Hidden species within the genus Ocys Stephens: the widespread species O. harpaloides (Audinet-Serville) and O. tachysoides (Antoine) (Coleoptera, Carabidae, Bembidiini) Author Maddison, David R. Author Anderson, Roy text Deutsche Entomologische Zeitschrift 2016 63 2 287 301 http://dx.doi.org/10.3897/dez.63.10748 journal article http://dx.doi.org/10.3897/dez.63.10748 1860-1324-2-287 C09ADD81A914416A93388525B0E35BB0 Ocys harpaloides (Audinet-Serville, 1821) Figs 2A; 3A; 6A, B; 7A, B; 8A; 9A, B; 10A, B; 11A, B Bembidion harpaloides Audinet-Serville, 1821:78. Lectotype male, here designated, in the Dufour collection of the MNHN, examined (see Nomenclatural notes, below), with two labels: "5244" [handwritten], "LECTOTYPE Bembidion harpaloides Audinet-Serville 1821 designated 2016 D.R. Maddison and R. Anderson" [partly handwritten, with red lines around the border]. Type locality: Clavados, Bretagne, France. Tachis rufescens Guerin-Meneville , 1823:123. Type specimens lost (see below). Type locality as specified in the original description: an island on the Seine River (presumably Ile Seguin or Ile Saint-Germain), Meudon, France. Neotype male, here designated, in the MNHN, labeled "JUVISY Aval RD 4-VI-45 M. DEWAILLY" [partly handwritten], " Ocys harpaloides Serv. M. Dewailly det." [partly handwritten], "NEOTYPE Tachis rufescens Guerin-Meneville designated 2016 D.R. Maddison & R. Anderson" [partly handwritten]. This specimen is from Juvisy-sur-Orge, Essonne, France, which is 18km SE of Meudon; it was formerly housed in the collection of Mr. Dominique Echaroux. Ocys melanocephalus Stephens, 1828:10. Lectotype male, here designated, in BMNH, examined, with six labels: "BRITISH ISLES J. Stephens Coll. BM 1853- 46", " Ocys melanocephalum " [handwritten], "♂" , " harpaloides det Netolitzky" [handwritten], "NHMUK010363535" [with matrix barcode], "LECTOTYPE Ocys melanocephalus Stephens designated 2016 D.R. Maddison and R. Anderson" [partly handwritten, with red lines around the border]. Type locality: British Isles. Bembidium dubium Wollaston, 1857:23. Holotype female, in BMNH, examined. Type locality: Madeira. Junior primary homonym of Bembidium dubium Heer, 1838. Synonymy tentative, based primarily on the colour of the type. Nomenclatural notes. Interpretation of Audinet-Serville's name rests in part on discovery of his specimens, as his description is not sufficient to distinguish between the two species. Audinet-Serville notes that his specimen or specimens of Bembidion harpaloides are from "M. de Brebisson" , presumably the botanist Louis Alphonse de Brebisson . Audinet-Serville's Coleoptera specimens were acquired by Leon Jean Marie Dufour (Thierry Deuve, pers. comm.), whose collection was incorporated into the Museum National d'Histoire Naturelle in Paris. In March 2016, David H. Kavanaugh searched for several hours for the type series of Bembidion harpaloides Audinet-Serville in Dufour's collection at the MNHN, with help from Azadeh Taghavian. The following notes are in part from Kavanaugh (pers. comm.). Dufour's collection is well-organized, and in the section containing bembidiines, there is only one box with Ocys . It contains four pins with Ocys specimens. These are housed under the heading " Bembidion (Tachys) rufescens ", a name in use around that time for this complex (e.g., Dejean 1831 ). In that region of the box, there are no additional pin holes, suggesting that there have been no specimens of Ocys removed from Dufour's collection. The four pins contain a total of eight specimens: the first two pins contain one specimen each, the third pin houses five specimens, each on a separate card, and the last pin houses a single specimen. The last specimen can be excluded from consideration, as its label contains a locality ( "Carcassonne" , in southern France) that differs from the locality stated by Audinet-Serville. The other three pins each have one or two labels: the first specimen has a label that apparently contains the number 249; the second the number 5244, and the remaining pin has two labels, one with a small amount of indecipherable text, the other with the number 124. All of the Ocys in Dufour's collection belong to this species, as determined