A collection of 457 specimens of Formicidae in 195 taxa from all over the World donated by Auguste Forel has been conserved since 1901 at the Naturmuseum Solothurn, Switzerland (NMSO). Hardly noticed by the scientific community, the collection includes 40 syntypes of 16 species and subspecies, and five paralectotypes from two species. To spread this knowledge, and to encourage the inclusion of the Formicidae collection in future taxonomic studies, all species from the collection are listed, and the types are catalogued and imaged.
INTRODUCTION
Auguste Forel (1848–1931) had been a passionate myrmecologist since his childhood. Born in Switzerland near Morges in the canton of Vaud he completed his medical studies in the year 1871 in Zurich. After a short stay in Vienna, where he deepened his knowledge in the field of neuroanatomy and neuropsychiatry, he passed the cantonal medical examination in Lausanne one year later. In 1873 Forel moved to Munich, where he intensively worked on his anatomical studies of the human brain (Parent, 2003) as well as on the Swiss ant fauna, crowned with his extensive work “Les fourmis de la Suisse” (Forel, 1874). From 1879 to 1898 he was director at the Burghölzli asylum in Zurich, and professor of psychiatry at the University of Zurich. Auguste Forel travelled a lot, and in 1889 and 1893 went on journeys to Tunisia and Algeria, in 1891 to Bulgaria (Forel, 1892b), and in the last years of his working career and after his retirement, he intensified travelling and visited Colombia in 1896, the United States and Canada in 1899, Eastern Europe and Russia in 1902, Egypt, Algeria, Tunisia, Italy in 1908, and in the following year he travelled from the Balkans to Greece and Turkey (Parent, 2003; Banani, 2005). After having suffered a stroke with partial paralysis in 1912 Forel was forced to stop his excursions.
In 1922 the main collection of Auguste Forel was sold to the Muséum d'histoire naturelle de Genève (MHNG). Apart from that collection doublets were given in 1923/1924 to the Zoological Museum Berlin, and a specific Swiss collection is conserved since the same date in the Musée de zoologie de Lausanne (MZL). A small overview collection remained in the possession of Forel's family (Horn & Kahle, 1935) and is conserved, presumably entirely, since 1981 also in the MZL (teste A. Freitag). In 1911 Forel visited the Zoologische Staatssammlung München (ZSM) (at that time Königlich Zoologisches Museum München) where he determined and described Formicidae, which was not mentioned by Horn & Kahle (1935). A series of types are therefore also at the ZSM (Merta, 1999). Similarly at the Naturhistorisches Museum der Burgergemeinde Bern (NMBE), not mentioned either in Horn & Kahle (1935), part of the Forel collection is conserved since 1911, comprising ant species from all over the World and syntype specimens. Interestingly, in the extensive online catalogue on biographies of entomologists of the Senckenberg Deutsches Entomologisches Institut, Müncheberg (SDEI Senckenberg, 2015) the Forel collection at the Naturmuseum Solothurn (NMSO) is mentioned.
Bloch (1940) provided an overview on all 457 specimens (from 174 species out of 64 genera at that time) of the Forel collection from localities all over the World conserved at the Naturmuseum Solothurn. The collection was first mentioned by Bloch (1911), where Forel himself is mentioned as donator of the collection. This was part of a common effort to contribute to the growing museum collection supported by other contemporaries (as Paul Born, Arnold Wullschlegel, E. Christ, Wilhelm Roos, Eduard Von Jenner, Fritz Rohr, Karl Ris, Emile Frey-Gessner, M. Bedot) under the lead of August Rätzer. At the NMSO the Forel collection is registered under the number “E. K. [Eingangskatalog = accessions catalogue] 112″ and is indicated as a donation dating back to 27th December 1901, long before his main collection was sold to the MHNG.
