notes on collecting site and/or habitat, date of capture, year, identity, and, in the case of longer series, the names of those to whom Becker gave specimens and how many. It appears that Becker did not always send all his material to Stein: Stein sometimes lists a specific number of syntypes from individual localities (Egypt, Canary Islands, Madeira), whilst Becker’s notebooks mention many more specimens. Budapest, Hungary. Stein (1900b, 1900d, 1904a, 1910c, 1913a, 1918) described a very large number of species from the collections of the Hungarian Natural History Museum, Budapest, mainly from South America, Africa (collectors Katona and Kov cs), and the Oriental Region and Australia (collector Bir ). Unfortunately, most of the Diptera collection in that museum, including all the Fanniidae and Muscidae, was destroyed by fire in 1956 (Fldv ry & Papp 2007), except for the types of six Stein species of Lispe (and some others), which I found in 1986 (Lispe albimaculata, L. cyrtoneurina, L. geniseta, L. ignobilis, L. metatarsata, L. vittata) and some species of Atherigona which were out on loan at the time. It is fortunate that Stein kept duplicates of the species that he described in his personal collection, sometimes even the second of two specimens, but there were a number of unique holotypes that were destroyed and many of those species can no longer be recognised. It should be noted that on the Kov cs labels the month “11” refers to February (ii) and not to November (xi). Emden (and the Natural History Museum, London) had some exchanges with the Hungarian Natural History Museum (1913a and 1918 papers) and so small numbers of syntypes are also in the Natural History Museum, London. Cambridge University, UK. This material is mounted in a very characteristic manner. The flies are pinned with minuten pins on to either square or oblong cork mounts, and some locality data are written on the mount. Stein (1913a, 1918) described a number of species from the Cambridge University collection, most of which have apparently been lost. Emden (1940: 92) wrote: “My thanks are due to . . . Mr. G. C. Varley of the University Museum of Zoology, Cambridge, who informed me that the species described by Stein from the Cambridge Museum have never been returned and seem to be lost.” This is indeed the case, but the specimens that Stein himself kept from Cambridge (collectors Frederick Muir and Guy Marshall) have been discovered by me in the Museum fr Naturkunde, Berlin. Stein (1910b) also reported on “Anthomyidae” from the Seychelles Islands, collected by the Cambridge coleopterist Hugh Scott. This collection was split between the Cambridge University Museum and the Natural History Museum, London, with some duplicates being retained by Stein in his collection. Data on the mounts consist of the name of the individual island and the


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years of the expedition, and a number written on the cork mount, which may not always be clearly legible (e.g. “104” and “109” are not always distinct). I have had access to a manuscript list compiled by Hugh Scott of the localities where collections were made during the Seychelles Expedition, July 1908 to March 1909, which gives a key to the numbers on the mounts. The following list enumerates the numbers found on the muscid types and gives the localities as provided by Scott.