by colour pattern and microsculpture. In addition, the first two are males, and dissection revealed genitalia that match genitalic type A (i.e., as in Fig. 6A, B). The labels on the three pins are in different handwriting, and the pins themselves differ in structure. Thus, it appears as if the three came from different sources, and likely only one of them represents authentic Audinet-Serville material. According to Antoine Mantilleri (Thierry Deuve, pers. comm.), the handwriting on the first specimen is likely that of Dufour. This suggests that the first specimen is not an authentic Audinet-Serville specimen, but rather an original Dufour specimen; however, it is possible that Dufour added a label to the specimen after having acquired Audinet-Serville's collection. The writer of the other labels is not known. We could discover no information about Audinet-Serville's handwriting, other than a single example of his signature, and thus we do not know if any of the labels might have been written by him, or whether they may have been added by later researchers. Thus, it is not clear which specimen on the three candidate pins is Audinet-Serville's , and it likely will never be determined with certainty. We consider the value in stabilizing the name more important than the uncertainty of the choice. As all of these specimens belong to the same species, as the specimen on the second pin is a male in sufficient condition and with undamaged genitalia, and as there is hint that the specimen on the first pin is not an Audinet-Serville specimen, we have chosen the second specimen as the lectotype of Bembidion harpaloides Audinet-Serville. The collection of Felix Edouard Guerin-Meneville was incorporated into the Chaudoir collection at the MNHN ( Cambefort 2006 ) in the Chaudoir magasin. David Kavanaugh and Azadeh Taghavian searched for types of Tachis rufescens Guerin-Meneville in the Chaudoir magasin. They found numerous Ocys , but every one could be eliminated as Guerin-Meneville specimens through consideration of the labels, as all specimens had a mechanically printed label (against other Guerin-Meneville specimens), and the labels specified a locality, collector, or collection that did not coincide with those of the Ocys examined by Guerin-Meneville (Kavanaugh, pers. comm.). The Guerin-Meneville specimens of Ocys are thus lost. In the interests of stabilizing the nomenclature, we sought a neotype for Tachis rufescens . Within the MNHN, there are specimens from three localities around Paris (Vaires, Chelles, and Chalifert). All of these specimens belong to the following species, Ocys tachysoides (Thierry Deuve, pers. comm.); we have confirmed this for the specimens from Vaires by examination of male genitalia. However, the lack of Ocys harpaloides in the twentieth-century specimens in the MNHN does not necessarily mean that the species was absent from Paris in 1823: in the 123 years that passed between the description of Tachis rufescens and the collection of the Viares specimens in 1946, the habitats around Paris may have changed. The possibility that Ocys harpaloides lived in Paris in the 1820s is made more likely by the presence, in the collection of Mr. Dominique Echaroux, of two series of specimens of Ocys harpaloides from just outside of Paris: one specimen from Bouray-sur-Juine and seven from Juvisy-sur-Orge. The latter is only 18km SE of the type locality of Tachis rufescens . Thus, both species have lived in the general Paris area, and a specimen of either species could be designated as a neotype. However, designation of the Vaires specimen, for example, would lead to Ocys rufescens as the name of the following species, with Ocys tachysoides as a junior synonym. This would change the name used for the following species in the Iberian Peninsula and Africa, where it is known as Ocys tachysoides (per Toribio 2013 ). In the interests of stability, we choose as neotype of Tachis rufescens one of the Juvisy-sur-Orge males; with this designation, Tachis rufescens is maintained as synonym of Ocys harpaloides . In the BMNH, under "melanocephalum" , are seven specimens (four males, three females) in the J. Stephens collection, all with the label "British Isles