In the meantime the collection, formerly housed in an external depot, remained unexplored and unfortunately single specimens were destroyed by feeding of carpet beetles (Dermestidae). These damages – if relevant, e.g. not only single legs or antennae are missing – are reported in the following. The re-examination of species described (and mostly collected) by Forel, and comparisons with the very helpful AntWeb (2016) – where all historical literature is available – resulted in the discovery of syntype specimens. These were not yet adequately labelled and hence not recognisable as types at first sight. The knowledge of the existence of this ant collection at the NMSO is important for future taxonomic examinations. Therefore these specimens are listed and pictured in the following.
The determinations are those of Forel himself. Therefore, difficult genera and species-groups with subsequent new discoveries and/or species that have been split in the meantime should be carefully revised by any interested myrmecologist; a necessity which should be emphasized with this present contribution.
MATERIAL AND METHODS
Photos were taken with a 3.15-megapixel digital camera (ProgRes CT3) on a stereomicroscope (Nikon SMZ 1000) for the ant specimens, and a digital camera (Canon G 11) for the labels. Series of images were captured with ProgRes Capture Pro 2.8.8 for Windows and stacked with the freely available CombineZP Image Stacking Software by Alan Hadley.
Additional remarks and corrections to label data are set in square brackets [ ], mistakes by Bloch (1940) are additionally highlighted by a preceding asterisk at the beginning of the square brackets [*]. Specimens from different localities are separated by a dash (—). The original combination used on the labels in the Forel collection to every species is given in curly brackets { }. All specimens of the Forel collection are registered in the database of the NMSO and these data are available on request for further investigations. The Forel collection at the NMSO is conserved in three regular sized insect boxes (51 × 42 × 6 cm). As a peculiarity, but usual at that time, only the first specimen of a series of separately pinned specimens is provided with a label.
RESULTS
Corrections, damaged and lost specimens
From the originally 457 specimens of the Forel collection, currently 426 from 195 taxa are still present in the collection of the NMSO (Appendix 1). Corrections as well as indications about damages or lost specimens are given for the 41 species below. 40 specimens are damaged and a few are lost. Bloch's paper (1940) contained a number of mistakes which are shown and corrected here under the respective species (ordered in alphabetical order):
- Acromyrmex disciger (Mayr, 1887) {Atta (Acromyrmex) discigera} 2 Blumenau [Brasilia] [leg.] Möller [head of one is lost].
- Aphaenogaster gibbosa (Latreille, 1798) 3 Souk Ahras [*Algeria; not Persia as in Bloch (1940)].
- A. treatae Forel, 1886 2 Tyorn 3400 “[feet] W.B. [has to be in the USA] 23.VII. [leg.] Forel.
- Camponotus cruentatus (Latreille, 1802) 3 près Montpellier, sous pierre [under stone] [one worker is heavily damaged; gaster, one leg and part of the head capsule are missing].
- C. maculatus (Fabricius, 1782) 3 Moyanga [=Moronga] Madagascar [the gaster of one of the workers is lost].
- Cyphomyrmex rimosus (Spinola, 1851) 3 , 1 ♀ St. Vincent [São Vicente, Brasil] [of the ♀ only head and mesosoma are left].
- Dolichoderus quadripunctatus (Linné, 1771) 2 Vaux [the head of one is missing].
- Dorylus helvolus (Linnaeus, 1764) 3 Basutoland [today Lesotho] 14.VI. [*not 1914, as the collection dates back to 1901] [leg.] Wronghton.
- Eciton carolinense Emery, 1894 3 Faisons [= Faison] N. C. [North Carolina, USA] ds.[dans] tronc [in (tree) trunk].
- E. hamatum (Fabricius, 1782) 3 Bonda Columbia [leg.] Forel [one specimen without head].
- Forelius maccooki (McCook, 1880) 2 Austin Texas [collector illegible] [*no locality was given by Bloch (1940)].
- Formica pallidefulva Latreille, 1802 3 V...mera [illegible] Morganton [several localities in the USA are possible] N....trnc à mat [illegible] [one is lost].