1 Silhouette: near Mont Pot--eau, all over 1000 feet, viii.1908. 19 Silhouette: Mare aux Cochons, ix.1908. 24 Silhouette: Mare aux Cochons, ix.1908. 29 Silhouette: forest above Mare aux Cochons, highest point of Silhouette, 2.ix.1908. 30 Silhouette: Mare aux Cochons and forest just above, 23.ix.1908. 36 Mah: from near Morne Blanc, x.1908. 48 Praslin: Ctes d’Or estate, especially from Cocode-Mer forest in the Valle de Mai, 28 & 29.xi.1908. 55 Mah: Port Victoria, from grass in Botanic Gardens, xii.1908. 60 Mah: Cascade Estate, 800–1000 feet (no date given). 65 Mah: Cascade Estate, 800–1000 feet, i.1909. 67 Mah: near Morne Blanc, 1908. 93 Mah: Cascade Estate, ii.1909. 98 Mah: Cascade Estate, forest, 1000–2000 feet, ii.1909. 101 Silhouette: plateau of Mare aux Cochons, ix.1908. 102 Silhouette: forest above Mare aux Cochons, about 1400 to over 2000 feet, ix.1908. 103 Silhouette: near Mont Pot--eau, about 1500 feet, viii.1908. 104 Mah: Cascade Estate, about 800 feet and over (H.P. Thomasset & H. Scott), x.1908–i.1909. 105 Mah: cultivated country about 1000 feet, xi.– xii.1908. 107 Mah: Long Island, 12–22.vii.1908. 108 Mah: coast-marsh at Port Glaud, 5.xi.1908. 109 Mah: Cascade Estate, about 1000 feet and over (no date). 110 Mah: Fort Noire district, x.–xi.1908. 111 Mah: high forest of Morne Blanc and Pilot, x.– xi.1908. 112 Mah: from near Morne Blanc, x.–xi.1908. 114 Mah: Morne Seychellois, over 1500 feet, 4.ii.1909. 115 Mah: summit of “Montagne Anse Major”, about 2000 feet, forest of rather stunted “Capucin” trees (Northea), 1.ii.1909. 117 Mah: Mare aux Cochons district, about 1500 feet, 26.i–2.ii.1909. 120 Mah: Cascade Estate, about 800–1500 feet, 1909.



131 Mah: various localities, from shady part of Fort Noire road near Casse-les-Dents, about 1000 feet, x.–xi.1908. 134 Mah: Cascade Estate, iii.1909. 141 Mah: Anonyme Island, from grass, ferns, trees, etc, i.1909. 142 Mah: marshy ground near sea-level, Cascade, 20.ii.1909. 146 Mah: marshes on coastal plain at Anse aux Pins and Anse Royale, 19.–21.i.1909. Field Museum of Natural History, Chicago, USA. The personal collections of G. de N. Hough and W. A. Nason are in this Museum. Hough’s collection is said to have suffered much damage and loss before it was acquired by the Museum, but a great deal still survives. His specimens are preserved together with his handwritten drawer labels, which are written in the following style: “HOMALOMYIA/depressa/Stein”, to which “Type” is added in pencil. Each specimen also has a printed label “Univ. of Chicago/G. H. Hough/Diptera Colln.”. There is no Nason material in the Illinois Natural History Survey, and his whole collection appears to be in the Field Museum. His material can be recognised by the following style of label: “Algonquin, Ill. [printed]/8.12.95 – 134 [handwritten]”. North America. Stein (1898) described a large number of new species, based on material sent to him by G. de N. Hough, J. M. Aldrich, C. W. Johnson and W. A. Nason. At the conclusion of the main body of his text (Stein 1898: 282–288) he included a series of “Verbesserungen und Nachtrge”, which contained descriptions of further new species and also additional material of some of the species described earlier in the paper. For example, to the species Limnophora cyrtoneurina n. sp. (described on page 203) he added “H. (La.) zahlreich” [Hough, Louisiana, abundant] (page 285). I regard all such additionally listed specimens as part of the typeseries and have listed them here as syntypes. St Petersburg, Russia. The specimens from Central Asia collected by V. I. Roborowsky and P. K. Kozlov that were sent to Stein (1907c) have labels in Cyrillic. But Stein relabelled most of those that he retained for his own collection and discarded the original Cyrillic labels. Taiwan. All the material collected on Formosa (Taiwan) and worked on by Stein (1915b, 1918) was collected by H. Sauter. That published in 1915 was from the then Deutsches Entomologisches Institut whilst that published in 1918 was from the Hungarian Natural History Museum. Emden (and the Natural History Museum, London) had exchanges with the Hungarian Natural History Museum (1913a and 1918 papers) and the Deutsches Entomologisches Institute (1915b paper) and so some Taiwan syntypes are now in the Natural History Museum, London.


