Hypoponera punctatissima

Hypoponera punctatissima (Roger) (Figs 97 – 102) Ponera punctatissima Roger, 1859: 246, pl. 7, fig. 7. Syntype workers and queen, POLAND: Rauden (now Rudy, Opole Prov.) and GERMANY: Berlin, in hothouses (Roger) (MNHN) [examined]. [Combination in Hypoponera: Taylor, 1967: 12.] (See note 1.) Ponera androgyna Roger, 1859: 246 (footnote). Syntype ergatoid males (not workers), POLAND: Rauden (now Rudy, Opole Prov.) (Roger) (ZMHB) [not seen]. [Junior synonym of punctatissima: Emery & Forel, 1879: 455; Seifert, 2003: 69.] Ponera tarda Charsley, 1877: 162. Syntype workers and queen, GREAT BRITAIN: Oxford, 1877 (R.S. Charsley) (OXUM) [not seen]. [Junior synonym of punctatissima: Dalla Torre, 1893: 41; Seifert, 2003: 69.] Ponera punctatissima r. jugata Forel, 1892: 251. Holotype queen, MADAGASCAR: Prov. d’Imerina (Sikora) (MHNG) [not seen]. [Raised to species: Dalla Torre, 1893: 39. Reverted to subspecies of punctatissima: Emery, 1899: 268. Combination in Hypoponera: Bolton, 1995: 215. Junior synonym of punctatissima: Seifert, 2003: 69.] Ponera ergatandria Forel, 1893: 365. Syntype workers, queens and ergatoid male, SAINT VINCENT I. (Antilles): 41b (H.H. Smith). (MHNG, BMNH) [BMNH ergatoid male examined]. [Junior synonym of punctatissima: Taylor, 1968: 65. Revived status as species and combination in Hypoponera: Kempf, 1972: 122. Reverted to junior synonym of punctatissima: Smith, D.R. 1979: 1343, here confirmed.] (See note 2). Ponera kalakauae Forel, 1899: 115. Syntype worker-queen intercaste (not worker) and queen, HAWAIIAN IS: Kauai I., Lahue, 2000 ft, vii.1896 (R.C.L. Perkins) (intercaste), and Honolulu (Oahu I.), xi.1896 (R.C.L. Perkins) (queen) (BMNH) [examined]. [Junior synonym of gleadowi: Wilson, 1958: 328. Junior synonym of punctatissima: Wilson & Taylor, 1967: 29 (in text).] (See note 3.) Ponera punctatissima subsp. schauinslandi Emery, 1899: 439. Syntype queens, HAWAIIAN IS: Laysan I. (now Laycan I.) (Schauinsland) (MSNG) [not seen] Syn. rev. [Combination in Hypoponera and junior synonym of punctatissima: Wilson & Taylor, 1967: 28. Revived from synonymy and raised to species: Seifert, 2003: 69.] Ponera ergatandria subsp. bondroiti Forel, 1911: 285. Syntype workers, queen and ergatoid male, BELGIUM: Bruxelles, Jardin Bot., xi.1909, 5.xi.1909, 15.xi.1909 (Bondroit) (MHNG) [not seen] Syn. n. [Raised to species: Santschi, 1937b: 364. Combination in Hypoponera: Onoyama, 1989: 2. Previously junior synonym of schauinslandi: Seifert, 2003: 69.] Ponera dulcis var. aemula Santschi, 1911: 351. Lectotype and paralectotype workers (designated by Seifert, 2003: 68), TANZANIA: Kilimandjaro, zone des cultures, Kiboscho, 1400 m, 1904 (Ch. Alluaud) (NHMB) [examined] Syn. n. [Combination in Hypoponera: Bolton, 1995: 213. Previously junior synonym of schauinslandi: Seifert, 2003: 69.] (See note 4.) Ponera ergatandria st. cognata Santschi, 1912: 153. Syntype workers, ANGOLA: Benguela, Cucula (J. Cruchet) (NHMB) [examined]. Syn. n. [Unresolved junior primary homonym of Ponera cognata Emery, 1896: 56 (now in Pachycondyla). Raised to species: Bernard, 1953: 202. Combination in Hypoponera: Bolton, 1995: 213.] (See note 5.) Ponera ragusai var. sordida Santschi, 1914a: 54. Holotype worker, KENYA: région côtière, Shimoni, st. no. 9, xi.1911 (Alluaud& Jeannel) (NHMB) [examined]. Syn. n. [Combination in Hypoponera: Bolton, 1995: 216.] Ponera incisa Santschi, 1914b: 320, fig. 8. Holotype ergatoid male (not worker), NIGERIA: Lagos (F. Silvestri) (not in NHMB, presumably in DEUN) [not seen]. Syn. n. [Combination in Hypoponera: Bolton, 1995: 214.] (See note 6). Ponera sulcatinasis r. durbanensis Forel, 1914: 213. Syntype workers, SOUTH AFRICA: Natal, Durban, 16.i.1914, no. 323 (G. Arnold) (MHNG) [examined]. Syn. n. [Combination in Hypoponera: Bolton, 1995: 214.] Ponera ergatandria r. petri Forel, 1916: 397. Holotype worker-queen intercaste (not worker), DEMOCRATIC REPUBLIC OF CONGO: St Gabriel (Kohl) (MHNG) [examined]. Syn. n. [Raised to species: Santschi, 1938: 78. Combination in Hypoponera: Bolton, 1995: 216.] Ponera brevis Santschi, 1921a: 113, fig. 1a-c. Holotype worker, BENIN (Dahomey on label): Porto Novo (Le Moult) (NHMB) [examined]. Syn. n. [Combination in Hypoponera: Bolton, 1995: 213.] (See note 7.) Ponera punctatissima var. exacta Santschi, 1923: 134. Syntype workers, TUNISIA: Hammamet (Santschi) (NHMB, BMNH) [BMNH syntype examined]. [Junior synonym of punctatissima: Atanassov & Dlussky, 1992: 71; Seifert, 2003: 69.] Ponera mina Wheeler, W.M. 1927b: 131, fig. 2. Syntype workers, queens and ergatoid male, AUSTRALIA: Norfolk I., 1915 (A.M. Lea) (MCZC and possibly SAMA) [not seen]. [Junior synonym of gleadowi: Wilson, 1958: 329. Revived from synonymy and combination in Hypoponera: Taylor & Brown, D.R. 1985: 31. Junior synonym of punctatissima: Wilson & Taylor, 1967: 29 (in text); Taylor, 1987: 30.] Ponera argonautorum Arnol'di, 1932: 66, figs. 6, 7. Holotype queen, RUSSIA: Black Sea Coast, Anapa, 26.viii.1925, Nr. 422 (K.V. Arnol’di) (probably in ZMUM) [not seen]. Syn. n. (provisional). [Combination in Hypoponera: Bolton, 1995b: 213.] (See note 8.) Ponera mumfordi Wheeler, W.M. 1933: 141. Syntype workers, FRENCH POLYNESIA: Marquesas Is, Uapou; Kohepu (Tekohepu) summit, alt. 3000 ft, 3.xi.1931 (LeBronnec) (probably in MCZC) [not seen]. [Junior synonym of gleadowi: Wilson, 1958: 328. Junior synonym of punctatissima: Wilson & Taylor, 1967: 29 (in text).] Ponera mesoepinotalis Weber, 1942: 44, fig. 4. Holotype worker, SUDAN: Imatong Mts, 2.viii.1939, 6400 ft, no. 1395 (N.A. We b e r) (not in MCZC, presumed lost) Syn. n. [Combination in Hypoponera: Bolton, 1995: 215.] (See note 9.) Ponera breviceps Bernard, 1953: 202, fig. 3A. LECTOTYPE dealate queen (by present designation) and two paralectotype dealate queens, GUINEA: Nimba (Lamotte) (MNHN) [examined]. Syn. n. [Combination in Hypoponera: Bolton, 1995: 213.] (See note 10.) Ponera ursoidea Bernard, 1953: 203, fig. 3C. LECTOTYPE worker-queen intercaste (not worker) (by present designation), GUINEA: Crête de Nion, 1300 m. (no collector’s name, probably Lamotte) (MNHN) [examined]. Syn. n. [Combination in Hypoponera: Bolton, 1995: 216.] (See note 11.) Ponera sulcitana Stefani, 1970: 1, figs. 1 – 12. Holotype worker; paratype workers, queens and ergatoid male, ITALY: Sardinia, Grotta dei Fiori presso Carbonia (A. Serra) (IZUC) [not seen] Syn. n. [Combination in Hypoponera: Bolton, 1995: 216.] (See note 12.) NOTES. 1 Seifert (2003) refers to each of the punctatissima type-specimens he examined as a paralectotype. While there seems no doubt that the material originated in Roger’s collection and is type-material, no trace of a lectotype designation can be found. Therefore the two MNHN specimens examined here, and those recorded by Seifert as present in ZMHB and DEIC, revert to syntype status. 2 The BMNH ergatoid male has the data: “41b” and “ Ponera ergatandria Forel, male type, St. Vincent ”. This is certainly a minor ergatoid male of punctatissima (yellow, eyeless and with 12 antennal segments). In the publication, Forel (1893) gives the following localities for ergatandria, all of which are syntypes. 41a: Bowwood Valley, nr Kingston, 800 ft, 15.x.; 41b: Islet fronting Châteaubelais Bay, 31.x.; 41c: Richmond Estate, 31.x.; 41d: Villa Estate, south end of island, 14.x. 3 Wilson (1958: 328-329) recorded the names decipiens, kalakauae, mina and mumfordi as junior synonyms of gleadowi. In the text of Wilson & Taylor (1967: 29) all these names were transferred to the synonymy of punctatissima, but decipiens was later reverted to the synonymy of gleadowi by Onoyama (1989) following his examination of its holotype. These transfers of synonymy were inadvertently omitted from the Bolton (1995) catalogue. 4 The short history of the name aemula amply illustrates the confused nature of the early taxonomy of Afrotropical Hypoponera. When introduced, aemula was described as an infraspecific form of dulcis, now in H. dulcis group. At its next appearance (Santschi, 1914a: 53) aemula had become an infrasubspecific name attached to coarctata subsp. boerorum. The first of these names is now in a separate genus (Ponera) and boerorum is a member of the H. boerorum group. The first critical examination of the syntypes of aemula, by Seifert (2003), placed it firmly in what is now the H. punctatissima group, which is confirmed here. 5 In the original description of cognata, Santschi mentions three workers, with the data recorded above. Of these, two are normally mounted and the head of the third is mounted on a small perspex slide, without any remnants of its body. In NHMB are two more workers, also labelled as “types” but from “ Benguela, Caconda (Cruchet) ”. These are identical to the Cucula specimens, but because they were not mentioned in the original description they have no type-status. 