- F. rubicunda Emery, 1893 {F. sanguinea rubicunda Emery, 1893} 2 Tyson N.C. [North Carolina] coll. excl. subsericea 20.VII. [collected from Formica subsericea Say, 1836; provenance unclear according to Bloch (1940)].
- Formicoxenus nitidulus (Nylander, 1846) 1 luz. Ural geb. [Ural mountains ?].
- Goniomma hispanicum (André, 1883) 2 Camargue [*France; instead of unknown locality in Bloch (1940)] [leg.] Forel.
- Harpegnathos saltator cruentatus (Smith, 1858) {H. cruentatus} 1 Kanara [*India; instead of “Kanada” in Bloch (1940)] [leg.] Bell 8. [18]66.
- Iridomyrmex purpureus (Smith, 1858): 2 Mackay, Queensland [Australia] [leg.] Turner [gasters of both specimens are missing].
- Lasius alienus (Foerster, 1850) 2 : Adelsberg [not specified if in Germany or Austria; several localities are possible; one is partly destroyed, gaster and two legs remain].
- L. emarginatus (Olivier, 1792) 3 : Adelsberg [same as above: not specified if in Germany or Austria; several localities are possible] 21.IX. [the head of one specimen is lost].
- L. flavus (Fabricius, 1782) 2 ♂, 1 ♀ Fisibach [canton AG, Switzerland] 7.VIII. [both ♂ are lost]. — 3 Fisibach.
- L. umbratus (Nylander, 1846) 2 ♂, 1 ♀ Burghölzli [Zurich] 30.IX. [one ♂ is lost]. — 2 : München [Bayern, Germany].
- Leptogenys processionalis (Jerdon, 1851) {L. ocellifera (Roger, 1861)} 3 [*instead of no sex indicated] India, Ceylon.
- Liometopum microcephalum (Panzer, 1798) 2 : Sare-Mussa [Bulgarien; the head of one specimen is lost].
- Manica rubida (Latreille, 1802) {Myrmica} 1 ♀, 1 Murgthal [Murgtal near Quarten canton SG, Switzerland] 9.IV. [both specimens almost destroyed].
- Messor arenarius (Fabricius, 1787) 2 [both severely damaged] Gabès [Tunesia] [leg.] Forel.
- M. barbarus (Linné, 1767) 1 ♀ Perrégaux [*today Mohammadia; not Perrigana as given by Bloch (1940)] Algérie 29.III. — 3 [one is completely destroyed] Laverdure [today Mechroha] Algérie, [leg.] Forel.
- M. structor (Latreille, 1798) 1 ♀, 2 [*not 3 as mentioned in Bloch (1940); and additionally 4 plant seeds glued on the edges of a label] Miramar Baléares [Spain; not Switzerland as interpreted by Bloch (1940)].
- Myrmica lobicornis Nylander, 1846 3 Elinine Vrh [*Rilo-Dagh mountains, Bulgaria; not “Schweizer Alpen” as mentioned by Bloch (1940)].
- M. nigrocincta Smith, 1858 1 : Zullusland [Australia] [whole specimen lost; labels still present in the box].
- M. rubra (Linné, 1758) {M. laevinodis Nylander, 1846} 1 ♀, 3 Bali – Effendi [=Aféndis, Crete, Greece] [the ♀ is completely destroyed].
- M. ruginodis Nylander, 1846 3 [one is completely destroyed] Rilo mo…otis [*illegible, most probably Rilo mountains; Bulgaria instead of “In der ganzen Schweiz” (Bloch 1940)] .
- M. rugulosa Nylander, 1849 3 Murg [near Quarten canton SG]; [one completely destroyed, another one partly destroyed].
- M. scabrinodis Nylander, 1846 2 ♂, 1 ♀ and 1 [without head] Vaux [near Morges] 8.VIII. — 1 : Vaux [near Morges] [the ♀ and one ♂ are completely destroyed].