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Stein appears to have made a few errors in giving the dates of collection. It seems most likely that he was overwhelmed by the sheer number of specimens before him and the amount of data that he was having to transcribe. This was also the case with his 1911 report on the South American collection of Schnuse (see Pont 2001). Collectors. The following list gives a selection of the collectors of material seen by Stein. Some of these are not mentioned in Stein’s papers but their names appear on collection labels. Where known, the depository of their collections is given in square brackets (for collection acronyms, see further below): Aldrich, John Merton (1866–1934), American dipterist [USNM] Alfers: see under Olfers Alluaud, Charles (1861–1949), French coleopterist [MNHNP] Baker, Charles Fuller (1872–1927), American entomologist and botanist [USNM] Beaufort, Lieven Ferdinand de (1879–1968) Dutch zoologist and one-time director of the Zoological Museum of Amsterdam [NBCL] Beaufort-Boissevain, Catharina Josephina (1885–1922), wife of de Beaufort [NBCL] Beccari, Odoardo (1843–1920), Italian traveller and naturalist [MCSNG] Becker, Theodor (1840–1928), German architect and dipterist [ZMHU] Bescke, Carl Heinrich (1798–1851), German commercial agent and collector [ZMHU and others] Beyrich, perhaps Heinrich Karl (1796–1834), German botanist [ZMHU] Bezzi, Mario (1868–1927), Italian university professor and entomologist [MCSNM] Bir , Lajos (1856–1931), Hungarian traveller and collector [HNHM (destroyed)] Bolsius, Alphonsus Maria Johannes (1844–1903), Dutch medical doctor in Indonesia [NBCL] Bonsdorf, Evert Julius (1810–1898), Finnish entomologist [ZMUH] Bradley, James Chester (1884–1975), American entomologist [CUIC] Brauns, Johannes (1857–1929), German hymenopterist [ZMHU & SDEI] Brunetti, Enrico Adelelmo (1862–1927), Italian bandmaster and dipterist [HNHM (destroyed) & BMNH] Bttner, Oskar Alexander Richard (1858–1927), German collector [ZMHU] Conradt, Leopold [Leopoldo] (fl. 1875–1922), German collector in Africa and natural history professor in Mexico [ZMHU & SDEI] Czerny, Leander Franz (1859–1944), Austrian abbot and dipterist [NMW] Czwalina, Gustav (1841–1894), German [East Prussian] entomologist [ZMKR] Dahl, Karl Friedrich Otto Theodor (1856–1929), German collector and zoologist [ZMHU]


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Dmel, C. F. Eduard (1821–1900), German collector and entomological dealer [ZMHU] Dammermann, Karel Willem (1885–1951), Dutch agricultural entomologist in Indonesia [NBCL] Dampf, Alfons (1884–1948), Estonian dipterist [ZMKR] Dedem, Frederik Karel Baron van (1873–1959), Dutch postal official, ornithologist and collector [NBCL] Drge, Charles Frederic or Carl Friedrich (1791–1867), German collector and entomological dealer [scattered] Ehrenberg, Christian Gottfried (1795–1876), German traveller and collector [ZMHU] Fischer (??–??), Austrian, collector [NMW] Frey, Karl Richard Hjalmar (1886–1965), Finnish dipterist [ZMUH] Fruhstorfer, Hans (1866–1922), German insect dealer and lepidopterist [scattered] Flleborn, Friedrich (1866–1933), German collector and scientist [ZMHU] Garlepp, Gustav (1862–1907), German collector in South America [SMT] Girschner, Ernst (1860–1914), German schoolmaster and dipterist [BMNH] Gredler, Vincenz Maria (1823–1912), German priest and coleopterist [?] Gssfeld, Paul (1840–1920), German geologist, mountaineer and explorer in Africa [?ZMHU] Hensel, Reinhold Friedrich (1826–1881), German zoologist and collector [ZMHU] Hine, James Stewart (1866–1930), American dipterist [OhSU] Homeyer, Alexander von (1834–1903), German army officer and collector [ZMHU] Hsemann, Paul Alfred (1868–1922), German collector [ZMHU] Hough, Garry de Neuville (1861–1927), American medical doctor and dipterist [FMNH] Jacobson, Edward Richard (1870–1944), German/ Dutch self-taught zoologist and collector in Indonesia [NBCL] Jeannel, Ren Gabriel (1879–1965), French coleopterist [MNHNP] Jensen-Haarup, Anders Christian (1863–1934), Danish schoolteacher, hymenopterist and collector in Argentina [ZMUC] Johannsen, Oskar Augustus (1870–1961), American dipterist [CUIC] Johnson, Charles Willison (1863–1932), American dipterist [MCZ] Kampen, Pieter Nicolaas van (1878–1937), Dutch agriculturalist in Indonesia, later university professor in The Netherlands [NBCL] Karl, Otto Johann Hermann (1868–1945), German schoolteacher and dipterist [PAN, mostly destroyed, and SDEI] Katona, label name for K lm n [Coloman] Kittenberger (1881–1958), Hungarian zoologist and collector in Africa [HNHM (destroyed)]