6 The holotype of incisa is not among Santschi’s collection in NHMB. If it still exists it is probably in the Silvestri collection at Portici (DEUN), which is not currently available. The original description gives the holotype as a worker, but it seems most likely that it is really an ergatoid male of punctatissima. Characters noted or figured by Santschi that point towards this identification include: extremely short scapes (from the drawing SI ca 65; measured ergatoid males SI 68 – 72), coupled with worker-like mandibles; large, rectangular, worker-like head; metanotal groove sharp and deep; gastral segments somewhat enlarged. Unfortunately the apical gastral segments are missing from Santschi’s drawing, which may imply that they are absent from the specimen. But the description and figure suggest most strongly that the holotype is an ergatoid male of punctatissima. It is also interesting to note that on the same page Santschi (1914b: 320) recorded a queen of punctatissima, collected from the same locality as incisa. 7 The holotype of brevis is in reasonable condition but is flat-mounted on card and much of the lower and posterior petiole, and the base of the gaster, are obscured by glue. Several characters noted by Santschi in the original description and figures are misleading. For instance, the metanotal groove is not shown in his fig. 1c, but in the text it is stated as present and it is conspicuous on the holotype. The propodeum is not marginate between the declivity and the sides. The lower petiole is obscured not only by glue but by the position of the legs. However, the node in profile appears to taper slightly towards the apex and is not “ as thick at the summit as at the base ”. 8 Ponera argonautorum was provisionally described as new by Arnol’di, based on a single alate queen discovered flying in a garden at Anapa, on the Russian north-east coast of the Black Sea. Its holotype has not been examined but it seems obvious, from the description and figures, that this is a queen of punctatissima. It is provisionally synonymised here to draw attention to our opinion, until the holotype can be examined in detail. 9 Stefan Cover (MCZC) informs us that although the holotype of mesoepinotalis is noted in the MCZC typeregister, the specimen has disappeared from the collection and cannot be found. Weber’s description and figure are sufficient to place the name with considerable confidence as a junior synonym of punctatissima. 10 Bernard’s syntype series of breviceps consisted of four specimens mounted on two pins. The upper specimen on the first pin is a dealate queen of punctatissima, with its gaster detached and mounted separately. The lower specimen on this pin is a headless male of a small species of Leptogenys; its pectinate pretarsal claw is clearly visible on the right foreleg. This pin bears three additional labels, as follows: “ Queen lectotype Ponera breviceps Ber. Designated by R.W. Taylor, 10 Aug. 63 ”; “ Leptogenys sp. male det. R.W. Taylor, 10 Aug. 63 ”; “ Synonym of Ponera gleadowi Forel. Det. R.W. Taylor, 10 Aug. 63 ”. On the second pin both specimens are dealate queens of punctatissima and bear a secondary label: “ Paralectotypes Ponera breviceps Bern. Designated by R.W. Taylor, 10 Aug. 63 ”. Taylor’s designations were never published and the lectotype designation is formalised here. The Leptogenys specimen is excluded from further consideration. 11 Bernard’s syntype series of ursoidea included three workers and a queen from Guinea: Crête de Nion (all on one pin) and a single worker from Guinea: Nimba (Lamotte) on a second pin. On the first pin the top specimen belongs to punctatissima but the two workers and the alate queen mounted below it are all dulcis. The worker on the Nimba pin is a specimen of dulcis that has the right side of its petiole scale damaged by indentation so that it is deformed. Bernard’s description contains a mixture of characters from both species. The first pin bears two secondary labels: “ Lectotype (top specimen with marked pin) and paralectotypes of Ponera ursoidea Ber. Designated by R.W. Taylor ”, and “ A synonym of Ponera gleadowi Forel. Det. R.W. Taylor, 10 Aug. 63 ”. The lectotype designation was never published by Taylor and is now formally established here. The remaining specimens, all dulcis, are hereby excluded from the type-series. One small point remains: the lectotype is not a worker but an intercaste, as indicated by its larger than worker-sized eye and the presence of a mesonotal-mesopleural suture, a character absent from genuine workers of this species. 