- M. sulcinodis Nylander, 1846 3 , 1 ♀ [*not only 3 as mentioned in Bloch (1940)]: Schluderbach [= Carbonin, Italian; South Tirol] [1 is completely destroyed].
- Pheidole megacephala (Fabricius, 1793) 1 Manà Prov. Rio [Brasilia] [leg.] Göldi. — 2 : Madeira [Portugal] [leg.] Schmitz [one of them is completely destroyed]. — 1 ♀ Nosibé [Madagascar] [leg.] Völtzkow.
- P. pallidula (Nylander, 1849) 1 ♂ Ghadame [= Ghadamès, Lybia] (Ali). — 2 , 1 ♀ Onsernone [*canton of Ticino, Switzerland; not mentioned in Bloch (1940)] [one is almost destroyed (gaster remaining), and parts of the gaster of the second one are missing]. — 2 : Algier [leg.] Moser. 1 [only gaster plus postpetiolus and petiolus are remaining] Tebessa [leg.] Forel.
- Tapinoma erraticum (Latreille, 1798) 1 ♂ Vaux. — 1 ♀ Horgen 1.VII. — 2 Vaux [gasters of both workers are missing].
- Temnothorax recedens (Nylander, 1856) 2 [*not suberis Forel, 1894 as mentioned in Bloch (1940)] parc près Montpellier.
- T. tuberum (Fabricius, 1775) 3 , 1 ♂ [1 and the ♂ are destroyed] Salève [France, near Geneva] 6.VI.
- T. unifasciatus (Latreille, 1798) 3 [where one is destroyed and another one heavily damaged (head missing)] Vaux [above Morges, Switzerland].
- Tetramorium guineense (Bernard, 1953) {T. guineensis (Fabricius, 1775)} 1 [completely destroyed] MacKay Queensland [Australia] 107 (Tarus) [?].
Annotated and illustrated catalogue of type specimens
In the following, 40 syntype specimens from 16 species and subspecies, 5 paralectotypes from two species and three former syntypes from a mixed type series are listed. Label data are mentioned verbatim in quotation marks. All type specimens were not yet marked or labelled as type specimens and hence difficult or not at all recognizable as such. Photos of the specimens and its original labels are given (Figs 1A-4S), and are available on request as maximally resolved images for scientific purposes. The types belong to species and subspecies published by Forel in the years 1887, 1890, 1892a, b, 1894a, b, 1899, 1901, and 1902. All species were compared with the original descriptions and – if type specimens were available – with AntWeb (2016). Details given in the descriptions, when necessary, are provided and additional references are included under the respective species (listed here in alphabetical order within the respective subfamilies):
Dolichoderinae
Azteca aurita Emery, 1893 {A. lacrymosa Forel, 1899} 3 ancient syntypes of a mixed type series (), which have lost their syntypical status: “A. lacrymosa Forel type Esperanza, Colombie ([leg. ] Forel) ...arton [illegible]”. Longino (2007) revised the Azteca aurita-species group, and designated lectotypes. The syntype series is a mixed one and consists of two species: Azteca pilosula Forel, 1899 and A. aurita, to which the Columbian specimens are assigned nowadays.
A. delpini antillana Forel, 1899 (Figs 3C, 4P) 3 syntypes (): “A. Delpini Sm r. [race] antillana For Sta Lucia (Antille) [leg.] Forel”. The locality fits perfectly with the description by Forel (1899: 111): “Hab. [Habitat] Antilles, St. Lucia (Forel)”.
A. velox Forel, 1899 (Figs 3B, 4O) 3 ): “A. velox Forel Burithaka (Columbien) [=Buritaca, Colombia] [leg.] Forel”. Forel (1899) mentioned his (re-)description: “…Colombie, pied de la Sierra Nevada (Forel)” which fits well with the label's locality, located north of the Sierra Nevada de Santa Marta. However, A. velox is a replacement name for A. coeruleipennis var. fasciata Pergande, 1896, based on homonymy. Hence Forel's specimens of A. velox are not syntypes, but are included and illustrated here to avoid misinterpretations.