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Kaznakov, Alexander Nikolaevich [Kaznakov Aleksandr Nikolaeviq] (1872–1933), Russian zoologist [ASSP] Kertsz, Kalman (1867–1922), Hungarian dipterist [HNHM (destroyed)] Kittenberger: see under Katona Kobus, Jan Derk (1858–1910), Dutch businessman and collector in Indonesia [NBCL] Knig, Eugen Gregor (fl. 1893–1931). Georgian coleopterist and entomological dealer, curator at the Kaukasus Museum in Tiflis (now Tbilisi) [ASSP] Koningsberger, Jacob Christian (1867–1951), Dutch naturalist and director of the botanical garden in Buitenzorg, Indonesia [NBCL] Kov cs, dn [Edmund] (1886–1919), Hungarian zoologist and collector in Africa [HNHM (destroyed)] Kowarz, Ferdinand (1838–1914), Austrian postal official and dipterist [OUMNH & ASSP] Kozlov, Peter [Pjotr] Kus’mich [Kozlov, Petr Kuzvmiq] (1863–1935), Russian traveller and collector [ASSP] Krebs, Ludswig (1792–1844), German traveller and collector in Africa [ZMHU] Krieghoff, Edmund Heinrich Christian (1844–1906), German pastor and collector [SMF] Kuntze, Albert Friedrich Arthur (1842–1933), German traveller and collector [SMT] Leeuwen, Jan van (1850–1924), Dutch entomologist [NBCL] Lichtwardt, Bernhard (1857–1943), German businessman and dipterist [SDEI] Lorentz, Hendrikus Albertus (1871–1944), Dutch lawyer and diplomat, collector in New Guinea [NBCL] Loria, Lamberto (1855–1913), Italian traveller and collector [MCSNG] Lutz, Adolfo (1855–1940), Brazilian doctor and medical zoologist [MNRJ] Mann, William Montana (1886–1960), American entomologist and zoologist [USNM & MCZ] Mannerheim, Carl Gustav von, Count (1797–1854), Finnish coleopterist [ZMUH] Marshall, Guy Anstruther Knox (1871–1959), British coleopterist [BMNH] Meade, Richard Henry (1814–1899), British surgeon and dipterist [BMNH] Meijere, Johannes Cornelis Hendrick de (1866–1947), Dutch dipterist [NBCL] Melander, Axel Leonard (1878–1962), American dipterist [USNM] Mik, Josef (1839–1900), Austrian dipterist [NMW] Monteiro, Rose (1840–1897), botanist and wife of Joaquin John Monteiro (1833–1878), naturalist and collector in Maputo, Mozambique [?] Moritz, Johann Wilhelm Karl (1797–1866), German collector in the West Indies and South America [?] Morrison, Herbert Knowles (1854–1885), American collector and insect dealer [USNM] Muir, Frederick Arthur Godfrey (1872–1931), British Hemipterist [BMNH]