12 There is nothing in Stefani’s descriptions and figures of the various sexes and castes of sulcitana that separates it from punctatissima. The worker description and drawings (figs. 1, 6, 11) show nothing that would detract from this conclusion, and the worker LPeI ca 50, estimated from his fig. 1, is within the range of punctatissima. Stefani’s “apterous queen ” (his fig. 3) appears to be a normal dealate gyne, not an intercaste, as the mesosoma seems to have a full complement of flight sclerites. His description and drawing of the ergatoid male makes the synonymy confident as the specimen is said to have 12 antennal segments, worker-like mandibles and minute eyes. Some years ago, Prof. Cesare Baroni Urbani (pers. com.) said that he had tried to obtain specimens from Stefani several times, but without success. WORKER. Measurements: HL 0.56 – 0.72, HW 0.46 – 0.60, HS 0.515 – 0.660, SL 0.35 – 0.48, PrW 0.33 – 0.43, WL 0.70 – 0.90, HFL 0.36 – 0.48, PeNL 0.14 – 0.20, PeH 0.30 – 0.39, PeNW 0.22 – 0.29, PeS 0.223 – 0.290 (60 measured). Indices: CI 79 – 87, SI 75 – 84, PeNI 63 – 74, LPeI 43 – 53, DPeI 140 – 165. Eyes small but conspicuous, of 1 – 6 often poorly defined or partially fused ommatidia, located far forward on the side of the head. Impression that extends back along midline of head from the frontal lobes reaches at least the midlength of the vertex and usually further. Apex of scape, when laid straight back from its insertion, usually fails to reach or less commonly just touches the midpoint of the posterior margin in full-face view; SL/HL 0.62 – 0.70. Minute punctulate sculpture of cephalic dorsum extremely fine and superficial. Mesonotal-mesopleural suture absent. Metanotal groove deeply incised across dorsum of mesosoma; mesonotum with a well-defined posterior margin. Propodeal declivity rounds into the sides, without sharp marginations or carinae. In profile the anterior margination of the mesopleuron rounded to very bluntly angular behind base of anterior coxa; without a prominent sharp angle or tooth. Mesopleuron smooth and shining, at most with a few scattered punctures; usually with a small reticulate patch in the extreme posteroventral corner above the mesocoxa. Petiole in profile with the anterior and posterior faces of the node weakly convergent dorsally; node longer just above the anterior tubercle than at the dorsum. Sternite of petiole in profile a rounded lobe, without angles anteriorly or posteriorly. Maximum width of first gastral tergite in dorsal view less than the width of the second tergite at its midlength. Base of cinctus of second gastral tergite smooth and shining in dorsal view, without cross-ribs. Posttergite of second gastral segment, from posterior margin of cinctus to apex, much broader than long. Disc of second gastral tergite minutely punctulate, appearing weakly microreticulate in places. Full adult colour varies from dull brownish yellow to very dark brown or almost black. In general, workers at the bottom end of the size range given above tend to be lighter and the colour becomes darker with increasing size, but this is by no means an entirely consistent rule. Apart from variations in size and colour, all the worker material of this widely distributed species is morphologically uniform and answers the description above. H. punctatissima is without doubt the world’s most accomplished ponerine tramp-species. Its range incorporates all tropical and subtropical zoogeographical regions, including most oceanic islands, and it also penetrates well into the temperate zones of both hemispheres where it is frequently synanthropic; for a brief synopsis of world distribution see Delabie & Blard (2002); European distribution is summarised in Seifert (2003). As well as the usual workers and alate queens, it also produces worker-queen intercastes (ergatoids) and dimorphic large and small ergatoid males, but never the usual alate male form. Because of its tramping