Formicinae
Camponotus alii Forel, 1890 (Figs 3F, 4S) 3 syntypes (): “C. alii Forel Djebel Ozmor p. Tebessa, 1380 m (Forel) Type”. The label data corresponds perfectly with the original description by Forel (1890: LXIII) where he stated: “Forêts de pins du Djébel Ozmor près Tébessa de 1100 à 1300 mètres...”.
C. bugnioni Forel, 1899 (Figs 1E, 4E) 3 syntypes (): “C. Bugnioni Dibulla, Colombie (Forel)”. Fits perfectly with the indicated type locality-area given by Forel (1899: 131): “Hab. [Habitat] Colombie, forêts du pied de la Sierra Nevada de Santa Marta, de Dibulla à Santa Marta.”
Cataglyphis savignyi (Dufour, 1862) {Myrmecocystus viaticus desertorum Forel, 1894} (Figs 3E, 4R) 2 syntypes (): “M. viaticus v. desertorum Gabès, Tunesien”. Forel (1894b) mentioned Gabès as the locality of C. desertorum, described as n. v. [nouvelle variété]. However, C. desertorum is at present a junior synonym of C. savignyi.
Lasius myops Forel, 1894 (Figs 3D, 4Q) 2 syntypes (): “L. myops Forel Terni [Algeria; not Umbria as mentioned by Bloch 1940] 9.IV.” One of the – originally three – is lost. The locality fits with the description (Forel, 1894a: 12), where he indicated: “Vallon près de Terni à plus de 1400 mètres d'élévation, sous les pierres, dans les lieux boisés.” Furthermore, Forel's journey to Algeria was undertaken from 27th March to 19th April 1893, which corresponds with the given collection date (9th April).
Myrmicinae
Cardiocondyla stambuloffii Forel, 1892 (Figs 2C, 4J) 3 syntypes (): “C. Stambuloffii ForelBurgas [Bulgaria] 13. VIII”. The type locality and the time fit perfectly with Forel (1892b), who gave Burgas, Anchialo and Sozopolis as localities and 13th to 16th August 1891 as time frame of his excursion.
Crematogaster daisyi Forel, 1901 (Figs 2F, 4M) 4 paralectotypes (3 , 1 ♀) [*not 4 ♀ as mentioned by Bloch (1940)] : “Cr. Daisyi Forel ♀ type Sarawak 76. [1876 ?] Haviland [=Borneo]”. As a lectotype and paralectotypes have been designated very recently by Hosoishi (2015), these formerly syntype specimens are paralectotypes according to article 73.2.2. of the ICZN (2016).
C. ranavalonae Forel, 1887 [*not ranovalonae (sic!) as cited by Bloch (1940)] (Figs 2E, 4L) 3 syntypes (): “C. ranavalonis [sic!] Forel Bois Ivondro p. [près] Tamatavé (Madagascar)”. The locality corresponds perfectly with the one given by Forel (1887: 389).
Messor lobicornis Forel, 1894 (Figs 1D, 4D) 1 syntype (): “Terni [*Algeria, Tlemcen; not Umbrien as mentioned by Bloch (1940)] 9.IV”. Forel (1894a: 32) gives the locality as follows: Caravansérail d'Aïn Ghoraba, près de Terni, dans une prairie. The label is identical with the one pictured from another syntype from the Forel collection of the MHNG on AntWeb (2016).
Monomorium indicum Forel, 1902 (Figs 1C, 4C) 3 syntypes (): “M. Salomonis r. [race] indicum Belgaum [=Belagavi, Karnataka, India] XXXI.12 (Wronghton)”. Although Forel (1902: 213) does not give any localities under indicum, he mentions “Belgaum” and the collector (Wronghton) under the species described above in the text (which is Monomorium dichroum Forel, 1902). Therefore it seems logical that M. indicum was collected at the same locality as M. dichroum, and therefore the locality on the label is the type locality, and that the present specimens belong to the syntype series.