Mller, Heinrich Ludwig Hermann (1829–1883), German botanist, entomologist and schoolmaster [?] Nason, William Abbott (1841–1918), American naturalist and collector [FMNH] Nietner, Johannes Werner Theodor (1828–1874), German collector, scientist and plantation owner in Sri Lanka [ZMHU] Oldenberg, Lorenz (1863–1931), German dipterist [SDEI] Olfers, Ignaz Franz Werner Maria von (1793–1871), German travelling naturalist and collector [ZMHU] Osten Sacken, Carl [Charles] Robert, Baron von der [Osten-Saken Robert Romanoviq] (1828–1906), Russian diplomat and dipterist [MCZ, ASSP, SDEI] Palmn, Johan Axel (1845–1919), Finnish dipterist [ZMUH] Pandell, Louis (1824–1905), French entomologist [MNHNP] Penther, Arnold (1865–1931), Austrian traveller and collector [NMW] Peters, Wilhelm Carl Hartwig (1815–1883), German traveller and collector in Africa [ZMHU] Petersen, Wilhelm Konstantin Frommhold (1854– 1933), German entomologist [ZMHU] Philippi, Rodolfo Amando (1808–1904), German-Chilean palaeontologist and zoologist [some ZMHU] Piepers, Marius Cornelis (1835–1919), Dutch-Indonesian lawyer and government official, lepidopterist in Indonesia [NBCL] Pokorny, Emanuel (1838–1908), Austrian dipterist [HNHM (mostly destroyed)] Pll, Josef (1874–1940), Austrian naturalist and schoolteacher in Innsbruck [?] Pppig, Eduard Friedrich (1798–1868), German botanist, zoologist and traveller [MNSL] Poppius, Bertil Robert (1876–1916), Finnish entomologist [ZMUH] Preuss, Paul R (1862–1926), German collector [ZMHU] Reimoser, Eduard (1864–1940), Austrian arachnologist and coleopterist [NMW] Ribbe, Carl (1860–1934), German entomologist [?SMT] Riedel, Max Paul (1870–1941), German dipterist [ZMHU] Ringdahl, Oscar (1885–1966), Swedish schoolmaster and dipterist [ZML] Roborowsky [Roborovsky], Vsevolod Ivanovich [Roborovskij Vsevolod Ivanoviq] (1856–1910), Russian traveller and collector [ASSP] Rder, Viktor von (1841–1910), German dipterist [MLUH] Rolle, Hermann (1864–1929), German insect dealer in Berlin [scattered] Sahlberg, Johan Reinhold (1845–1920), Finnish entomologist [ZMUH] Sauter, Hans (1871–1948), German collector in Taiwan [SDEI, some formerly in HNHM (destroyed)] Schillings, Carl Georg (1865–1921, German collector in East Africa [ZMHU]