ability, oddly polymorphic sex/caste system and the considerable confusion about the taxonomy within the genus, punctatissima has amassed an inordinately large synonymy. A glance at the synonymic synopsis above will illustrate how often intercastes and ergatoid males have been misinterpreted as workers of separate taxa by authors dating back to Roger (1859) and how often supposed infraspecific forms were incorrectly associated with other species. In a recent analysis, Seifert (2003) reached the conclusion that punctatissima actually consisted of two species, punctatissima and schauinslandi, that were not distinguishable by standard morphology but could be separated by micro-morphometrics subjected to discriminant analysis. He concluded that punctatissima had the junior synonyms androgyna, exacta, jugata and tarda (of which only exacta had not been previously proposed as a synonym), and that schauinslandi was the senior synonym of aemula and bondroiti. He stated that he had examined type-material of all the available names that could probably represent punctatissima or schauinslandi, to ensure that no possible senior name remained unconsidered. Unfortunately, a quick glance at the synopsis above shows that many names associated with punctatissima remained unexamined for the Afrotropical region, that several synonyms proposed by Wilson & Taylor (1967) had been omitted, and that one name from the latter publication has the same date as schauinslandi. It is not surprising that Seifert failed to detect the Afrotropical relations as their identities could not possibly be assessed based solely on their often inadequate and frequently misleading original descriptions. Taken together, the Afrotropical type-material and the mass of other material examined here did not fall neatly into the two presumed species, or into any one of the two, which raises the possibility that either there are more than two extremely cryptic and virtually inseparable species currently confused here, or that there is only one somewhat variable genuine species present. The possibility remains strong that the discriminant functions applied by Seifert do not isolate discrete species, but rather indicate allopatric populations of the same species, or even different ecomorphs of a single species. The problem could not be completely resolved here, though the ecomorph possibility is favoured, so it was decided that the most parsimonious approach would be to regard all as a single species, under the earliest available name, until a complete analysis of all material, world-wide, becomes feasible. Whether this can be accomplished by morphometrics or must await a DNA analysis of various populations remains to be seen, but a dedicated study is certainly essential. Therefore, at present, H. punctatissima workers are regarded as any specimen that possesses the characters of the punctatissima group (see introduction) and fits the description above. Its worker-queen intercastes are very worker-like but have larger eyes (usually 10 to about 20 ommatidia) and a developed mesonotal-mesopleural suture that is absent in the worker. Ergatoid males fall into major (larger, brown and with small eyes present, Figs 100 – 102) and minor (smaller, yellow and eyeless) categories (Yamauchi, et al. 1996) and are again very worker-like, especially in head structure, but have shorter scapes (SI 68 – 72), only 12-segmented antennae (as do workers, intercastes and queens), and of course fully developed but retractile male genitalia. Alate males have never been seen and it is probably now safe to say are never produced. True queens are alate, with much larger eyes (obviously>50 ommatidia) that have short setae projecting between the ommatidia, and have a full complement of flight sclerites on the mesosoma. A closely related species, ragusai, has been confused (under the name of its junior synonym, gleadowi) with punctatissima in the past (e.g. Wilson, 1958). H. ragusai has much the same sex/caste system as punctatissima, including monomorphic ergatoid males which have 13-segmented antennae, as is most frequently the case in male ants. This species is very uncommon in the Afrotropical region and indeed seems rare everywhere. In general the worker of ragusai is small and yellowish. Its petiole in profile is lower and longer than in punctatissima and in d