J. Stephens Coll. BM. 1853 - 46". We consider this to be the type series of Ocys melanocephalus Stephens. Three of the males in this series were dissected, and genitalia examined; they all match genitalic type A (i.e., as in Fig. 6A, D). We have chosen one of these specimens as the lectotype. In the Wollaston collection of the BMNH, in Insecta Maderensia, Drawer 2, is a single teneral female, without a determination label. This specimen is presumed to be the holotype of Bembidium dubium Wollaston, 1854. Its colour suggests that it belongs to this species. However, the species membership of the type is of no nomenclatural importance, as the name is a junior primary homonym of Bembidium dubium Heer, 1838, itself a junior synonym of Bemidion assimile (Gyllenhal, 1810). An additional name ( Carabus tempestivus Panzer, 1799) is listed as a synonym of Ocys harpaloides by some authors (e.g. Stephens, 1828). However, this name is a synonym of Trechus quadristriatus (Schrank, 1781), as documented by Erichson (1837) . Diagnosis. Body length 4.2-6.1 mm ( Toribio 2013 and our observations; average length of males 5.0 mm (n=5), of females 5.51 mm (n=5)). Head and pronotum a clear orange-brown; anterior and central part of elytra of the same colour, but sides and posterior region of elytra dark brown (Fig. 3), and with the epipleural gutter and suture yellow-brown. Microsculpture of elytral disc more transverse, with more close-set parallel lines and fewer meshes (Fig. 7A, B); density of microsculpture lines 25-26 per 0.1 mm (n=4 males). Hind margin of pronotum in most specimens straighter than in Ocys tachysoides , such that the hind margin is directed more or less laterally near the hind angles (Fig. 10A), occasionally with a slight emargination laterally (Fig. 10B). Elytra more parallel-sided, greatest width behind middle. Basal margin at shoulder slightly arcuate with a forward-directed concavity (Fig. 11A, B). Elytral striae 2 through 4 less marked in the apical third. Aedeagus with ventral margin bend slightly downward toward apex (Fig. 6A, B); apex more rounded. Anterior edge of central sclerite complex of internal sac more rounded; brush sclerite larger; dorsal membranes of internal sac darker. Gonocoxite relatively long (Fig. 8A), GCR=0.64-0.69 (average 0.67, n=5); spermathecal margin (opposite the efferent duct of the spermathecal gland) curved (Fig. 8A, B; n=5), with tip pointed toward duct of gland. Figure 10. Hind margin of pronotum. A, B, Ocys harpaloides (A: North Slob, Wexford, Ireland, voucher DNA2759, B: Halve Maan, Oostende, Belgium, voucher DNA2853). C, D. Ocys tachysoides (B: Belvoir Forest, Belfast, Northern Ireland, voucher DNA2763, D: Murlough NNR, Co. Down, Northern Ireland, voucher DNA2899). Scale bar 0.1 mm. Figure 11. Left shoulder. A, B, Ocys harpaloides (A: North Slob, Wexford, Ireland, voucher DNA2759, B: Bos Ter Rijst, Schorisse, Belgium, voucher DNA2937) C, D. Ocys tachysoides (B: River Schelde, De Kramp, Moerzeke, Belgium, voucher DNA2936, D: Murlough NNR, Co. Down, Northern Ireland, voucher DNA2899). Scale bar 0.1 mm. Geographic distribution (Fig. 12). In Africa, from Morocco, Algeria, and Tunisia. In Europe from Spain, France, Belgium, Italy, Ireland, and the United Kingdom. Examination of specimens in additional collections will likely show it to be more widely distributed. Figure 12. Distribution map of Ocys harpaloides and Ocys tachysoides . Only records of which we are confident are included. In addition to specimens we have examined, and those from Toribio (2013) , Antoine (1956), and Sciaky (2009), we have included three publically available records from the Barcode of Life Database (http://www.boldsystems.org), corresponding to BOLD records GBCOU8213-14, GBCOU8667-14, and FBCOH116-12, as well as specimens identified by Luca Toledano and Paolo Bonavita in their collections (CTVR and PB, respectively) based upon images and diagnoses we provided to them. Specimens examined. In addition to type specimens, and those listed in Table 1, we examined specimens from Morocco: Tetouan (BMNH); Algeria (BMNH); France: Carcassone (MNHN), Bouray-sur-Juine (DE); Spain: San Roque (BMNH), Beuda, Girona (OSAC), and Sant Carles de Peralta (RAC).