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Schmidt, Carl Wilhelm (1859–1924), German collector [?] Schnabl, Johann Andreas [Jan Sznabl], (1838–1912), Polish medical doctor and dipterist [PAN (destroyed)] Schnuse, Carl August Wilhelm (1850–1909), German collector in South America [SMT] Scholtz, Heinrich (1812–1859), Silesian conchologist [ZMHU] Schultze, Leonhard Sigmund (1872–1955), German zoologist and anthropologist in SW Africa (Namibia) [ZMHU] Schulz, Wilhelm Albert (1871–??), German insect dealer [ZMHU] Scott, Hugh (1885–1960), British coleopterist [CUM and BMNH] Siebeck, Alexander (fl. 1885–1913), Austrian forest superintendent [?] Sikora [Sicora], Franz (1863–1902), Austrian naturalist, fossil and insect collector in Madagascar [NMW] Silvestri, Filippo (1873–1949), Italian economic entomologist [MCSNG] Simony, Oskar (1852–1913), Austrian entomologist [NMW]. Sintenis, Franz (1835–1911), Estonian entomologist [PAN (destroyed)] Staudinger, Otto (1830–1900), German lepidopterist [ZMHU & SMT] Steinheil, Eduard (1830–1879), German collector [?] Stieglmayr, Fr. (fl. 1895–1907), collector in South America [NMW & SMT] Strobl, Gabriel (1846–1925), Austrian monk and naturalist [BSA] Tessmann, Theodor Gnther (1884–1969), German/ Brazilian collector and scientist [ZMHU] Thalhammer, J nos (1847–1934), Hungarian Jesuit priest and collector [HNHM (destroyed)] Thieme, Theodor Otto (1857–1907), German collector and scientist [ZMHU] Tief, Wilhelm (1846–1896), Austrian entomologist [NHMK] Ude, M. (??–??), German collector [ZMHU] Uhle, E. (??–??), German collector or dealer [ZMHU] Verrall, George Henry (1848–1911), British racecourse manager and dipterist [OUMNH] Villeneuve de Janti, Joseph (1868–1944), French medical doctor and dipterist [IRSNB] Werner, Franz Josef Maria (1867–1939), Austrian traveller and collector [NMW] Woldstedt, Frederik Wilhelm (1847–1888), Finnish hymenopterist [ZMUH] Wstnei, Wilhelm Heinrich Carl Franz (1839–1907), Danish entomologist [ZMUC] Zarudny, Nikolai Alexeievich [Zarudnyj, Nikolaj Alekseeviq] (1859–1919), Russian ornithologist, traveller and collector [ASSP] Zenker, Georg August (1855–1922), German collector [ZMHU] Zimmermann, Charles Christoph Andrew (1800–1867), American entomologist [MCZ & ANSP]


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Types and their recognition When Stein’s collection was received by the Berlin Zoologisches Museum in 1921, it was dealt with personally by the then Diptera curator, Gnther Enderlein (1872–1968). He incorporated the Muscidae (including Fanniidae and Anthomyiidae) into the Museum’s Palaearctic and Exotic collections. Stein’s locality labels on specimens that he collected personally contain only the most basic information and are frequently difficult to decipher. Typically, he gives only the date and an initial for the locality, for example: “G” for Genthin “Rg” for Rgen “Tr” for Treptow “W” for Wittenberg “Kiss” for Kissingen Enderlein added to many of these specimens a label with the locality and collector printed and with the date handwritten up the left-hand margin, e.g. “Prov. Sachsen / Genthin / Prof. P. Stein S.” and “30.5.1890”. “S” ¼ “Sammler”, collector. However, he sometimes misread the date, and so it is always necessary to check Stein’s own label and not to rely on the Enderlein label. Stein’s label is pinned beneath Enderlein’s label, and the date is usually clearly legible although the locality cipher frequently is not. It is necessary to correct here an error in Pont & Werner (2006). The abbreviation “R” refers to an unnamed locality in the vicinity of Genthin and not to Rgen island in the Baltic Sea. In Enderlein’s handwritten notes on Stein’s localities and labels, preserved in the Diptera section of the Zoologisches Museum, “R” and “Rg” are listed as referring to Rgen. However, two pencil corrections have subsequently been made, which were not visible in the photocopy from which I was working at that time: the “R” has been deleted, and has been added to Genthin: “ G. — R: Genthin”. As a result of this, some changes have been made to the composition of type-series as listed in Pont & Werner (2006). Stein rarely attached an identification label to his own material or indeed to much of the European material in his personal collection. However, there is sometimes a label, presumably the original species label pinned with the series in his collection, which is attached to one of the specimens in the series. It is a small rectangular black-edged label, bearing the species name and author in Stein’s hand. In the text below I refer to Stein’s own locality or identification label as “Stein’s label”, and the additional label printed and attached by Enderlein as “Enderlein’s label”. So far as the Exotic material is concerned, Enderlein added a red printed “TYPUS” label to specimens that he considered to be types. In some cases, especially those that were returned to museums outside Germany, he also attached a printed label “det. P. Stein” or “P.