M. smithii Forel, 1892 (Figs 1F, 4F) 3 syntypes (): “M. Smithii Type Forel, Ashburton N. Zealand (Smith)”. The label data of the syntypes fits perfectly with the locality given by Forel (1892a: 13), where he mentions “Ashburton, Neu-Seeland [New Zealand] (W. W. Smith)”.
Myrmica smythiesii Forel, 1902 (Figs 1B, 4B) 2 syntypes (): “M. Smythiesii Forel Cas ha mere [Cashmere] Smythies H. [18]79.” Forel (1902: 227) wrote in the description: “Diverses localités de l'Himalaya, de 7000′ à 12000′ [feet] (Smythies).” This includes the present indication. Furthermore the specimen was collected by Smythies himself while he travelled in the Himalayas.
Pogonomyrmex mayri Forel, 1899 (Figs 3A, 4N) 1 paralectotype (): “Sta. Martha [Santa Marta, Colombia]”. Johnson (2015) designated the lectotype and paralectotypes based on syntypes examined at the NHMW.
Solenopsis latro Forel, 1894 (Figs 2D, 4K) 3 syntypes (): “S. latro Forel Souk Ahras [Algeria] chez A. [Aphaenogaster] sardoa (Forel)”. In his description Forel (1894a: 21) gives the same data: “Souk Ahras (Algérie orientale), en nid double avec l’Aphaenogaster sardoa.”.
Strongylognathus huberi Forel, 1874 (Figs 1A, 4A) 2 syntypes (): “S. huberi Fully” [Switzerland, canton Valais]; 1 ♀ “Str huberi ♀ Type Fully 28.VIII”. Forel (1874) mentioned that both males and females, were unknown to him when describing S. huberi. Therefore, the single female – although marked with the word “Type” – obviously does not belong to the syntype series. The label of the workers fits perfectly with further syntype specimens shown on AntWeb (2016).
Temnothorax algiricus trabutii (Forel, 1894) (Figs 1G, 4G) {Leptothorax angustulus r. [race] trabutii Forel, 1894} 2 syntypes (): “L. angustulus Nyl. r. [race] Trabutii [sic!] Forel Les Trembles Algérie”. The given locality fits perfectly with the indications by Forel (1894a: 3738): “Tas de bois et de racines d'arbustes à Tlemcen, Terni, les Trembles, Hammam bou Hadjar.”
T. delaparti (Forel, 1890) {Leptothorax} (Figs 2A, 4H) 2 syntypes (): “L. de la Parti ForelDjebel Ozmor p. [près] Tebessa 1380 m. [Algeria] Forel”. Fits perfectly with the description (Forel, 1890: LXXIV), where he stated: “Sommet du Djébel Ozmor (1380 mètres), près de Tebessa (Algérie)...”.
T. oraniensis (Forel, 1894) {Leptothorax nigrita r. [race] oraniensis} (Figs 2B, 4I) 3 syntypes (): “L. nigrita r. [race] oraniensis Forel type Aïn Fezza Algérie (Forel)”. Fits perfectly with the indications in the description (Forel, 1894a: 35): “Franchetti, Aïn Fezza, Tlemcen, en fourmilières fort populeuses, situées sous les pierres des prairies un peu, mais pas trop rocailleuses. A Aïn Fezza, j’en ai trouvé plusieurs fourmilières dans une prairie située à côté de la gare.”
ACKNOWLEDGEMENTS
I am grateful to Elsa Obrecht (Küttigkofen) for her linguistic advice and the referees for their helpful comments and additions.
REFERENCES
Appendices
Appendix 1.
All 426 Formicidae from the Auguste Forel collection at the NMSO are listed with given locality, canton (for Swiss localities), states/nations, the year they were collected (if given), the numbers of specimens (m = males; q = queens; w = workers), the collector (leg.) and the presence of type specimens.