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Stein det.” to which he added the identification and a comment such as “Cotypus”. Sometimes he replaced the original data label with one of his own, handwritten on coloured paper according to the zoogeographic region, e.g. pale green for Nearctic, pale purple for Australia. Sometimes he added a one-line label with the country handwritten on paper coloured according to the zoogeographic region. Sometimes he made handwritten additions to Stein’s own handwritten determination labels, adding the sex or “Typus” or the year of description. From time to time his labelling appears to have been haphazard, as red “TYPUS” labels have been attached to specimens that are not types and, conversely, undoubted syntypes have not been recognised and labelled with their status. Specimens that Stein borrowed for study from other museums vary in the standard and quality of the labelling. Old specimens, naturally, have little more than a country and a collector, and are handwritten. More recent specimens, such as those collected by Bir or by Jacobson, have printed labels with country, locality, date and collector. To these, Stein attached a handwritten identification label. Stein’s identification labels are helpful only to a limited extent, whether he was dealing with old or new species. If specimens were unique he labelled them, but if there was a series of specimens he would label only one or two. In labelling new species, he was rather careful at the start of his descriptive activity, labelling his 1906b new species as, for example, “Spilogaster / abnormis / sp. nov. / < Stein”. Later the labels become less discursive, e.g. “Fannia / minuta / sp. n.”. The fact that he did not label every syntype certainly creates problems, and difficulties also arise in cases where he synonymised one of his own species: the syntypes were then transferred to a series bearing the older name, and he apparently removed his original “sp. n.” labels. It is also the case that he appears not to have written “sp. n.” in every case but sometimes to have written “Stein” after the species name. It could be thought that specimens labelled in this way are later identifications of his own species rather than syntypes, but this appears not to be the case as there are examples of undoubted syntypes with “Stein” rather than “sp. n.” on the label. In a few cases the generic name on the label is different from that under which the species was published, e.g. “Mydaea rufitibia” on the label for the published Aricia rufitibia. In all these cases, it could be that he actually wrote labels for the syntypes long after he had described them and published them, and so incorporated the later information such as “Stein” rather than “sp. n.” and a different generic combination on to his labels. It does not follow that we are dealing with a later identification simply because “sp. n.” does not appear on the label. It also has to be borne in mind that some of Stein’s labels may have been transferred from one specimen to another when his collection was being incorporated into the main Zoologisches Museum collection. This would certainly explain some of the anomalies in the labelling.



At all events, one has to act pragmatically, and my purpose in compiling this account is to spread the net widely and to list all possible syntypes. Those working on generic revisions in the future may disagree in detail, but in any case the designation of a lectotype to fix the identity of a name will finalise the nomenclatural status of all actual and putative syntypes.

Stein’s species of Fanniidae and Muscidae Unless otherwise stated, all the material listed here has been seen by me at some time or another over the last half-century. Others who have reported on Stein’s types include H.C. Huckett (1890–1989), who visited Berlin in 1932 to study the Nearctic types, and W. Hennig (1913–1976), who studied the Palaearctic types whilst preparing his revision of the Palaearctic Muscidae (including Fanniidae) Except where stated, the condition of specimens is good. A comment “right hind leg missing” means that the specimen is in good condition and well preserved except for the missing leg. In citing label data, a forward slash (/) indicates the end of a line. It is not necessary, nor is it practicable at this point in time, to cite every label on every specimen verbatim. I have given what is necessary for the accurate recognition of types, referring back to my previous reviews of certain of Stein’s species (Pont 1966a, 1966c, 1968, 1969b, 1970, 2001, 2004; Pont & Werner 2006). The information contained in those papers is not repeated here, but reference is made back to them where all relevant details may be found. The following abbreviations are used for museums and institutes referred to in the text below (and in the section under “collectors” above): ANSP ASSP BMNH BSA CUIC CUM FMNH HNHM INHS IRSNB

Philadelphia Academy of Natural Sciences, Pennsylvania, U.S.A. Zoological Institute of the Russian Academy of Sciences, St Petersburg, Russia The Natural History Museum, formerly British Museum (Natural History), London, U.K. Naturhistorisches Museum der BenediktinerStift, Admont, Austria Cornell University Insect Collection, Ithaca, New York, U.S.A. University Museum of Zoology, Cambridge, U.K. Field Museum of Natural History, Chicago, Illinois, U.S.A. Hungarian Natural History Museum, Budapest, Hungary Illinois Natural History Survey, Urbana, Illinois, U.S.A. Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium


Zoosyst. Evol. 89 (1) 2013, 31– 166

LACM MCSNG MCSNM MCZ MLUH MNHNP MNRJ MNSL NBCL NHMK NRS NMW OhSU OUMNH PAN SDEI SMF SMT UKaL USNM ZMHU

ZMKR ZML ZMUC ZMUH

Los Angeles County Museum, Los Angeles, California, U.S.A. Museo Civico di Storia Naturale di Genova, Genova, Italy Museo Civico di Storia Naturale di Milano, Milan, Italy Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A. Martin-Luther-Universitt, Halle a.S., Germany Musum National d’Histoire Naturelle, Paris, France Museo Nacional, Rio de Janeiro, Brazil Museum of Natural Sciences, Leipzig, Germany Naturalis Biodiversity Centre, Leiden, The Netherlands Landesmuseum fr Krnten, Klagenfurt, Austria Naturhistoriska Riksmuseet, Stockholm, Sweden Naturhistorisches Museum Wien, Vienna, Austria Ohio State University, Columbus, Ohio, U.S.A. Oxford University Museum of Natural History, Oxford, U.K. Polish Academy of Sciences, Warsaw, Poland Senckenberg Deutsches Entomologisches Institut, Mncheberg, Germany Senckenbergisches Museum, Frankfurt am Main, Germany Staatliches Museum fr Tierkunde, Dresden, Germany Snow Entomological Collection, University of Kansas, Lawrence, Kansas, U.S.A. National Museum of Natural History, Washington, D.C., U.S.A. Museum fr Naturkunde, Leibniz Institut fr Evolutions- und Biodiversittsforschung an der Humboldt-Universitt zu Berlin, Berlin, Germany Zoological Museum, Knigsberg [now Kaliningrad], Russia Zoological Museum, Lund University, Lund, Sweden Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark Zoological Museum, Finnish Museum of Natural History, Helsinki Finland

FANNIIDAE abnormis Stein, 1900d: 210 (,; Homalomyia). “2 , aus S. Antonio und Songo” [BOLIVIA]. HNHM: Syntypes 2 , destroyed in 1956. ZMHU: No syntypes found – there are specimens identified by Stein and some (from Callanga) incorrectly labelled as types.


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Current name: Fannia abnormis (Stein, 1900). albitarsis Stein, 1911: 105 (< ,; Fannia). “6 Mnnchen und 5 Weibchen aus Guayacan (Chile) 22.9.02, Coquimbo (Chile) 23.9.02, Caldera 25.9.02, vom Titicacasee (Bolivia) 25.11.02, 29.5.03, Juliaca (Peru) 15.6.02 und Sicuani (Peru) 19.6.03. In der Garlepp’schen Sammlung findet sich ein in Cuzco 23.9.02 gefangenes Mnnchen” [BOLIVIA, PERU, CHILE]. SMT: Lectotype < (des. Pont & Carvalho 1994: 232; see also Pont 2001: 459–460), from Guayacan, 22.ix.1902; paralectotypes 1 < 3 ,. ZMHU: Paralectotypes 2 < 1 , (Pont 2001: 482; Pont & Werner 2006: 6). Note: 5 paralectotypes have not been traced. Current name: Fannia albitarsis Stein, 1911. antennata Stein, 1911: 103 (